Pengantar

Human Nutrition

Edisi Kedua

Diedit atas nama The Nutrition Society oleh

Michael J Gibney
Susan A Lanham-New
Aedin Cassidy
Hester H Vorster

A John Wiley & Sons, Ltd., Publikasi

Pengantar
 Nutrisi
Manusia
Seri Buku Teks Masyarakat Nutrisi Menyusui
Hasil yang merugikan pada kehamilan

Pengantar Nutrisi Manusia

Pengantar Nutrisi Manusia: perspektif global tentang
makanan dan nutrisi Nutrisi dan Metabolisme
Komposisi tubuh Konsep inti nutrisi Nutrisi
Metabolisme energi molekuler
Nutrisi dan metabolisme protein dan asam amino Pengaturan asupan makanan
Pencernaan dan metabolisme karbohidrat Integrasi metabolisme 1:energi
Nutrisi dan metabolisme lipid Integrasimetabolisme 2: Karbohidrat dan lipid
Standar referensi diet Integrasi metabolisme 3: Protein dan asam amino
Vitamin Fitokimia
Mineral dan elemen jejak Kehamilan dan menyusui
Mengukur asupan Pertumbuhan dan penuaan
makanan Komposisi Saluran gastrointestinal
makanan Makanan dan gizi: masalah kebijakan Sistem kardiovaskular Sistem
dan peraturan Metodologi penelitian gizi rangka Sistem
Keamanan pangan: masalah kesehatan masyarakat imun dan inflamasi Sistem
yang semakin penting Pangan dan penyakit terkait gizi: sensorik
tantangan global Aktivitas fisik
Overnutrisi
Kurang
Otak

Gizi Kesehatan Masyarakat

Gambaran umum gizi kesehatan masyarakat GiziNutrisi Klinis
Epidemiologi gizi Prinsip umum cnutrisi linical
Pilihan pangan metabolik dan penilaian gizi
Penilaian status gizi pada individu dan penduduk kelebihan gizi
tingkat ion gizi
Penilaian aktivitas fisik Gangguanmetabolik
Gizi Gangguan makan
berlebihan Efek samping untuk makanan
Gangguan makan Gangguan makan, diet dan mode dukungan nutrisi
makanan Etis dan masalah hukum
Strategi PHN untuk nutrisi: intervensi pada tingkat Saluran cerna
individu Hati
Strategi PHN untuk nutrisi: intervensi pada Pankreas
tingkat ekologi Panduan ginjal
makanan dan nutrisi Darahdan sumsum tulang
Pemrograman janin The paru-paru
Penyakit kardiovaskular kekebalan dan infl sistem inflamasi
Kanker jantung dan pembuluh darah
Osteoporosis Penyakitkerangka
Diabetes Traumatic
Vitamin A defi defisiensi Infectious penyakit Penyakit
yodium Defisiensi zat ganas
besi Gizi anak Gizi anak
Kesehatan ibu dan anak
Cystic fibrosis Air dan elektrolit
Kasus klinis

Pengantar
Human Nutrition

Edisi Kedua

Diedit atas nama The Nutrition Society oleh

Michael J Gibney
Susan A Lanham-New
Aedin Cassidy
Hester H Vorster

A John Wiley & Sons, Ltd., Publikasi

Edisi pertama diterbitkan 2009
Edisi pertama diterbitkan 2002
© 2009, 2002 oleh The Nutrition Society

Blackwell Publishing diakuisisi oleh John Wiley & Sons pada Februari 2007. Program penerbitan Blackwell
telah digabungkan dengan bisnis Ilmiah, Teknis, dan Medis global Wiley untuk membentuk Wiley-Blackwell.

Kantor terdaftar
John Wiley & Sons Ltd, The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, United Kingdom Kantor

editorial
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2121 State Avenue, Ames, Iowa 50014-8300, Amerika Serikat

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mengajukan izin untuk menggunakan kembali materi hak cipta dalam buku ini, silakan lihat situs web kami di
www.wiley.com/wiley-blackwell.

Hak penulis untuk diidentifikasi sebagai penulis karya ini telah ditegaskan sesuai dengan Undang-Undang
Hak Cipta, Desain dan Paten 1988.

Semua hak dilindungi undang-undang. Tidak ada bagian dari publikasi ini yang boleh direproduksi,
disimpan dalam sistem pengambilan, atau ditransmisikan, dalam bentuk apa pun atau dengan cara apa
pun, elektronik, mekanis, fotokopi, rekaman, atau lainnya, kecuali sebagaimana diizinkan oleh Undang-
Undang Hak Cipta, Desain dan Paten Inggris 1988, tanpa izin sebelumnya dari penerbit.

Wiley juga menerbitkan buku-bukunya dalam berbagai format elektronik. Beberapa konten yang muncul di
cetakan mungkin tidak tersedia di buku elektronik.

Sebutan yang digunakan oleh perusahaan untuk membedakan produk mereka sering kali diklaim sebagai
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profesional atau bantuan ahli lainnya diperlukan, layanan profesional yang kompeten harus dicari.

Library of Congress Katalogisasi-dalam-Data Publikasi

Pengantar nutrisi manusia / diedit atas nama Nutrition Society oleh Michael J. Gibney. . . [dkk.]. - edisi ke-2.
p. ; cm. - (Seri buku teks nutrisi manusia)
Termasuk referensi bibliografi dan indeks.
ISBN 978-1-4051-6807-6 (pbk.: Alk. Paper) 1. Nutrisi. I. Gibney, Michael J. II. Nutrition Society
(Inggris Raya) III. Seri.
[DNLM: 1. Fisiologi Gizi 2. Pangan. QU 145 I623 2009]
QP141.I665 2009
612.3 – dc22
2008035123

Catatan katalog untuk buku ini tersedia dari British Library.

Ditetapkan dalam 10 di 12 pt Minion oleh SNP Best-set typesetter Ltd., Hong Kong
Dicetak di Singapura oleh Fabulous Printers Pte Ltd

1 2009
 Isi
Kontributor vii Seri Kata Pengantar ix Kata Pengantar xi Ucapan Terima Kasih xii
1. Pengantar Nutrisi Manusia: Perspektif Global tentang Makanan dan Nutrisi 1 HH Vorster
2. Komposisi Tubuh 12 P Deurenberg
3. Metabolisme Energi 31 A Astrup dan A Tremblay
4. Nutrisi dan Metabolisme Protein dan Asam Amino 49 NK Fukagawa dan YM Yu
5. Pencernaan dan Metabolisme Karbohidrat 74 J Mathers dan TMS Wolever
6. Nutrisi dan Metabolisme Lipid 86 BA Griffi n dan SC Cunnane
7. Standar Referensi Diet 122 KM Lebih Muda
8. Vitamin 132 DA Bender
9. Mineral dan Elemen Jejak 188 JJ Strain dan KD Cashman
10. Mengukur Asupan Makanan 238 UE MacIntyre
11. Komposisi Pangan 276 HC Schönfeldt dan JM Holden
12. Pangan dan Gizi: Masalah Kebijakan dan Peraturan 293 MJ Gibney dan A McKevitt
13. Metodologi Penelitian Gizi 305 JA Martínez dan MA Martínez-González
14. Makanan Keselamatan: Masalah Kesehatan Masyarakat yang Meningkat Pentingnya 324
A Reilly, C Tlustos, J O'Connor, dan L O'Connor
15. Penyakit Terkait Makanan dan Gizi: The Global Challenge 350 HH Vorster dan MJ Gibney
Index 361
Dr Stephen Cunnane
Kontributor Departemen Kedokteran, Fisiologi dan
Biofisika dan Pusat Penelitian
Penuaan,Université de Sherbrooke
Kanada
Profesor Arne Astrup Profesor
Kepala, Departemen Nutrisi Manusia, Associate dari,
Fakultas Ilmu Kehidupan, Departemen Nutrisi Manusia,
Universitas Kopenhagen, Universitas Wageningen,
Kopenhagen, Denmark Belanda
Profesor Tamu,
Dr David A Bender Universitas Tor Vergata , Rome,
Sub-Dekan (Pendidikan), Italy Nutrition Consultant,
Sekolah Kedokteran Universitas College Singapore
London, London, Inggris
Professor Naomi K Fukagawa
Profesor Kevin D Cashman Department of Medicine,
Departemen Ilmu Pangan dan Gizi, University University of Vermont,
College Cork, Burlington, Vermont, USA
Irlandia
Pro fessor Michael J Gibney Dr Lisa O'Connor
Departemen Kedokteran Klinis, Badan Keamanan Pangan Irlandia,
Trinity College, Dublin, Dublin, Irlandia
Irlandia
Alan Reilly
Otoritas Keamanan Pangan Irlandia,
Dublin, Irlandia

Profesor Hettie C Schönfeldt

Sekolah Pertanian dan Ilmu Pangan
Dr Bruce A Griffi n Universitas Pretoria,
Reader di Metabolisme Afrika Selatan
Nutrisi, Divisi Ilmu Gizi,
Fakultas Ilmu Kesehatan dan
Profesor JJ (Sean) Strain
Kedokteran, Universitas Surrey,
Profesor Nutrisi Manusia,
Guildford, UK
Irlandia Utara Pusat Pangan dan
Kesehatan, Universitas Ulster,
Joanne M Holden Coleraine, Irlandia Utara
Nutrient Data Laboratory,
Beltsville, Christina Tlustos
Maryland, AS Food Safety Authority of Ireland,
Dublin, Irlandia
Una E MacIntyre
Institute for Human Nutrition, Angelo Tremblay
University of Limpopo, Preventive and Social Medicine,
Medunsa, Laval University,
Afrika Selatan Ste-Foy, Québec,
Kanada
Dr Aideen McKevitt
School of Biomedical Sciences,
University of Ulster,
Northern Ireland Profesor Hester H Vorster
Direktur Centre of Excellence for Nutrition
Professor J Alfredo Martínez Faculty of Health Sciences,
Intitute of Nutrition and Food Sciences, North-West University
University of Navarra, Potchefstroom, Afrika Selatan
Spanyol
Dr Thomas MS Wolever
ProfesorMiguel A Martínez-González Departemen Ilmu Gizi,
Departemen Pengobatan Pencegahan dan Fakultas Kedokteran,
Kesehatan Masyarakat, Universitas Toronto,
Universitas Navarra, Kanada
Spanyol
Dr Kate MMuda
Profesor John Mathers DosenNutrisi Manusia,
Pusat Penelitian Nutrisi Manusia, Sekolah Ilmu Biologi,
Institut untuk Penuaan dan Kesehatan Institut Teknologi Dublin,
Universitas Newcastle, Inggris Irlandia

Dr Judith O'Connor Dr Yong-Ming Yu

Otoritas Keamanan Pangan Irlandia, Departemen Bedah,
Dublin , Irlandia Rumah Sakit Umum Massachusetts dan Rumah
viii Kontributor
 Sakit Shriners Burns,
Sekolah Kedokteran Harvard,
Boston, Massachusetts, AS.
S eries Kata
Pengantar Dekade
-dekade awal abad ke-20 adalah periode
penelitian intensif tentang unsur makanan yang
penting untuk pertumbuhan dan perkembangan
normal, dan menyaksikan penemuan sebagian
besar vitamin, mineral, asam amino, dan asam
lemak esensial. Pada tahun 1941, sekelompok
ahli fisiologi, ahli biokimia, dan ilmuwan medis
terkemuka mengakui bahwa disiplin ilmu gizi
yang muncul membutuhkan masyarakat
terpelajarnya sendiri dan Masyarakat Nutrisi
didirikan. Misi kami adalah, dan tetap, "untuk
memajukan studi ilmiah nutrisi dan aplikasinya
untuk pemeliharaan kesehatan manusia dan
hewan". Nutrition Society adalah komunitas
terpelajar nutrisi terbesar di Eropa dan kami
memiliki lebih dari 2000 anggota di seluruh dunia.
Anda dapat mengetahui lebih lanjut tentang
Society dan cara menjadi anggota dengan
mengunjungi situs web kami di
www.nutsoc.org.uk
Revolusi berkelanjutan dalam biologi yang
diprakarsai oleh pemetaan genom skala besar
dan difasilitasi oleh pengembangan yang andal,
sederhana untuk -menggunakan alat biologi
molekuler menjadikan ini waktu yang sangat
menyenangkan untuk bekerja di bidang nutrisi.
Sekarang kita memiliki kesempatan untuk
memperoleh pemahaman yang lebih baik tentang
bagaimana
gen tertentu berinteraksi dengan asupan nutrisi
dan faktor gaya hidup lainnya untuk
memengaruhi ekspresi gen dalam sel dan
jaringan individu dan, pada akhirnya,
memengaruhi kesehatan kita. Pengetahuan
tentang polimorfisme dalam gen kunci yang
dibawa oleh pasien akan memungkinkan resep
perawatan diet yang lebih efektif dan aman. Di
tingkat populasi, epidemiologi molekuler
membuka pendekatan yang jauh lebih tajam
untuk memahami peran pola makan tertentu
dalam penyebab penyakit
. Kegembiraan ini tercermin dalam beberapa
pertemuan ilmiah yang diselenggarakan oleh
Nutrition Society, yang sering kali bekerja sama
dengan komunitas sister terpelajar di Eropa,
setiap tahun. Kami memberikan hibah perjalanan
dan bantuan lainnya untuk mendorong siswa dan
peneliti muda untuk menghadiri dan berpartisipasi
dalam pertemuan ini.
Sepanjang sejarahnya, tujuan utama
Perhimpunan adalah untuk mendorong penelitian
nutrisi dan menyebarkan hasil penelitian tersebut.
Jurnal pertama kami, The Proceedings of the
Nutrition Society,
mencatat, seperti yang masih terjadi, presentasi
ilmiah yang dibuat untuk Society. Tak lama
kemudian, The British Journal of Nutrition
didirikan untuk menyediakan
media publikasi penelitian utama tentang semua
aspek gizi manusia dan hewan oleh para ilmuwan
dari seluruh dunia. Menyadari kebutuhan siswa
dan guru mereka untuk tinjauan otoritatif tentang
isu-isu topikal dalam nutrisi, Society mulai
menerbitkan Tinjauan Penelitian Nutrisi pada
tahun 1988. Pada tahun 1997, kami meluncurkan
Nutrisi Kesehatan Masyarakat, jurnal
internasional pertama yang didedikasikan untuk
bidang yang penting dan berkembang ini . Semua
jurnal ini tersedia dalam bentuk elektronik, serta
dalam bentuk kertas konvensional dan kami
sedang menjajaki peluang baru untuk
memanfaatkan web agar hasil penelitian nutrisi
lebih cepat dan mudah diakses. Untuk melindungi
publik dan meningkatkan prospek karir ahli gizi,
Masyarakat Nutrisi berkomitmen untuk
memastikan bahwa mereka yang berpraktik
sebagai ahli gizi mendapatkan pelatihan dan
kualifikasi yang tepat. Hal ini diakui dengan
menempatkan nama-nama individu yang
memenuhi kualifikasi yang sesuai pada daftar
profesional kami dan dengan penghargaan
kualifikasi Ahli Gizi Kesehatan Masyarakat
Terdaftar (RPHNutr) dan Ahli Gizi Terdaftar
(RNutr). Lulusan dengan gelar yang sesuai tetapi
belum memiliki pengalaman pascasarjana yang
memadai dapat bergabung dengan register
Associate Ahli Gizi kami. Kami melakukan
akreditasi program gelar universitas di bidang gizi
kesehatan masyarakat dan sedang
mengembangkan proses akreditasi untuk
program gelar gizi lainnya.
Seperti halnya dalam penelitian, memiliki alat
terbaik adalah keuntungan besar dalam proses
 belajar mengajar. Inilah alasan di balik inisiatif The Nutrition Society Textbook
untuk meluncurkan seri buku teks nutrisi manusia
yang dirancang untuk digunakan di seluruh dunia.
Hal ini dicapai dengan berhasil meluncurkan seri Outgoing Editor-in-Chief
pertama dalam berbagai bahasa termasuk
Spanyol, Portugis dan Yunani. Society sangat University College Dublin, Irlandia
berhutang budi kepada Profesor Mike Gibney dan
tim editornya atas kerja tak kenal lelah mereka
dalam 10 tahun terakhir untuk membawa edisi
pertama dari seriteks ini
bukumembuahkan hasil yang sukses di seluruh
dunia. Kami menantikan edisi baru ini di bawah
pengawasan Dr Susan Lanham-New dalam
ukuran yang sama. Baca, pelajari, dan nikmati.
Profesor Ian McDonald
Presiden Masyarakat Nutrisi
Kata Pengantar
Seri Buku Teks Masyarakat Nutrisi dimulai
sepuluh tahun yang lalu sebagai proyek ambisius
untuk menyediakan siswa sarjana dan
pascasarjana dengan rangkaian buku teks yang
komprehensif untuk memenuhi kebutuhan
mereka dalam hal bahan referensi untuk studi
mereka. Secara keseluruhan, proyek ini telah
berhasil dan Seri Buku Teks Masyarakat Nutrisi
telah diadaptasi oleh semua unit nutrisi akademik
terbaik di seluruh dunia. Serial tersebut telah
diterjemahkan ke dalam bahasa Spanyol dan
Portugis.
edisi kedua Pengantar Nutrisi Manusia ini
merupakan pembaruan dari fondasi paling dasar
untuk studi nutrisi manusia. Meskipun sedikit
yang berubah, semua penulis telah membuat
pembaruan apa pun yang diperlukan dan kami
telah mengatur ulang beberapa bab. Studi
tentang nutrisi manusia di universitas di seluruh
dunia berkembang pesat karena peran diet dalam
kesehatan menjadi lebih jelas. Memang,
sekuensing genom manusia telah menyoroti
kisaran gen yang lebih sempit yang
mengendalikan biologi manusia, menekankan
peran penting lingkungan termasuk pola makan
dalam kesehatan manusia. Selain itu, kami
sekarang menyadari peran penting diet dalam
berinteraksi dengan genom kita baik di dalam
rahim maupun dalam periode langsung
perkembangan pascapersalinan.
Series
Editor
Profesor Michael J Gibney
Asisten Editor
Julie Dowsett
University College Dublin, Irlandia
Studi tentang nutrisi manusia membutuhkan
dasar yang kuat dalam fisiologi dan biokimia
metabo lisme manusia dan itu adalah dasar dari
buku teks Nutrition and Metabolism. Buku teks ini
dirancang untuk melayani dua kebutuhan.
Pertama, banyak yang akan menggunakan buku
ini sebagai pengantar nutrisi manusia dan tidak
melangkah lebih jauh. Pelaku di bidang farmasi,
ilmu pangan, pertanian dan sejenisnya dapat
mengambil modul pengantar nutrisi manusia dan
meninggalkan subjek di sana tetapi mendapat
informasi yang baik di daerah tersebut. Mereka
yang akan mempelajari gizi manusia akan
menemukan dalam buku teks ini pengantar
tentang banyak bidang diet dan kesehatan yang
akan mereka pelajari secara lebih mendalam
dengan menggunakan buku teks yang tersisa
dalam seri Nutrition Society. Selain biologi dasar,
siswa akan diperkenalkan dengan konsep
kebijakan pangan dan tantangan ganda pada
pasokan pangan global, baik kelebihan maupun
kekurangan gizi.
Saat saya menulis, saya menyerahkan
kepemimpinan Seri Buku Teks Masyarakat Nutrisi
kepada Dr Susan Lanham-New di Universitas
Surrey yang telah setuju untuk mengambil tugas
penting ini bagi Masyarakat. Saya ingin berterima
kasih kepada semua orang yang telah bekerja
sama dengan saya dalam proyek ini dan
mendoakan yang terbaik bagi Sue dan tim
barunya.
Michael J Gibney
Pemimpin Redaksi Masuk
 Susan A Lanham-New semua yang telah melayani di Komite Ilmiah
University of Surrey, Internasional dan Editor Buku Teks, tanpa
kepada siapa tugas ini tidak dapat diatasi dan
Asisten Editor Inggris kepada semua penulis yang memberikan
Jennifer Norton
The Nutrition Society, UK

Ucapan Terima
waktu untuk membuat edisi ini mungkin. Terima
Kasih kasih yang sangat khusus harus diberikan
kepada Mike Gibney dan Julie Dowsett, atas
upaya dan dedikasinya dalam melihat buku teks
edisi kedua ini hingga penerbitan.
Dengan penghargaan yang bersyukur kepada

1
Pengantar Nutrisi Manusia:
Perspektif Global tentang Pangan
dan Gizi
Hester H Vorster

Pesan-pesan utama

peserta magang dalam ilmu gizi dan praktik,

• Nutrisi manusia adalah domain ilmiah multifaset yang
kompleks yang menunjukkan bagaimana zat dalam siswa akan belajar bagaimana mengumpulkan,
makanan memberikan nutrisi penting untuk pemeliharaan
kehidupan. menyusun, dan mengklasifikasikan pengetahuan
dengan membaca, eksperimen, observasi, dan
• Untuk memahami, mengkaji, meneliti, dan mempraktekkan
penalaran. Jalan untuk perjalanan ini telah
gizi diperlukan pendekatan terpadu yang holistik dari tingkat
dipetakan ribuan tahun yang lalu. Pengetahuan
molekuler hingga kemasyarakatan. • Nutrisi yang optimal
dan seimbang adalah penentu utama kesehatan. Ini dapat
bahwa nutrisi - apa yang kita pilih untuk makan
digunakan untuk meningkatkan kesehatan dan
dan minum - mempengaruhi kesehatan,
kesejahteraan, untuk mencegah kesehatan yang buruk dan
untuk mengobati penyakit. kesejahteraan, dan kualitas hidup kita sudah
setua sejarah manusia. Selama jutaan tahun
• Studi tentang struktur, sifat kimia dan fisik, serta efek
pencarian makanan telah membantu membentuk
fisiologis dan biokimia dari lebih dari 50 nutrisi yang
ditemukan dalam makanan mendukung pemahaman perkembangan manusia, organisasi masyarakat
tentang nutrisi.
dan sejarah itu sendiri. Ini telah menimbulkan
perang, pertumbuhan penduduk, perluasan
perkotaan, teori ekonomi dan politik, agama,
1.1 Orientasi pada nutrisi manusia sains, kedokteran
, dan perkembangan teknologi.
Tujuan utama dari seri empat buku teks tentang Barulah di paruh kedua abad kedelapan belas
nutrisi ini adalah untuk membimbing siswa nutrisi nutrisi mulai mengalami pengalaman pertama
melalui perjalanan yang mengasyikkan dalam
penemuan nutrisi sebagai sebuah ilmu. Sebagai © 2009 HH Vorster.
 • Ratusan juta orang yang mengalami rawan pangan dantubuh secara keseluruhan memperoleh dan
gizi secara global, kekurangan gizi dan kelebihan gizi,
menggunakan substansi yang diperlukan yang
serta perilaku gizi yang tidak tepat merupakan tantangan
yang dihadapi ahli gizi saat ini. diperoleh dari makanan (nutrisi) untuk menjaga
integritas struktural dan fungsional. Untuk
• Praktik gizi memiliki dasar penelitian dan pengetahuan
yang kuat dan berkembang dengan baik. Namun memahami bagaimana manusia memperoleh dan
memanfaatkan makanan dan nutrisi dari tingkat
demikian, terdapat banyak bidang di mana lebih banyak
informasi diperlukan untuk menyelesaikan masalah gizi
molekuler ke tingkat masyarakat, dan faktor-
global, regional, komunal dan individu.
• Pengembangan norma, standar, dan nilai etika dalam
faktor yang menentukan dan mempengaruhi
penelitian dan praktik gizi diperlukan. proses ini, studi dan praktik nutrisi manusia
melibatkan spektrum ilmu pengetahuan dasar
dan terapan lainnya. c disiplin ilmu. Ini termasuk
biologi molekuler, genetika, biokimia, kimia, fisika,
ilmu pangan, mikrobiologi, fisiologi, patologi,
imunologi, psikologi, sosiologi, ilmu politik,
kebangkitan kembali dengan pengamatan oleh
antropologi, pertanian, farmakologi, komunikasi,
para ilmuwan bahwa asupan makanan tertentu,
dan ekonomi. Departemen Gizi, oleh karena itu,
yang kemudian disebut nutrisi, dan akhirnya zat
sering ditemukan di Fakultas Kedokteran
lain yang belum diklasifikasikan sebagai nutrisi,
(Kesehatan) atau Ilmu Sosial, atau Farmasi, atau
mempengaruhi fungsi tubuh, melindungi dari
Pertanian di lembaga pelatihan tersier. Sifat
penyakit, memulihkan kesehatan, dan
multidisiplin ilmu gizi, baik dalam bidang ilmu
menentukan respons orang terhadap perubahan
alam (biologi) maupun sosial, menuntut
lingkungan. Selama periode ini, nutrisi dipelajari
mahasiswa ilmu gizi harus memiliki pemahaman
dari model atau paradigma medis dengan
dasar tentang berbagai cabang ilmu dan harus
mendefinisikan struktur kimia dan karakteristik
mampu mengintegrasikan konsep yang berbeda.
nutrisi yang ditemukan dalam makanan, fungsi
dari disiplin ilmu yang berbeda ini. Ini menyiratkan
fisiologisnya, reaksi biokimia dan kebutuhan
bahwa siswa harus memilih mata pelajaran yang
manusia untuk mencegah, pertama, penyakit
menyertainya (mata pelajaran pilihan) dengan
defisiensi dan, kemudian, juga penyakit tidak
hati-hati dan bahwa mereka harus membaca
menular kronis.
secara luas di berbagai bidang ini.
Sejak akhir 1980-an nutrisi telah mengalami
kebangkitan kedua dengan tumbuhnya persepsi
bahwa pengetahuan yang diperoleh tidak 1.3 Kerangka konseptional
membekali umat manusia untuk memecahkan
untuk studi nutrisi
masalah kerawanan pangan dan malnutrisi global
. Penekanannya bergeser dari paradigma medis Dalam perjalanan penemuan ke dalam ilmu
atau jalur ologis ke yang lebih psikososial, nutrisi, seringkali perlu untuk menempatkan
perilaku di mana gizi didefinisikan sebagai pengetahuan baru, atau aplikasi baru dari
manusia dasar pengetahuan lama, ke dalam perspektif
2 Pengenalan Hak Gizi Manusia gambaran holistik. Untuk ini, kerangkakonseptual
kerjadari sifat multidisiplin ilmu dan praktik nutrisi
mungkin bernilai.seperti itu
, tidak hanya penting untuk perkembangan
manusia tetapi juga sebagai hasil dari
pembangunan.
Dalam teks pengantar pertama ini, fokusnya Kerangka konseptual, yang menggambarkan
adalah pada prinsip-prinsip dan esensi nutrisi interaksi kompleks antara faktor internal atau
manusia, dengan tujuan utama membantu konstitusional dan faktor lingkungan eksternal
mahasiswa nutrisi untuk mengembangkan yang menentukangizi
pemahaman yang holistik dan terintegrasi dari statusdan kesehatan, disajikan pada Gambar 1.1.
domain ilmiah yang kompleks dan multifaset ini. Pada tingkat genetik sekarang diterima bahwa
nutrisi menentukan ekspresi fenotipik dari gen
otipe individu dengan mempengaruhi proses
1.2 Pendekatan Terpadu transkripsi, translasi, atau reaksi pasca-translasi.
Dengan kata lain, nutrisi dapat secara langsung
Nutrisi manusia menggambarkan proses di mana
mempengaruhi ekspresi genetik (DNA),
organel seluler, sel, jaringan, organ, sistem, dan
menentukan jenis RNA yang terbentuk Gambar 1.1 Kerangka konseptual untuk pemahaman yang
(transkripsi) dan juga protein yang disintesis holistik dan terintegrasi tentang nutrisi manusia.
Perspektif Global tentang Pangan dan Gizi 3
(terjemahan). Misalnya, glukosa, suatu
makronutrien karbohidrat, meningkatkan Tingkat fungsi manusia
transkripsi untuk sintesis glukokinase, zat besi (faktor)
mikronutrien meningkatkan translasi untuk
Konstitusional
sintesis ferritin, sedangkan vitamin K
meningkatkan karboksilasi pasca-translasi dari • Inti sel
residu asam glu tamic untuk sintesis protrombin. DNA RNA
(tingkat Genetik)
Nutrisi, oleh karena itu, memengaruhi sintesis
protein struktural dan fungsional, dengan
memengaruhi ekspresi gen di dalam sel.
Nutrisi juga bertindak sebagai substrat dan
• Sel: metabolisme
kofaktor dalam semua reaksi metabolisme dalam • Lingkungan internal
sel yang diperlukan untuk pertumbuhan dan • Sirkulasi
pemeliharaan struktur dan fungsi. Sel mengambil • Semua sistem organ, juga
nutrisi (melalui mekanisme kompleks • Sistem saraf pusat
melintasi membran sel) dari lingkungan
terdekatnya, yang juga dikenal sebagai
lingkungan internal tubuh. Komposisi lingkungan Status gizi
ini diatur secara cermat untuk memastikan fungsi Kesehatan / sakit-sakitan
dan kelangsungan hidup sel yang optimal, suatu
proses yang dikenal sebagai homeostasis, yang
melahirkan pendekatan sistem dalam studi Lingkungan eksternal
nutrisi. • Ketahanan / ketidakamanan
Nutrisi dan oksigen disediakan untuk pangan
lingkungan internal oleh darah yang bersirkulasi, • Karakteristik rumah
yang juga menghilangkan produk akhir metabolik tangga;perawatan •
keadaanSosial
dan zat berbahaya dari lingkungan ini untuk • keadaan ekonomi
dikeluarkan melalui kulit, ginjal, dan usus besar. • Perumahan, sanitasi,
Fungsi bersama dari berbagai organ dan politik • Pertanian,
pelayanan kesehatan
sistem tubuh memastikan bahwa nutrisi dan (sumber; ideologi)
oksigen diekstraksi atau diambil dari lingkungan
eksternal dan ditransfer ke darah untuk diangkut
dan dikirim ke lingkungan internal dan sel. Sistem
pencernaan, misalnya, bertanggung jawab atas
konsumsi makanan dan minuman, pemecahan sistem dalam menanggapi komposisi kimia dan
(pencernaan dan fermentasi) ini untuk ekstraksi fisik darah dan lingkungan internal, dan untuk
nutrisi, dan penyerapan nutrisi ke dalam sirkulasi, kebutuhan seluler.
sedangkan sistem pernapasan mengekstraksi Keadaan kesehatan atau penyakit dari
oksigen dari udara. . Fungsi-fungsi ini berbagai organ dan sistem akan menentukan
dikoordinasikan dan diatur oleh endokrin dan kebutuhan nutrisi tubuh secara keseluruhan.
saraf pusat. Sistem saraf pusat juga merupakan tempat
atau "markas" dari fungsi mental yang lebih tinggi
yang terkait dengan perilaku sadar atau kognitif,
spiritual, religius, dan budaya, yang akan
menentukan, dalam menanggapi lingkungan
internal dan eksternal, apa dan bagaimana
banyak yang akan dimakan. Apa dan berapa
banyak yang dimakan selanjutnya akan
bergantung pada apa yang tersedia, yang
dipengaruhi oleh sejumlah faktor yang
menentukan ketahanan pangan. Semua faktor ini,
pada tingkat individu, rumah tangga, komunitas,
nasional, atau internasional, membentuk dapat dimodifikasi, dan kuat dalam
lingkungan eksternal. mempromosikan kesehatan, mencegah dan
Selama kebangkitan pertama nutrisi, mengobati penyakit, dan meningkatkan kualitas.
penekanan ditempatkan pada studi nutrisi dan hidup.
fungsinya
. Sebuah model medis, ilmu alam atau biologi
mendukung studi tentang hubungan antara nutrisi 1.5 Nutrisi: dasar-dasar
dan kesehatan atau kesehatan yang buruk.
Selama kebangkitan kedua, aspek-aspek ini tidak Orang makan makanan, bukan nutrisi; Namun,
diabaikan, tetapi diperluas untuk mencakup studi kombinasi dan jumlah nutrisi dalam makanan
tentang semua faktor lingkungan eksternal yang dikonsumsilah yang menentukan
lainnya yang menentukan apa dan berapa kesehatan. Untuk membaca seseorang harus
banyak makanan dan nutrisi yang tersedia di mengetahui huruf-huruf alfabet; untuk melakukan
tingkat global. Kajian ini didukung oleh ilmu penjumlahan, seseorang harus dapat
sosial, perilaku, ekonomi, pertanian, dan politik. menghitung, menambah, mengurangi,
Oleh karena itu, studi tentang gizi manusia mengalikan, dan membagi. Untuk memahami
berupaya memahami kompleksitassosial dan bio nutrisi, seseorang harus mengetahui tentang
faktorlogis tentang bagaimana individu dan
populasi mempertahankan fungsi dan kesehatan
yang optimal, bagaimana kualitas, kuantitas dan nutrisi. Pembelajaran nutrisi, ABC dan kalkulasi
keseimbangan pasokan makanan dipengaruhi, numerik nutrisi, akan menjadi bagian utama dari
apa yang terjadi pada makanan setelahnya. perjalanan nutrisi siswa, dan harus mencakup:
dimakan, dan cara diet memengaruhi kesehatan
dan kesejahteraan. Pendekatan terpadu ini ● struktur kimia dan fisik dan karakteristik nutrisi
menghasilkan pemahaman yang lebih baik ● sumber makanan dari nutrisi, termasuk makanan
tentang penyebab dan konsekuensi malnutrisi, komposisi, cara makanan ditanam, dipanen,
dan hubungan antara gizi dan kesehatan. disimpan, diproses dan disiapkan, dan
4 Pengantar Nutrisi Manusia pengaruhnya terhadap komposisi nutrisi dangizi
nilai
● pencernaan, penyerapan, transportasi peredaran
1.4 Hubungan antara gizi dan darah, dan pengambilan nutrisi oleh sel, serta
kesehatan pengaturan lation dari semua proses ini
● metabolisme nutrisi, fungsinya, penyimpanan,
Gambar 1.2 menunjukkan bahwa individu dapat dan ekskresi
dikategorikan secara luas untuk memiliki status ● kebutuhan fisiologis (tuntutan atau persyaratan)
gizi yang optimal atau menjadi kurang gizi, gizi untuk nutrisi dalam kesehatan dan penyakit,
berlebihan, atau kekurangan gizi. Penyebab dan dan selama keadaan khusus (kehamilan,
konsekuensi utama dari status nutrisi ini menyusui, acara olahraga), juga sebagai
ditunjukkan. Penting untuk disadari bahwa variabilitas individu
banyak faktor gaya hidup dan lingkungan lainnya, ● interaksi dengan nutrisi lain, non-nutrisi
selain nutrisi, memengaruhi kesehatan dan (fitokimia), antinutrien, dan obat-obatan
kesejahteraan, tetapi nutrisi adalah faktor utama,
 Situasi gizi Konsekuensi kesehatan, hasil

Gizi optimal
Individu yang aman pangan dengan
Kesehatan, kesejahteraan, perkembangan normal,
memadai, seimbang dan bijaksana
berkualitas tinggi yang
pola makan

Kurang Gizi: kelaparan

Hidup individu yang tidak aman pangan
• Penurunan perkembangan fisik dan mental
dalam kemiskinan, ketidaktahuan, politik
• Sistem kekebalan yang
terganggu lingkungan yang tidak stabil ,
• penyakit menular Peningkatan
terganggu masyarakat,perang
• lingkaranVicious dari gizi,
keterbelakangan, kemiskinan

kelebihan gizi
berlebihan dari makanan,
Obesitas, sindrom metabolik, kardiovaskular
terutama macronutrients, ditambah:
penyakit, tipe 2 diabetes mellitus, tertentu
• aktivitas fisik yang
kankerrendah:NCD kronis , sering ditandai
• merokok, stres, penyalahgunaan alkohol
oleh kelebihan gizi macronutrients dan
gizi mikronutrien

Malnutrisi
Gizi transisi: Individu
beban ganda penyakit menular ditambah NCD,
dan masyarakat yang sebelumnya
sering ditandai dengan kelebihan gizi dari
makanan yang tidak aman → dihadapkan dengan
makronutrisi dan gizi dari
melimpah tarianenak
mikronutrien yang
makanan → beberapa kurang gizi, yang
lain terlalu banyak makronutrien
dan terlalu sedikit mikronutrien

Gambar 1.2 Hubungan antara gizi dan kesehatan. NCD, penyakit tidak menular.

● konsekuensi dari konsumsi yang kurang dan
konsumsi berlebihan dari nutrisi
● penggunaan terapeutik dari nutrisi
● faktor-faktor yang mempengaruhi keamanan
pangan dan nutrisi serta keamanan pangan.
Ada lebih dari 50 nutrisi yang diketahui
(termasuk asam amino dan asam lemak) dan
lebih banyak lagi bahan kimia dalam makanan
yang diduga mempengaruhi fungsi dan
kesehatan manusia (Kotak 1.1). Nutrisi tidak ada
dalam isolasi, kecuali air dan lainnya dalam
beberapa sediaan farmasi. Dalam makanan, di
usus selama pencernaan, fermentasi dan
penyerapan, di dalam darah selama
pengangkutan, dan di dalam sel selama
metabolisme, nutrisi berinteraksi satu sama lain.
Oleh karena itu, nutrisi tertentu tidak boleh
dipelajari secara terpisah, tetapi diintegrasikan
dengan nutrisi lain dan dilihat dalam konteks
fungsi tubuh total. Kajian gizi juga mencakup
bagaimana menentukan kebutuhan gizi untuk
membuat rekomendasi asupan dan bagaimana
status gizi dipantau dengan mengukur asupan,
antropometri, komposisi tubuh, penanda biokimia
yang mencerminkan status gizi, dan tanda klinis
malnutrisi.
Pengetahuan tentang nutrisi dan fungsinya
akan memungkinkan ahli gizi untuk menasihati
individu apa dan berapa banyak yang harus
dimakan. Namun demikian, pengetahuan tersebut
adalah
Kotak 1.1 Kelas Gizi untuk Gizi Manusia
Perspektif Global tentang Pangan dan Gizi 5
Merupakan tragedi besar bahwa jutaan orang
saat ini hidup dengan kelaparan, dan takut
kelaparan. Hal ini terlepas dari kenyataan bahwa
ketahanan pangan atau “akses untuk setiap saat,
ke pasokan berkelanjutan dari makanan yang
cukup dan aman bergizi untuk perkembangan
fisik dan mental yang normal dan
kehidupan yang sehat dan produktif” adalah hak
asasi manusia yang tertanam dalam konstitusi
sebagian besar negara berkembang. Terlepas
dari kenyataan bahwa makanan yang cukup
diproduksi di tingkat global (lihat Kotak
1.2).Makanan

belum cukup untuk memahami dan mengatasi

masalah malnutrisi global yang dihadapi umat
manusia saat ini. Persepsi ini telah menghasilkan
pengembangan disiplin ilmu sosial untuk
mendukung pengetahuan dari ilmu biologi untuk
mengatasi malnutrisi global.
1.6 Malnutrisi Global
Kotak1.2
Kerawanan pangan: ketika orang hidup dengan kelaparan,
dan takut kelaparan. Ketahanan pangan: akses untuk
semua, setiap saat, ke pasokan pangan yang memadai
dan aman secara bergizi dan terjangkau untuk
perkembangan fisik dan mental yang normal serta
kehidupan yang sehat dan produktif.

Kelas / kategori Subkelas / kategori Contoh nutrisi

Karbohidrat (makronutrien) Monosakarida Disakarida Glukosa, fruktosa, galaktosa Sukrosa, maltosa, laktosa Pati dan
Polisakarida serat makanan
Protein (makronutrien) Protein sumber nabati dan hewani Asam amino (n = 20): alifatik, aromatik , mengandung
sulfur, asam,dasar
lemakdan minyak (lipid) (makronutrien) tak jenuh tunggaltak jenuh ganda (n-3, n-6, Oleat (cis) dan elaidat ( trans) asam lemak
Asam lemak jenuh Asam lemakAsam n-9) Linoleat, α-linolenic, arachidonic,
lemak Asam palmitat dan stearat eicosapentaenoic, docosahexaenoic acid
Mineral (mikronutrien) Mineral dan elektrolit Unsur jejak selenium, tembaga, mangan, molibdenum,
Kalsium, natrium, fosfat, kalium, besi, seng, fluorida,kromium
Vitamin(mikronutrien) Retinol larut lemak (A), kalsiferol (D), tokoferol (E), vitamin K
larut dalam air Asam askorbat (C), tiamin (B1), ribofl avin (B2), niacin (B3),
piridoksin (B6) , Folat, cobalamin (B12)
Air Air Air
6 Pengenalan ManusiaGizi bawah usia 5 tahun menderita defisiensi protein
dan energi akut atau kronis. Diperkirakan 3,5–5
miliar orang kekurangan zat besi, 2,2 miliar
Ketidakamananmerupakan penghambat hak kekurangan yodium, dan 140–250 juta
asasi manusia, kualitas hidup, dan martabat kekurangan vitamin A. Hal ini telah menghasilkan
manusia. Diperkirakan, selama dekade terakhir beberapa inisiatif dan komitmen global, yang
abad ke-20, 826 juta orang kekurangan gizi: 792 dipelopori oleh sejumlah organisasi Perserikatan
juta di Bangsa-Bangsa, untuk mengurangi kekurangan
negara berkembang dan 34 juta di negara maju. gizi global, kerawanan pangan, kelaparan,
Di negara berkembang, lebih dari 199 juta anak di kelaparan, dan kekurangan gizi mikro. Beberapa
kemajuan telah dibuat dalam mengurangi angka-
angka ini, tetapi masalahnya masih jauh dari
terpecahkan. Beberapa inisiatifnya adalah:
● Pertemuan Puncak Dunia untuk Anak-anak yang
didukung oleh United Nations Children's
(Emergency) Fund (UNICEF) 1990, dengan
seruan untuk mengurangi kekurangan gizi
parah dan sedang di antara anak-anak di
bawah usia 5 tahun hingga setengah dari
angka tahun 1990 per tahun 2000, termasuk
tujuan untuk menghilangkan malnutrisi
mikronutrien
● Konferensi Internasional Organisasi Kesehatan
Dunia / Pangan dan Pertanian (WHO / FAO)
1992 tentang Gizi yang memperkuat tujuan
sebelumnya dan memperluasnya ke
penghapusan kematian akibat kelaparan
● FAO- 1996 mendukung KTT Pangan Dunia di
mana 186 kepala negara dan pemerintahan
menjanjikan kemauan politik dan komitmen
mereka terhadap rencana tindakan untuk
mengurangi jumlah orang yang kurang gizi
menjadi setengah dari jumlah mereka pada
tahun 1996 pada tahun 2015 ● pembentukan
Informasi Kerawanan dan Kerentanan Pangan
pada tahun 1997 dan Sistem Pemetaan (FIVIMS)
dan Kelompok Kerja Antar Lembaga (IAWG),
yang terdiri dari 26 organisasi dan lembaga
internasional dengan komitmen untuk
mengurangi kerawanan dan kerentanan pangan
serta penyebab multidimensi yang berakar pada
kemiskinan; Informasi tentang inisiatif ini dapat
diakses di: http://www.fao.org/
● Tujuan Pembangunan Milenium: Perserikatan
Bangsa-Bangsa mengartikulasikan delapan
tujuan, mulai dari mengurangi separuh
kemiskinan dan kelaparan yang ekstrim,
menghentikan penyebaran virus human menjadi agen pengembangan ketahanan pangan
immunodefi ciency ( HIV) / Acquired Immuno
Defi Ciency Syndrome (AIDS) dan
penyelenggaraan pendidikan dasar universal
tercapai pada target
tahun 2015; cetak biru dari tujuan ini telah yang menghancurkan, terutama di sub-Sahara
disetujui oleh semua negara di dunia dan
lembaga pembangunan terkemuka.
Laporan FAO tahun 2001 menunjukkan bahwa fisik
pada tahun 1997-1999 terdapat 815 juta orang
yang kekurangan gizi di dunia, 777 juta di
antaranya berada di negara berkembang, 27 juta
mereka seperti yang diikuti di negara industri.
di negara transisi
dan 11 juta di negara industri. Penurunan
tahunan orang yang kekurangan gizi dari periode
1990-1992 adalah 6 juta. Untuk mencapai tujuan
World Food Summit untuk mengurangi separuh
jumlah kekurangan gizi di negara berkembang
pada tahun 2015, diperkirakan penurunan
tahunan yang dibutuhkan adalah 22 juta.
Jelas, ini adalah tantangan besar bagi para
ilmuwan dan praktisi pangan dan gizi. Dibutuhkan
pendekatan holistik dan pemahaman tentang
faktor kompleks yang saling berinteraksi yang
berkontribusi terhadap malnutrisi di berbagai
tingkatan. Ini termasuk penyebab langsung,
sedang, mendasar, dan mendasar:
● tingkat individu atau penyebab langsung: asupan
makanan dan nutrisi, aktivitas fisik, status
kesehatan, struktur sosial, perawatan, tabu,
pertumbuhan, pilihan pribadi
● penyebab tingkat rumah tangga atau menengah:
keluarga ukuran dan komposisi, kesetaraan
gender, aturan distribusi makanan di dalam
rumah tangga, pendapatan, ketersediaan
makanan, akses ke pangan
● tingkat nasional atau penyebab yang
mendasarinya: kesehatan, pendidikan, sanitasi,
pertanian dan ketahanan pangan, perang,
ketidakstabilan politik , urbanisasi, pertumbuhan
populasi, distribusi dan konflik, perang,
bencana alam, sumber daya yang menurun
● penyebab tingkat internasional atau dasar:
struktur sosial, ekonomi dan politik, perjanjian
perdagangan, ukuran populasi, distribusi
pertumbuhan populasi, degradasi lingkungan.
Untuk mengatasi penyebab kekurangan gizi ini,
kerawanan pangan dan masyarakat yang
kelaparan dan individu harus diberdayakan untuk
dan mata pencaharian mereka sendiri. Yang
memperumit tugas memerangi kerawanan
pangan dan kelaparan adalah bencana alam
seperti kekeringan, banjir, angin topan dan suhu
ekstrim, perang yang sedang berlangsung dan
konflik regional, serta dampak HIV dan AIDS
Afrika.
Di banyak negara berkembang, penduduk asli
telah mengubah pola makan dan pola aktivitas
Program pemberian makanan mentary yang
luwes di negara-negara ini sering dikaitkan
dengan meningkatnya kecenderungan obesitas,
resistensi insulin, dan munculnya penyakit kronis
gaya hidup di beberapa segmen populasi ini,
sementara segmen lain masih kekurangan gizi.
Koeksistensi kekurangan gizi dan kelebihan
gizi, yang menyebabkan beban ganda penyakit
menular dan kronis, penyakit tidak menular, dan
penyebab multi faktor malnutrisi, membutuhkan
pendekatan inovatif untuk mengatasi kekurangan
gizi dan kelebihan gizi dalam program gizi
terpadu dan peningkatan kesehatan, dengan
fokus tentang nutrisi optimal untuk semua.
1.7 Hubungan antara ilmu gizi
dan praktek
Perjalanan melalui domain ilmiah gizi akan, pada
tahap khusus, bercabang ke jalan yang berbeda.
Jalan ini akan mengarah pada cakupan atau
cabang ilmu nutrisi yang berbeda yang tercakup
dalam teks kedua, ketiga, dan keempat dari seri
ini. Cabang-cabang ilmu gizi yang berbeda ini
dapat mengarah pada pelatihan spesialis gizi
untuk bidang praktik tertentu.
Tujuan utama dari ahli gizi adalah menerapkan
prinsip-prinsip gizi untuk meningkatkan kesehatan
dan kesejahteraan, mencegah penyakit, dan /
atau memulihkan kesehatan (mengobati penyakit)
pada individu, keluarga, komunitas, dan populasi.
Untuk membantu individu atau kelompok orang
untuk makan makanan yang seimbang, di mana
persediaan makanan memenuhi kebutuhan gizi,
melibatkan penerapan prinsip-prinsip gizi dari
bidang yang sangat luas ke hampir setiap aspek
kehidupan manusia. Oleh karena itu tidak
mengherankan bahwa cabang atau spesialisasi
nutrisi yang berbeda ini telah berevolusi dan
berkembang. Ini termasuk gizi klinis, gizi
masyarakat, kesehatan masyarakat, dan gizi
masyarakat. Diharapkan akan ada tumpang tindih
di bidang praktik spesialisasi ini.
● Ahli gizi klinis akan menasihati individu dari
paradigma biomedis – penyakit – perilaku untuk
meningkatkan kesehatan, mencegah penyakit,
atau mengobati penyakit. Ahli gizi klinis
sebagian besar akan bekerja dalam layanan
kesehatan (pengaturan berbasis fasilitas seperti
rumah
sakit, klinik, praktik swasta).
● Ahli gizi komunitas, dengan keterampilan
tambahan dari ilmu perilaku psikososial,
sebaiknya
Perspektif Global tentang Pangan dan Gizi 7
menyadari dinamika dalam komunitas tertentu
yang bertanggung jawab atas masalah gizi. Ini
mencakup ketahanan pangan rumah tangga,
latar belakang sosial ekonomi, tingkat
pendidikan, praktik pengasuhan anak, sanitasi,
air, sumber energi, layanan kesehatan,
dankualitas
indikatorhidup lainnya. Ahli gizi komunitas akan
merancang, menerapkan, dan memantau
program partisipatif komunitas yang sesuai
untuk mengatasi masalah ini.
● Kesehatan masyarakat atau ahli gizi masyarakat
mencakup bidang praktik kesehatan dan
perawatan tetapi juga akan peduli dengan
masalah keamanan pangan (pertanian) dan
lingkungan di tingkat publik. Kesehatan
masyarakat atau ahli gizi masyarakat, misalnya,
akan bertanggung jawab atas pengawasan gizi,
dan desain, implementasi, dan pemantauan
pedoman diet yang menangani masalah
kesehatan masyarakat yang relevan.
Pengetahuan latar belakang di bidang ekonomi,
pertanian, ilmu politik, dan desain kebijakan
sangat penting untuk perumusan dan
penerapan kebijakan gizi di suatu negara.
Banyak negara berkembang tidak akan
memiliki kapasitas atau sumber daya keuangan
untuk melatih dan mempekerjakan para
profesional untuk spesialisasi yang berbeda.
Akan tetapi, pelatihan khusus di masa depan dan
pekerjaan dari para profesional yang berbeda
dapat menghasilkan kapasitas untuk mengatasi
masalah gizi secara lebih efektif.
1.8 Tonggak Nutrisi: Perkembangan
Gizi sebagai Ilmu
Kepercayaan Kuno
Sepanjang keberadaan manusia, orang telah
mengaitkan kekuatan khusus dengan makanan
tertentu dan mengembangkan kepercayaan dan
tabu tentang makanan. Ini sering didasarkan
padaiklim, ekonomi, politik, atau agama
keadaan dan prinsip, tetapi juga pada
pengamatan tentang hubungan antara konsumsi
makanan dan kesehatan tertentu.
Contoh yang tercatat adalah filsuf China dan
India kuno yang menasihati tentang penggunaan
menghangatkan dan mendinginkan makanan dan
 rempah-rempah untuk kondisi tertentu dan untuk banyak contoh yang dicatat dengan cermat
"mengangkat jiwa", hukum Mosaik yang tentang bagaimana pengamatan awal (seringkali
didokumentasikan dalam Perjanjian Lama yang kuno dan primitif) tentang pola makan dan
membedakan antara makanan bersih dan najis, hubungan kesehatan mengarah pada penemuan,
puasa danhalal penjelasan fungsi, isolasi, dan sintesis berbagai
praktekIslam, dan biarawan Benediktin dari nutrisi. Mungkin contoh yang paling sering dikutip
Salerno yang mengajarkan penggunaan panas adalah uraian James Lind
dan lembab versus pada tahun 1772 tentang bagaimana buah jeruk
8 Pengantar Nutrisi Manusia dapat menyembuhkan dan mencegah penyakit
kudis pada pelaut dalam perjalanan panjang.
Faktor anti kudis (asam askorbat atau vitamin C)
makanan dingin dan kering untuk berbagai hanya diisolasi pada tahun 1921, ditandai pada
tujuan. Hippocrates, bapak kedokteran modern, tahun 1932, dan disinkronkan secara kimiawi
yang hidup dari 460 hingga sekitar 377 SM, dan pada tahun 1933. Contoh lain dari batu mil nutrisi
kemudian Moses Maimonides, yang hidup pada adalah induksi beri-beri pada unggas domestik
abad ke-12, mendorong orang untuk oleh Eijkman pada tahun 1897, pengamatan dari
mempraktikkan penghematan dan gaya hidup Takaki pada tahun 1906
yang bijaksana. Mereka, dan yang lainnya,
menasihati agar, agar panjang umur dan sehat,
seseorang harus menghindari terlalu banyak
lemak dalam makanan, makan lebih banyak bahwa beri-beri pada pelaut Jepang dapat
buah, cukup tidur, dan aktif secara fisik - nasihat dicegah dengan melengkapi pola makan beras
yang masih dimasukkan dalam modern, berbasis mereka dengan roti gandum, dan, pada akhirnya,
sains. pedoman diet abad kedua puluh satu! isolasi faktor penyebabnya, tiamin atau vitamin
B1, oleh Funk pada tahun 1911. Lainnya adalah
Keyakinan budaya Hadiah Nobel- memenangkan penemuan oleh
Persepsi bahwa makanan mewakili lebih dari Minot dan Murphy pada tahun 1926 bahwa
bagian penyusunnya masih benar. Makan anemia pernisiosa adalah gangguan nutrisi
bersama adalah bentuk interaksi sosial yang karena kekurangan vitamin B12 dalam makanan,
diterima. Ini adalah cara di mana kebiasaan dan deskripsi kwashiorkor sebagai keadaan defisiensi
adat istiadat budaya, status sosial, kekerabatan, protein oleh Cecily Williams pada tahun 1935,
cinta, rasa hormat, berbagi, dan keramahan dan penemuan pati resisten dan pentingnya
diekspresikan. Ilmuwan dan ahli nutrisi menyadari fermentasi kolon bagi manusia oleh ahli gizi dari
bahwa, ketika merumuskan pedoman pola makan Dunn Clinical Nutrition Center pada 1980-an.
bagi masyarakat tradisional yang hidup, Sejarah nutrisi modern seperti yang
kepercayaan budaya dan tabu harus dipraktikkan saat ini sangat menarik untuk dibaca,
dipertimbangkan dan digabungkan. Ada banyak dan para siswa didorong untuk meluangkan
contoh kebiasaan dan pola makan makanan waktu untuk itu. Hal ini sering kali dicirikan oleh
tradisional, seringkali berdasarkan apa yang keberanian yang menghancurkan hati dan
tersedia. Saat ini, dengan dunia menjadi desa wawasan yang mengejutkan. Contoh yang
global, budaya telah belajar dari satu sama lain, pertama adalah konsekuensi klinis, metabolik,
dan pola makan yang diasosiasikan dengan dan patologis yang didokumentasikan dengan
kesehatan yang baik, seperti pola makan cermat dari kelaparan dan kelaparan oleh
Mediterania, menjadi populer di antara banyak sekelompok dokter Yahudi pada tahun 1940 di
budaya. ghetto Warsawa: dokter yang mereka sendiri
sekarat karena kelaparan. Contoh yang terakhir
Kebangkitan pertama: pengembangan adalah studi oleh Price, seorang dokter gigi
basis bukti Amerika, yang mencoba mengidentifikasi faktor
Pengetahuan tentang efek kesehatan spesifik makanan yang bertanggung jawab atas
dari diet, makanan, dan nutrisi tertentu sekarang kesehatan gigi dan kesehatan secara
benar-benar didasarkan pada hasil eksperimen keseluruhan pada orang yang menjalani gaya
ilmiah yang kaku. Nutrisi berkembang secara hidup tradisional. Dia tanpa sadar menggunakan
bertahap sebagai ilmu, tetapi maju dengan paradigma fortigenik dalam penelitiannya,
kemajuan pesat selama abad kedua puluh. Ada meneliti kekuatan dan faktor yang membuat
 orang tetap sehat, jauh sebelum istilah itu kalangan perempuan. Gizi buruk, hambatan
didefinisikan atau nilainya diakui. utama bagi pembangunan nasional, merupakan
Saat ini, ribuan ilmuwan nutrisi meneliti penyumbang tunggal terbesar pada angka
berbagai aspek nutrisi di laboratorium dan studi kematian anak, dan gangguan pertumbuhan fisik
lapangan di seluruh dunia dan dan perkembangan mental anak-anak di negara
mempublikasikannya di lebih dari 100 jurnal berkembang dan maju. Selain itu, kombinasi gizi
ilmiah nutrisi buruk dan gizi lebih di komunitas yang sama,
internasional. Artinya, ilmu gizi melahirkan dalam satu rumah tangga, dan bahkan pada
pengetahuan baru berdasarkan metodologi individu yang sama sering dilaporkan. Contohnya
penelitian yang mapan. Berbagai jenis adalah ibu obesitas dengan anak yang kurang
eksperimen, yang bervariasi dari eksperimen gizi dan wanita obesitas dengan defisiensi zat gizi
molekuler di laboratorium, melalui intervensi klinis mikro tertentu. Persepsi bahwa masalah
terkontrol plasebo, tersamar ganda, hingga survei malnutrisi global ini akan diselesaikan hanya
epidemiologi observasional, dan eksperimen dengan cara yang
berdasarkan paradigma kesehatan (fortigenik) inovatif, multidisiplin, dan multisektorial telah
atau penyakit (patogenik), akan dibahas dalam mengarah pada kebangkitan baru kedua dalam
volume ini (Bab 13). Proses peer review dari hasil penelitian dan praktik nutrisi.
yang dipublikasikan telah membantu dalam
pengembangan pedoman untuk menilai seberapa 1.9 Tantangan masa depan
mungkin, mungkin, meyakinkan, dan dapat untuk penelitian dan praktik
diterapkannya hasil dari studi ini. Pengetahuan nutrisi Nutrisi
baru tentang nutrisi, makanan, dan hubungan
pola makan dengan kesehatan dan penyakit, oleh
dasar dan molekulernutrisi
karena itu, dihasilkan melalui proses di mana
banyak ilmuwan memeriksa potongan teka-teki Perkembangan luar biasa dalam beberapa tahun
yang berbeda di seluruh terakhir dari biologi molekuler dan ketersediaan
teknik baru yang canggih membuka bidang di
mana interaksi-gen dan manipulasi diet ekspresi
genetik akan meningkat perhatian (lihat Bab 15).
dunia dalam eksperimen ilmiah terkontrol. Oleh Efek lebih dari 12.000
karena itu, praktik nutrisi saat ini memiliki basis Perspektif Global pada Pangan dan Nutrisi 9
penelitian yang kuat yang memungkinkan para
profesional nutrisi untuk mempraktikkan nutrisi
berbasis bukti. zat berbeda dalam makanan nabati, yang belum
diklasifikasikan sebagai nutrisi, juga akan
diperiksa. Zat-zat ini diproduksi oleh tanaman
Kebangkitan kedua:
untuk tujuan hormonal, atraktan, dan
mengatasi malnutrisi global
kemoprotektif, dan terdapat bukti bahwa banyak
Ada sedikit keraguan bahwa peningkatan nutrisi di antaranya menawarkan perlindungan terhadap
telah berkontribusi pada peningkatan kesehatan berbagai kondisi manusia. Ada kemungkinan
dan kelangsungan hidup yang dialami oleh bahwa fungsi baru dari nutrisi yang diketahui, dan
manusia modern. Namun, angka global tentang bahkan nutrisi baru, dapat ditemukan, dijelaskan,
prevalensi kurang gizi dan gizi lebih menunjukkan dan diterapkan di masa mendatang.
bahwa jutaan orang tidak memiliki cukup
makanan, sementara jutaan orang yang makan Nutrisi Klinis dan Komunitas
terlalu banyak menderita akibat obesitas. Sangat
menggoda untuk menyamakan situasi ini dengan Saat ini, fokusnya telah berpindah dari
kesenjangan antara si miskin dan si kaya atau eksperimen sederhana dengan jawaban yang
antara negara berkembang dan negara maju, jelas ke studi di mana statistik canggih harus
tetapi situasinya jauh lebih kompleks. Obesitas, digunakan untuk membedah peran nutrisi khusus,
akibat dari kelebihan gizi, saat ini menjadi makanan, dan diet dalam penyakit multifaktorial.
masalah kesehatan masyarakat tidak hanya di Epidemiologi nutrisi sekarang ditetapkan sebagai
negara kaya, disiplin ilmu di mana pertanyaan-pertanyaan ini
maju, dan aman pangan tetapi juga di negara dapat diatasi. Sejumlah masalah mendesak harus
berkembang dan rawan pangan, terutama di diteliti dan hasilnya diterapkan, misalnya:
 ● penyebab biologis dan sosiologis dari obesitas negara-negara miskin terlihat jelas.
pada masa kanak-kanak, yang muncul sebagai Ahli gizi memiliki peran penting dalam
masalah kesehatan masyarakat global memastikan keamanan pangan untuk semua, hak
● kebutuhan gizi orang tua: pada tahun 2000, lebih asasi manusia, di masa depan. Salah satu fungsi

dari 800 juta semut penghuni Bumi berusia utama mereka adalah mendidik dan
lebih dari 60 tahun; untuk memastikan kualitas menginformasikan penduduk untuk tidak terlalu
hidup yang tinggi pada populasi lansia yang bergantung pada produk hewani dalam makanan
terus berkembang, lebih banyak lagi yang perlu mereka, yang produksinya
diketahui tentang kebutuhan nutrisi mereka menempatkan beban yang jauh lebih berat pada
● hubungan antara nutrisi dan fungsi kekebalan lingkungan daripada makanan nabati. Tantangan

dan bagaimana nutrisi yang ditingkatkan dapat utama adalah meyakinkan para pemimpin politik
membantu mempertahankan diri dari serangan dan pemerintahan bahwa mengatasi kekurangan
mikroorganisme; Mengingat meningkatnya gizi (hambatan utama dalam pembangunan
pandemi HIV / AIDS, lebih banyak informasi di nasional) dalam program berkelanjutan harus
bidang ini sangat dibutuhkan menjadi prioritas utama dalam masyarakat
● rekomendasi pola makan: meskipun cukup, bukti berkembang dan miskin. Tantangan lainnya

meyakinkan tentang efek nutrisi dan makanan adalah mengembangkan model yang didasarkan
terhadap kesehatan, ahli gizi umumnya belum pada dinamika dalam masyarakat dan, dengan
terlalu berhasil dalam memotivasi masyarakat menggunakan pendekatan hak asasi manusia,
untuk mengubah pola makan mereka menjadi untuk mengatasi kekurangan gizi tanpa
lebih sehat. Kita perlu tahu lebih banyak menimbulkan masalah gizi lebih. Ada contoh di
tentang mengapa orang membuat pilihan mana model seperti itu, yang dimasukkan ke
makanan tertentu untuk merancang pedoman dalam program pengembangan masyarakat, telah
diet yang peka budaya dan praktis yang akan sangat berhasil (misalnya, di Thailand).
berdampak positif pada pilihan diet. Pedoman
pola makan berbasis makanan yang sekarang
sedang dikembangkan di banyak negara
merupakan langkah pertama ke arah ini. Pangan fungsional: perkembangan baru
Pangan fungsional adalah makanan baru atau
Gizi kesehatan masyarakat baru, yang dikembangkan untuk memiliki manfaat
kesehatan tertentu, selain fungsinya yang biasa.
Tantangan paling penting yang dihadapi umat
Contohnya adalah olesan dengan tambahan
manusia di masa depan mungkin untuk
fitosterol, untuk menurunkan serum kolesterol
menyediakan makanan yang aman dan memadai
10 Pengantar Nutrisi Manusia
lipoprotein densitas rendah dan risiko penyakit
jantung koroner, dan pengembangan produk
bertepung dengan pati resisten dan indeks
makanan dan air bersih untuk semua dengan glikemik yang lebih rendah, untuk membantu
cara yang aman bagi lingkungan yang tidak akan mengontrol kadar glukosa darah. Pengembangan
mengganggu kemampuan generasi mendatang dan pengujian pangan fungsional merupakan
untuk memenuhi kebutuhan mereka. Selain bidang baru yang menarik. Makanan ini dapat
ratusan juta orang yang tidak makan cukup membantu memperbaiki atau memulihkan status
makanan untuk memenuhi kebutuhan mereka gizi pada banyak orang. Namun, lebih banyak
untuk hidup sehat dan aktif, tambahan 80 juta yang harus diketahui tentang biomarker yang
orang harus diberi makan setiap tahun. cocok untuk menguji keefektifannya, variabilitas
Tantangan untuk memberi makan umat manusia respons manusia terhadap produk makanan
di masa depan menuntut perbaikanpertanian tertentu, keamanan, pemahaman konsumen, dan
budayadi daerah yang dilanda kekeringan seperti bagaimana pesan kesehatan mereka harus
sub-Sahara Afrika, penerapan bioteknologi dirumuskan, diberi label, dan dikomunikasikan.
dengan cara yang bertanggung jawab, kerjasama
lintas disiplin dan lintas sektoral dari semua yang
Keamanan Pangan
terlibat, dan distribusi yang lebih baik dari
pasokan makanan sehingga makanan yang Pemberian pangan yang aman secara
terjangkau dapat diakses oleh semua. Perlunya berkelanjutan, bebas dari mikroorganisme, racun,
pertumbuhan ekonomi yang berkelanjutan di dan zat berbahaya lainnya yang menyebabkan
 penyakit, tetap menjadi tantangan besar.
Pengalaman terkini dengan hewan yang
menderita bovine spongiform encephalopathy
(BSE atau penyakit sapi gila) atau dari penyakit
kaki-dan-mulut, atau burung yang terinfeksi virus
influenza A (flu burung), telah menunjukkan
Kontaminasi mikroba
Bakteri dan kapang (fungi) penghasil toksin dan aflatoksin
Racun menyebabkan “keracunan makanan” dan aflatoksin bersifat karsinogenik
Racun alami
seperti sianida pada singkong, solanin pada
kentang; dapat diproduksi oleh keadaan yang
tidak normal, dapat berupa penghambat enzim
atau antivitamin
Pencemaran lingkungan
Logam berat dan mineral
Pemalsuan pidana, polusi industri
Zat dari bahan kemasan
Perubahan selama memasak dan pemrosesan
makanan
Gambar 1.3 Potensi zat berbahaya dalam makanan. DDT, dikloro-difenil-trikloroetana.
1.10 Perspektif tentang Masa Depan
Penelitian dan praktik gizi, meskipun telah ada
selama bertahun-tahun, masih dalam tahap awal
sebagai disiplin ilmu dasar dan terapan.
Mahasiswa nutrisi saat ini dan masa depan akan
mengambil bagian dalam kebangkitan nutrisi
kedua yang sangat menarik ini dan melihat
kematangannya
. Namun, untuk mempengaruhi nutrisi dan
kesehatan individu dan populasi secara efektif,
ahli gizi harus menjalin hubungan dan kemitraan
dengan profesional kesehatan dan pembuat
kebijakan lainnya, dan harus mengembangkan
proses berpikir lateral. Besarnya dan
kompleksitas masalah gizi yang dihadapi umat
manusia saat ini menuntut upaya multidisiplin dan
multisektorial bersama dari semua yang terlibat
untuk menyelesaikannya. Oleh karena itu, pesan
utama untuk melakukan perjalanan ilmu nutrisi
adalah bahwa kerja tim itu penting: seseorang
tidak dapat menempuh jalan ini sendirian;
dibutuhkan mitra dari berbagai disiplin ilmu.
Kebutuhan esensial lainnya adalah
pengembangan kepemimpinan yang
berkelanjutan di bidang nutrisi. Pemimpin di
setiap tingkat penelitian dan praktik diperlukan
betapa cepatnya masalah nasional dapat
internasional karena pemasaran produk global.
Daftar kemungkinan zat berbahaya dalam
makanan menekankan perlunya pemantauan
terus menerus terhadap pasokan makanan oleh
petugas kesehatan (Gambar 1.3).
Residu pertanian
Pestisida seperti DDT atau hormon yang
digunakan untuk mendorong pertumbuhan
seperti somatotropin sapi
Aditif yang disengaja
Pemanis buatan
Pengawet
Fitokimia
Karbohidrat yang dimodifikasi (untuk makanan
fungsional)
untuk menanggapi tantangan malnutrisi global
yang ada dan untuk menghadapi tantangan masa
depan.
Kemajuan modern dalam biologi molekuler dan
bioteknologi di satu sisi, dan masih adanya
malnutrisi global di sisi lain, semakin menuntut
evaluasi ulang terhadap norma, standar, dan nilai
etika untuk ilmu dan praktik nutrisi. Diperlukan
arahan
dari pemimpin yang bertanggung jawab (Kotak
1.3). Ada kebutuhan mendesak akan pedoman
etika dan kode etik untuk kemitraan antara
industri makanan, badan PBB, pemerintah, dan
akademisi
. Kemitraan ini diperlukan untuk mengatasi
malnutrisi global dalam program berkelanjutan.
Perspektif Global tentang Pangan dan Gizi 11
Kotak 1.3 Tantangan masa depan yang membutuhkan
kepemimpinan yang luar biasa
• Nutrisi molekuler dasar
• Interaksi nutrisi-gen
• Peran fitokimia dalam kesehatan
• Nutrisi baru? Fungsi baru?
• Gizi masyarakat dan kesehatan masyarakat
• Obesitas pada masa kanak-kanak
 • Persyaratan orang tua 15% protein, dan 3–5% mineral.
• Rekomendasi diet • Variasi komposisi tubuh antar individu besar, terutama
• Nutrisi pasien dengan human immunodefiency virus / karena variasi massa lemak. Variasi massa bebas lemak
didapat dari sindrom imunodefisiensi lebih kecil.
• Gizi masyarakat
• Untuk memberi makan umat manusia
• Keamanan pangan
• Makanan fungsional
• Untuk memastikan bahwa makanan baru efektif dan
aman
2.1 Pendahuluan
• Keamanan pangan
• Pemantauan berkelanjutan Umat manusia telah lama terpesona dengan
• Kemitraan dengan disiplin ilmu lain komposisi tubuh manusia. Berabad-abad yang
• Kepemimpinan lalu, orang Yunani membedah mayat manusia
untuk mendapatkan wawasan tentang struktur
dan bentuk tubuh manusia, dan gambar dari
Abad Pertengahan struktur otot kasar menghiasi
Siswa di bidang nutrisi, pada awal perjalanan dinding banyak galeri seni terkenal. Mereka
penemuan nutrisi sebagai ilmu, harus dihargai tidak hanya karena prestasi artistik
memanfaatkan banyak kesempatan untuk mereka, tetapi juga untuk apa yang mereka
mengembangkan kualitas kepemimpinan. ungkapkan dari karya para diseksionis pada
Semoga ini menjadi perjalanan yang bahagia, masa itu. Dengan kemajuan dalam
berbuah, dan seumur hidup dengan banyak pengembangan metode kimiawi analisis di abad
pelajaran yang dapat diterapkan dalam penelitian ke-20, studi tentang komposisi tubuh ini
dan praktik nutrisi untuk membuat perbedaan diterapkan pada jaringan tubuh, janin, dan mayat
dalam kehidupan semua. bayi yang baru lahir. Ilmuwan seperti Mitchell,
Widdowson, dan Forbes melakukan pekerjaan
Bacaan lebih lanjut paling penting dari analisis kimiawi pada mayat
dewasa selama 1940-an dan 1950-an. Saat ini,
Situs Web analisis aktivasi neutron memungkinkan
http://whq.libdoc.who.int/trs/who_trs_916 komposisi kimiawi tubuh manusia dipelajari
http://www.who.int/nutrition/en secara in vivo. Analisis kimiawi awal tubuh ini
http://www.ifpri.org memberikan wawasan
http://fao.org/ag/ agn / nutrisi / profi les_en.stm

2 © 2009 P Deurenberg.

Komposisi
Tubuh Paul Deurenberg

Pesan-pesan utama
• Data komposisi tubuh digunakan untuk mengevaluasi
status gizi, pertumbuhan dan perkembangan,
homeostasis air, dan status penyakit tertentu.
• Komposisi tubuh manusia dipelajari pada tingkat atom,
molekuler, sel, jaringan, dan seluruh tubuh. Level-level
tersebut saling terkait. • Tubuh manusia yang “berbobot
normal” terdiri dari sekitar 98%
oksigen, karbon, hidrogen, nitrogen, dan kalsium; 60–
70% air, 10–35% lemak (tergantung jenis kelamin), 10–
• Beberapa teknik langsung, tidak langsung, dan tidak
langsung tersedia untuk mengukur komposisi tubuh,
masing-masing dengan kelebihan dan kekurangannya
masing-masing.
• Pilihan metode akan dipengaruhi oleh ketersediaan
instrumentasi, invasi, dan bahaya radiasi pada subjek,
harga, akurasi yang diperlukan, dan tujuan aplikasi.
 • Interpretasi dan penerapan data dari pengukuran ● cellular
komposisi tubuh harus dilakukan dengan hati-hati dan
● tissue
harus mempertimbangkan batasan metode yang
● whole body.
digunakan, usia, jenis kelamin, dan kelompok etnis.

Of the many methods available to measure

body composition, a few are highlighted and a
short description of each is given. For more
detailed infor mation, the books by Forbes (1987)
and Heymsfi eld et al. (2005) on human body
ke dalam perubahan yang terjadi selama
composition are recom mended for further
pertumbuhan dan pengembangan. Mereka juga
reading.
menjadi dasar untuk sejumlah metode yang
sekarang banyak digunakan untuk menilai
komposisi tubuh in vivo. 2.2 Five levels of body composition
Saat ini, diketahui bahwa banyak penyakit dan
kelainan yang berkaitan dengan komposisi tubuh Human body composition can be studied at the
yang tidak normal atau perubahan komposisi atomic, molecular, cellular, tissue, and whole
tubuh. Kondisi yang paling umum dari kondisi ini body level. These fi ve levels are related to each
adalah obesitas, di mana jumlah lemak tubuh other. For example, information at the atomic
terlalu tinggi, yang menyebabkan kelainan pada level can be used, subject to certain assumptions,
metabolisme lemak dan karbohidrat, tekanan to provide informa
darah tinggi, dan diabetes pada orang dewasa. Di tion at the whole body level.
ujung lain dari spektrum nutrisi, malnutrisi energi
Atomic level
dan protein mengakibatkan penurunan jumlah
simpanan lemak dan protein dalam tubuh, dan Many chemical elements (atoms) are found in the
banyak penyakit yang berhubungan dengan human body, but the six elements oxygen,
kelainan pada total air tubuh atau dengan carbon, hydrogen, nitrogen, calcium, and
distribusi air tubuh ke seluruh tubuh. ruang phosphorus are the most abundant and together
intraseluler dan ekstraseluler. account for more than 98% of body weight (Table
Because of the high variability between 2.1). Indeed, the 11 most common elements
subjects in chemical body composition, mainly account for 99.5% of the atomic body
due to the high variation in body fat stores, the composition. This information was initially based
concept of fat-free on chemical analysis of carcasses, but today the
information can also be obtained by in vivo
neutron activation analysis (IVNAA). The classical
chemical cadaver analysis, as carried out mainly
mass (FFM) was introduced at the end of the nine in the 1940s,
teenth century. If body composition data are Body Composition 13
expressed as a proportion of the FFM, data
become much more consistent between
individuals. For example, the fraction of water in Table 2.1 Body composition at the atomic level of a 70 kg
reference man
the FFM (0.73 ± 0.02) is very consistent across
individuals, whereas the between subject Atomic element Amount (kg) Amount (% body weight)
variation is two to three times higher if expressed
per kilogram of body weight. This high variability Oxygen 43 61
in body components led to the defi nition of a Carbon 16 23
Hydrogen 7 10
“reference man,” an imaginary person with a
Nitrogen 1.8 2.6
given body composition. Calcium 1.0 1.4
In this chapter a (global) description of the com Phosphorus 0.6 0.8
position of the healthy human body is given and Total 69.4 98.8
dis cussed at the following levels:

● atomic Box 2.1

● molecular
 The water content in the body varies with age. In a fetus, Total 69.8 99.4
the water content slowly decreases from more than 90%
after conception to about 80% before delivery at about 7
months of gestation. A newborn has about 70% body women. In severe obesity body fat can be as high
water, which is about 82% of the fat-free mass. This value
slowly decreases further to 72% of the fat-free mass until
as 60–70% of body weight.
the body is chemically mature at age 15–18 years. In Body protein varies between 10% and 15%. It
general, males have more body water (related to body is higher in males than in females, as males
weight) than females, as their body fat content is lower. generally have more muscles. There is no protein
storage in the body and, generally speaking, loss
of protein coincides with a loss of functionality
given the high protein content and high protein
still forms the basis for many in vivo techniques
turnover rates in vital organs.
that are used to assess body composition.
The amount of minerals in the body varies
Molecular level between 3% and 5%, again dependent on body
fat. Calcium and phosphorus are the two main
The chemical elements in the human body are minerals. They are found mainly in bones.
bound in molecules and, in very global terms, the Carbohydrates are found in the body as glucose
main com partments are water, lipids, proteins, (blood sugar) and glycogen, a polysaccharide in
minerals, and carbohydrates. The total amount of muscle and liver cells that serves as a short-term
water in the body is high and, depending on the energy store. The amount of carbohy
body fat content, can be as high as 60–70% of drates in the body rarely exceeds 500 g. Table
total body weight. Total body water can be 2.2 gives the body composition of the reference
divided into intracellular water and extracellular man at a molecular level.
water, and the ratio of the two is an important
health parameter that is disturbed in many Cellular level
diseases (Box 2.1).
At the cellular level, body composition can be
Lipids appear in the human body in different
described in terms of body cell mass, extracellular
forms. Essential structural lipids such as the
fl uids, and extracellular solids. The body cell
phospho lipids (cell membranes) and
mass includes the cells with all their contents,
sphingomyelin (nervous system) form only a
such as water, proteins, and minerals.
minor part of the total lipids in the body. The
Extracellular fl uid contains about 95% water,
nonessential lipids, mostly triglycerides or
which is plasma in the intravascular space and
triacylglycerol (fat), are the most abundant. They
interstitial fl uid in the extravascular space.
are the energy store of the adult human body,
Extracellular solids are mainly proteins (eg, colla
insulate against cold, protect vital organs such as
gen) and minerals (bone minerals and soluble
the kidneys against mechanical damage, and, to
miner
a certain extent, enhance the body's appearance.
als in the extracellular fl uid). Body composition at
In a “normal weight” healthy adult, the amount of
the cellular level is not easy to measure, owing to
body fat varies between 10% and 25% in men
its complex nature. As will be discussed later, the
and between 15% and 35% in 40
K method can be used to assess body cell mass
14 Introduction to Human Nutrition
and some dilution techniques, for example
bromide dilution, can be used to assess
Table 2.2 Body composition at the molecular level of a 70 kg extracellular water.
refer ence man

Component Amount (kg) Amount (% body weight)

Table 2.3 Body composition at the tissue level of a 70 kg
Water reference man
Extracellular 18 26
Intracellular 24 34 Tissue/organ Amount (kg) Amount (% body weight)
Lipid
Essential 1.5 2.1 Muscle 28 40
Nonessential 12 17 Adipose tissue 15 21.4
Protein 10.1 14.4 Blood 5.5 7.9
Mineral 3.7 5.3 Bone 5 7.1
Carbohydrate 0.5 0.6 Skin 2.6 3.7
 Liver 1.8 2.6 established in earlier studies between body
Total 57.9 82.7 parameters (eg, skinfold thick
ness) and information on body composition (eg,
body fat by density), also enable the assessment
of body composition. Another example is the
Tissue level assess ment of body water based on weight,
Cells with equal functions form tissues, including height, age, and gender.
muscular, connective, epithelial, and nervous
tissue. Bones are connective tissue and consist 2.3 Relationships between
mainly of hydroxyapatite, [Ca3(PO4)2]3Ca(OH)2, different levels of body
bedded in a protein matrix. A rather simple body composition
composition model at the tissue level would be:
Body weight = adipose tissue + skeletal The fi ve levels of body composition are
muscle + bone + organs + rest interrelated. This means that information at one
level can be trans
Several of these components can now be
measured with, for example, computed
tomography (CT) or magnetic resonance imaging
(MRI) for adipose tissue; creatinine excretion or Box 2.2
N-methyl-histidine excretion in 24 h urine for
skeletal muscle; dual-energy X-ray Adipose tissue is made of adipocytes, which are cells that
absorptiometry (DXA) for bones; and MRI or ultra store triglycerides in the form of small fat droplets. Adipose
sound for organs. Body composition at the tissue tissue con
level is given in Table 2.3. Body Composition 15

Whole body level

Table 2.4 Methods used to determine body composition
Body composition measurements at the whole
body level use simple body parameters to give an Direct Indirect Doubly indirect Carcass analyses
insight into body composition. Formulae, based Densitometry Weight/height indices
on statistical relationships that have been
tains about 80% triglycerides and some 1– part is water plus electrolytes. During IVNAA Deuterium oxide dilution
2% protein (enzymes), and the remaining weight Skinfolds/ultrasound

loss adipose tissue decreases: the actual fat loss will be about 40K counting Circumferences/diameters
80%
of the actual weight loss. More-compartment models Impedance

lated to another level. This is important as it forms
the basis of many techniques used to determine
body composition. In the context of this chapter,
only a few examples are given. After determining
the amount of calcium in the body by, for
example, IVNAA (atomic level), the amount of
bone can be calculated assuming that a certain
amount of total body calcium is in the skeletal
tissue. Determination of total body potassium (by
40
K or IVNAA) enables the assessment of the
body cell mass, as most of the body potassium is
known to be intracellular. Skinfold thickness
measurements (total body level) enable the
assessment of body fat (molecular level).
Formulae used for these calcula tions are
component based, property based, or some
times a combination. Component-based formulae
are based on fi xed relationships between
components. An example is the calculation of
total body water from measured hydrogen: the
chemical formula of water determines the factor.
Property-based formulae are based on
established statistical relationships between
variables. An example is the prediction of body fat
percentage (body composition parameter) from
skinfold thickness (property) (Box 2.2). Property-
based formulae tend to be population spe cifi c,
which limits the widespread application.
Most body composition techniques that are in
use today are based on assumptions, often
derived from carcass analyses or experimentally water losses occur during the analytical work. As
derived from observational studies. Violation of early as the nineteenth century, it was rec
these assumptions leads to biased results, and ognized that the variation in chemical body
some methods are more prone to bias than composi tion was reduced when results were
others. In the following short description of expressed as a
different methodologies, the most important 16 Introduction to Human Nutrition
assumptions are highlighted.
fraction of the fat-free body. The data on the
2.4 Body composition techniques chemical composition of only a few human
cadavers form the basis for the assumptions that
Body composition techniques can be described in
are normally used in indirect methods. These
terms of direct, indirect, and doubly indirect
chemical analyses were per
methods.
DXA Infrared interactance formed in fi ve men and one woman. It was
CT/MRI scans Creatinine excretion concluded that, on the basis of FFM, the mean
amounts of water, protein, and minerals in the
IVNAA, in vivo neutron activation analysis; DXA, dual-energy body are 72.6%, 20.5%, and 6.9%, respectively.
X-ray absorptiometry; CT, computed tomography; MRI, The variability in these fi gures is about 13% for
magnetic resonance imaging.
protein and minerals and 4% for water. Although
one can question the quality of these data as a
● In direct methods, the body component of basis for other methods (low number, high
interest is determined directly without or with variation in age, variation in gender, some
only minor assumptions. Examples are carcasses were not analyzed immediately after
chemical carcass analy ses and IVNAA for the death), they form the basis for many indirect and
atomic components. doubly indirect body composition methods.
● In indirect techniques, the body component of Chemical carcass analy sis also revealed that the

interest is determined indirectly. Examples are amount of potassium in the FFM is fairly constant.
the determination of body protein from body This fact is used as the basis for the calculation of
nitrogen, assuming a constant conversion factor the amount of FFM or for body cell mass from
40
of 6.25 from nitrogen to protein, and the total body potassium, determined by K
determination of body cell mass using K. In scanning.
40

both examples, assumptions are used. These In the 1980s, cadaver studies were performed
assumptions may not be valid in the given again in the “Brussels study.” Unfortunately, only
situation or for the subject(s) under study and informa tion at a tissue level and not at atomic or
hence could lead to biased results. molecular level was collected. However, the need
● Doubly indirect methods rely on a statistical rela for cadaver studies has greatly diminished given
tionship between easily measurable body para that the same information can now be obtained in
meter(s) and the body component of interest. vivo by IVNAA.
Examples are the assessment of skeletal
muscle mass by creatinine excretion and the In vivo neutron activation analysis IVNAA is a
assessment of body fat from skin-fold thickness. relatively new body composition tech nique that
Table 2.4 gives an over view of the most allows the determination of specifi c chemi cal
common methods. elements in the body. The body is bombarded
with fast neutrons of known energy level. The
2.5 Direct methods neutrons can be captured by chemical elements
(as part of mol ecules) in the body, resulting in a
Carcass analysis transition state of higher energy for that element –
energy that is fi nally emitted as gamma rays. For
The (chemical) analysis of carcasses is a time
example, capture of neutrons by nitrogen results
consuming exercise and requires very precise
in the formation of the isotope 15N, which will emit
approaches to the task. The carcass has to be
the excess energy as gamma rays:
carefully dissected into the different tissues that
are then exactly weighed, after which the 14
N + 1n → 15N* + gamma rays
chemical analyses have to be performed. To 14 15
avoid errors it is important that no unaccounted where N is nitrogen with atomic mass 14, N is
nitrogen with atomic mass 15, and 1n is a neutron. in American football players was not the result of
With IVNAA, many elements in the body can be excess fat but of enlarged muscle mass.
determined, including calcium, phosphorus, nitro In underwater weighing, the weight of the
gen, oxygen, potassium, and chlorine. subject is fi rst measured in air and then while
The information obtained at the atomic level totally immersed in water. The difference between
can be converted to more useful information. For weight in air and weight under water is the
upwards force, which equals the weight of the
displaced water (Archimedes' law),
example, from total body nitrogen total body
protein can be calculated as 6.25 times the total
nitrogen, assuming that body protein consists of Box 2.3
16% nitrogen. The advantage of the method is
that the chemical body composition can be The density of the fat-free mass (FFM) can be calculated if
determined in vivo and can be compared with its com position is known.
other, indirect, techniques. For fundamental In the calculation example below it is assumed that the
studies and for validation of existing techniques in FFM consists of 73.5% water, 19.6% protein, and 6.9%
minerals with densities (at 37°C) of 0.993, 1.340, and 3.038
special groups of subjects, for example in kg/l, respectively. In addition, it is assumed that the
different ethnic groups, elderly subjects, obese volumes of the separate com
sub jects, or in the diseased state, the partments can be added up to the total volume of the FFM
methodology can be of great importance. The (in fact, the compartments do not “mix”). Thus, the volume
disadvantage of IVNAA is not only the price. The of the FFM equals the sum of the other compartments:
subject is irradiated, with the radiation dose used FFMvolume = Watervolume + Mineralvolume + Proteinvolume As
depending on the number and kind of elements to volume is weight/density, the equation can be written
be determined. It is relatively low for nitrogen
as: 100/DensityFFM = 73.5/0.993 + 6.9/3.038 +
(0.26 mSv) but high for calcium (2.5 mSv).
19.6/1.340
From this, the density of the FFM can be calculated as
2.6 Indirect methods 1.0999 kg/l. It is obvious that differences in composition of
the FFM will result in a different density.
Densitometry
The densitometric method assumes that the body
consists of two components, a fat mass, in which Box 2.4
all “chemical” fat is located, and the FFM, which
Siri's formula can be derived assuming that the body
consists of (fat-free) bones, muscles, water, and consists of fat mass (FM) and fat-free mass (FFM). If body
organs. Chemically, the FFM consists of water, weight is assumed to be 100% and body fat is x%, then
minerals, protein, and a small amount of FFM is 100 − x%. It is assumed that the volumes of these
carbohydrate, the last often being neglected. The two compartments can be added up to total body volume.
density of the fat mass is 0.900 kg/l and, from Then:
carcass analysis data, the density of the FFM can Bodyvolume = FMvolume + FFMvolume
be calculated as 1.100 kg/l, depending on the As volume is weight/density, the equation can be
relative amount of minerals, protein, and water in
the FFM (Box 2.3). written as: 100/body density = x/0.9 + (100 − x)/1.1
The density of the total body depends on the From this, body fat percentage (BF%) can be
ratio of fat mass to FFM. Once the density of the
calculated as: BF% = 495/density − 450
body has been determined, the percentage of fat
in the body (BF%) can be calculated by Siri's The general formula to calculate BF% from body density
formula (Box 2.4): ⎞ 1
(Db) is: ⎝⎜ ⎠⎟ DDD
BF% = (495/body density) − 450
Body Composition 17
Body density can be determined by several tech
niques, the oldest and perhaps most accurate
being underwater weighing. Behnke fi rst used
the tech nique, showing that excess body weight
 ure 2.1 Underwater weighing. The subject is submerged com
pletely and breathes via a respirometer (1) for simultaneous
residual lung volume measurement. Weight (W) (2) under
water (uw) is recorded and density (D) is calculated as Dbody =
Wair/(Wair − Wuw). Corrections are made for water temperature
and lung volume: percent age of fat in the body = 495/Dbody −
450.

fat is assumed to be maximal 3% of BF%. This

error is mainly due to violation of the assumption
that the density of the FFM equals 1.100 kg/l in
the subject under study. It can be argued that in
certain subjects or groups of subjects this
assumption may be vio

Fig

⋅ ⎛ D
=⋅ −
⎞ −
⎝⎜ ⎠⎟ −
BF% FFM FM DD FFM FM lated, as for children and in preg Siri's formula will
b
⎛
FM example in young nant women. Use of then lead to
DD FFM FM
In general, a lower density of the FFM than 1.1 kg/l will volume is assessed simultaneously.
result in an overestimation of BF% if Siri's formula is used. 18 Introduction to Human Nutrition
It is likely that the density of the FFM is lower in elderly
people, owing to bone mineral loss (osteoporosis).
Densitometry (using Siri's equation) overestimates
body fat compared with a four-compartment model (see Research to date has generally shown good
Figure 2.7). agreement between underwater weighing and air
displacement. Air displacement is better accepted
by the volunteers, but some experience diffi culties
from which, after correction for the water tempera because of the breathing pattern to be followed or
ture (density), the displaced water volume (and because of claustrophobia.
thus the body volume) can be calculated.
Corrections must be made for residual lung Dilution techniques
volume and air in the gut. Figure 2.1 shows an Carcass analyses revealed that the amount of
underwater weighing. The technique gives very water in the FFM is relatively constant at about
reproducible results within about 1% of BF%. The 73%. Total body water (TBW) can be determined
absolute error in determined body by dilution techniques. Dilution techniques are
biased conclusions. Some laboratories have generally based on the equation:
attempted to use water displacement instead of C1 ⋅ V1 = C2 ⋅ V2 = Constant
underwater weighing, but the technique failed,
mainly because of the diffi culty in accurately where C is the tracer (deuterium oxide, tritium, or
18
reading the water level in the tank. O water) concentration and V is the volume.
An air-displacement method has been commer When a subject is given a known amount of a
cially available since 1995. This method tracer
measures body volume after placing the subject
in a small, air tight chamber and increasing the
pressure by adding a known amount of air into
the chamber. Boyle Gay Lussac's law enables the Box 2.5
calculation of body volume. Corrections are made
for temperature and humidity changes, and lung A person with a body weight of 75 kg is given an exactly
 weighed dose of 15 g deuterium oxide. This deuterium
oxide is allowed to be equally distributed in the body water
n

compartment for about 3–5 hours. Then, blood is taken and tritium oxide, and 18O-labeled water. Other tracers
the deuterium concentration in the sample is determined.
Assuming the plasma level to be 370 mg/kg, the “deuterium
space” can be calculated as 15 000/370 = 40.5 kg. As
deuterium exchanges in the body with hydroxyl groups from
other molecules, the deuterium space has to be corrected
for this nonaqueous dilution (4–5%). Thus, total body water
is 0.95 ⋅ 15 000/370 = 38.5 kg. Assuming a hydration of the
fat-free mass of 73%, the body fat percentage of this 75 kg
weight subject would be: 100 ⋅ [75 − (38.5/0.73)/75] =
29.7%.
i

can also be used, such as alcohol and urea, but

they
Box 2.6

(C1 ⋅ V1), which is known to be diluted in a given

c

body compartment, the volume of that body com

s

a
e

i
p

are less suitable because they are partly

partment can be calculated from the dose given metabolized B

and (alcohol) or because they are actively excreted from

7
6
5
4
t

the concentration of the tracer in that compartment 3

2
1
0
-1
70 71 72 73 74 75 Percent water in FFM
f

y
the body (urea) during the dilution period. After
after equilibrium has been reached. Suitable tracers giving a subject the tracer and allowing around 3–
5 hours for equal distribution throughout the body,
determination of the concentration of deuterium in
blood, saliva, or urine allows the calculation of
TBW (Box 2.5).
Alternatively, other tracers can be used, such
d

o
as tritium oxide and 18O-labeled water, and the
for the determination of TBW are deuterium oxide, tracer can be given intravenously, which is
advantageous when the subject has
gastrointestinal disorders. The reproducibility of
the method is 1–3%, depending on the tracer
used and the analytical method chosen. From
TBW, the FFM, and hence fat mass, can be
calculated, assuming that 73% of the FFM is
b

water:
 BF% = 100 ⋅ (Weight − TBW/0.73)/Weight
The precision for estimations of body fat is about
3–4% of body weight. As with the densitometric
method, this error is due to violations of the
assump tion used (ie, that the relative amount of
water in the FFM is constant and equals 73% of
the FFM). In sub
For the computation of body composition from dual-energy
X-ray absorptiometry, especially body fat and lean tissue,
several assumptions are made, one of which is a constant
hydration of the fat-free mass (FFM). The fi gure shows
that the bias in calculated body fat percentage depends on
the hydration of the FFM.
Reference is a four-compartment model.
jects with a larger than 73% water content in the
FFM (pregnant women, morbid obese subjects,
and patients with edema), the factor 0.73 will
result in an overesti mation of the FFM. A three-
compartment model of the body that contains fat
mass, water, and dry FFM has a lower bias than a
two-compartment model. An overestimation of
body fat by densitometry, for example because of
a relatively high amount of water in the FFM, will
be counteracted by an underestima
tion using the dilution method (see also Box 2.6).
The use of tracers that do not cross the cell mem
brane enables the determination of extracellular
Body Composition 19

water (ECW). Commonly used tracers in this respect

3
are bromide salts or sodium-24. Intracellular water
(ICW) cannot be determined directly and is calcu
lated as the difference between TBW and ECW. rays from the body reveals 1
the amount of radioactive potassium, from which
Total body potassium TBK and hence FFM can be calculated. The
Chemical carcass analysis has revealed that the chamber in which the subject is scanned has to be
amount of potassium in the fat-free body is carefully
relatively con Men

2
F

stant, although the amount of potassium in

Women

K
25 45 65 85 Age (years)
different tissues varies widely. The determination shielded to avoid any background radiation
(cosmic radiation). The scanning of the body for
potassium lasts for 20–30 min and the
reproducibility is 2–3%.
Several authors have shown that the amount of
potassium in the FFM is different between males
and females, is lower in obese subjects, and is
B

probably also age dependent. Thus, TBK is much

more useful as a measure of body cell mass
T

of total body potassium (TBK) is relatively easy,

(BCM) than as a measure of FFM. However, this
owing to the natural occurrence of three potassium
39 40 41 discrepancy can be used to calculate the “quality”
isotopes ( K, K, and K), in constant relative
40 of FFM, defi ned as the ratio of cellular to
amounts, of which K is radioactive (gamma
extracellular components of FFM, or operationally
emission). Counting the emission of the gamma
as BCM/FFM. Thus, when TBK is used to assess thickness of the tissue. Therefore, correction for
BCM, and another method such as the body size has to be made. Compared with
hydrodensitometry or DXA is used to assess FFM traditional methods, DXA scanning is easy and
independently, it can be shown that the quality of widely available which, in turn, leads to prediction
FFM declines with age, along with the quantity 20 Introduction to Human Nutrition
(Figure 2.2). When potassium values are used to
calculate intracellular water, BCM, or FFM,
assuming constant amounts of potassium in equations for body composition based on DXA.
these body components, the same errors can However, as with other methods, DXA relies on
occur as with densitometry and dilu tion certain assumptions (Box 2.6) and there are
techniques. many publications showing that the error in body
Although the technique is easy to apply in composi
patients, the high cost of the scanning tion measurements using DXA can be
instrumentation limits its use other than in considerable (Figure 2.3). Moreover, identical
research settings. machines, even using the same software
versions, can give different results in scanning the
Dual-energy X-ray absorptiometry During DXA same person.
(also known as DEXA), the body or part of the
body is scanned with X-rays of two distinct levels Multicompartment models
of energy. The attenuation of the tissues for the Two-compartment models, consisting of fat mass
two different levels of radiation depends on its and FFM, lack validity in many situations where
chemi cal composition and is detected by the composition of the body is “abnormal.”
photocells. The Examples already mentioned are pregnancy,
Figure 2.2 Difference in total body potassium (TBK) content of morbid obesity, and the elderly. A combination of
the fat-free mass (FFM) between men and women and the techniques often results in more valid estimates,
relationship with age. as is the case when, for example, body density
and body water are com bined. In this particular
case, the body is divided into three
compartments:
instrument's software generates a two-
dimensional picture of the body or the body Body weight = Fat mass + Body water
compartment under study. The software can + Dry fat-free mass
calculate several body com ponents: bone
mineral content and bone mineral density, lean In this three-compartment model the variation
mass, and adipose tissue fat mass. These of the water content in the FFM is accounted for.
calculations are possible for each of the body There are fewer assumptions in this model,
parts, eg, for legs, trunk, spine, femur, and arms. leading to more valid results. Modern techniques
However, the method cannot distinguish between such as DXA enable the valid and precise
subcutane ous adipose tissue and discrete measurement of bone mineral, from which total
adipose tissue sites such as perirenal adipose body mineral can be estimated. When the mineral
tissue. The reproducibility of DXA is very high, content of the body is com
varying from about 0.5% for bone mineral density bined with body density and body water, a four
to about 2% for total body com position. The compartment model of the body is generated:
reproducibility for regional body com position is Body weight = Fat mass + Water
less. The method is quick and easy to perform + Minerals + Protein
and places very few demands on the subject. The
radiation dose (0.02 mSv) is only a fraction of the In this model, most of the variation in the amounts
radiation dose of a normal chest radiograph, and of the chemical components is accounted for,
hardly higher than the normal background. Apart result ing in a very reliable body composition
from repeated scanning, the radiation dose measure (Box 2.7). Four-compartment models
should not be a limiting factor in terms of can also be obtained using other techniques. For
volunteers being exposed to hazardous levels of example, the measure ment of calcium,
radiation. A disadvan tage of the method is that phosphorus, and nitrogen with IVNAA in
the attenuation of the X rays depends on the combination with TBW provides informa tion for a
model consisting of fat, minerals, protein, and
water. In the literature, models based on FM FM FM M
six compartments are also
described. However, they do not provide much M + PP
additional information and the increased technical
error negates the methodological advantage.
More-compartment models enable the best
possi ble estimate of body composition for
populations as FFM TBW TBW

The fi rst bar represents a two-compartment model of body

compo sition, in which the body is divided into fat mass
and fat-free mass (FFM). In the second bar, the FFM is
divided into water and a “dry” FFM, consisting of protein
and mineral. The third bar shows a four compartment
model in which the body is divided into water, protein,
mineral, and fat. The four-compartment model shown has
only minor assumptions and provides body composition
data that are very accurate.

well as for individuals. Although some studies

com paring body composition from four-
compartment models show that mean values
generally agree with simpler approaches, there
are also studies showing directional bias of two-
compartment body composi tion models. For this
reason, more-compartment models should ideally
be used as a reference (gold standard). However,
only a limited number of labo ratories can perform
all of the necessary measure ments for the
calculation of maximum compartment models.
Moreover, the data are expensive to collect, and
measurements are time-consuming and not very
practical in clinical situations.

Imaging techniques
CT scanning enables the visualization of tissues
in cross-sectional slices of the body. The
Fig thickness of those slices can vary, but is normally
about 1 cm. During CT scanning a source of X-
rays rotates per
pendicularly around the body or a body segment,
while photodetectors, opposite to the source,
register the attenuation of the X-rays after they
have passed through the body in the various
directions. The infor
ure 2.3 Dual-energy X-ray absorptiometer (DXA) scan using a mation received by the photodetectors is used to
HOLOGIC whole-body DXA (QDR-4500). Subcutaneous body
fat, bone, and muscle are distinguished by different colors. gen erate images. Software enables the
calculation of the
Body Composition 21

Box 2.7
 rela tively high level of radiation, the method is
limited to subjects for whom scanning is indicated
on clinical grounds. An alternative method to CT
scanning is MRI, which has the advantage that no
ionizing radia tion is involved.
During MRI, the signals emitted when the body
is placed in a strong magnetic fi eld are collected
and, as with CT scanning, the data are used to
generate a visual cross-sectional slice of the body
in a certain region. The determination of adipose
Fi tissue versus nonadipose tissue is based on the
shorter relaxation time of adipose tissue than of
other tissues that contain more protons or differ in
resonance frequency. MRI has the advantage
gure 2.4 Magnetic resonance imaging scan at the L4 level in
an obese subject. The white areas in the image are adipose over CT scanning that the subject is not exposed
tissue. Sub cutaneous adipose tissue and intra-abdominal to ionizing radiation. However, the time necessary
adipose tissue are sepa rated by the abdominal muscles. to make an MRI image is relatively long (minutes
versus seconds using CT), which has impli
amounts of tissues with different attenuation, for cations for the quality of the image. Any
example adipose tissue against nonadipose movement of the subject, even the movements of
tissue. The CT technique was introduced for body the intestinal tract when making images in the
composi tion assessments in the 1980s and is abdominal region, will decrease the quality of the
now widely used, predominantly for image.
measurements of body fat distri bution. Figure 2.4 As with CT scanning, images can be combined
shows a scan of the abdomen at the level of the to obtain information on total body composition.
umbilicus, made by MRI, a technique that gives Infor mation about organ size can be obtained
comparable information. The precision of the with a high accuracy. For example, MRI is used to
calculation of a tissue area or tissue volume from study the con tribution of various organs to the
the same scan(s) is very accurate, with an error resting metabolic rate of the total body.
of about 1%. Partial volume effects (pixels that Both CT scanning and MRI are expensive, and
contain tissue with different attenuation) may infl therefore their use will remain limited to a few
uence the accu racy and reproducibility of the labo ratories and for very specifi c situations.
method.
A single CT scan provides only relative data,
for example in a scan of the abdomen the relative 2.7 Doubly indirect methods
amount of visceral adipose tissue to
subcutaneous adipose tissue. Multiple CT Anthropometry
scanning allows the calculation of tissue volumes. Weight/height indices
From adipose tissue volumes (tissue level) and A weight/height index aims to correct body weight
an assumed density and composition of the for height. As a measure of body composition, for
adipose tissue, the amount of fat mass (molecular example body fat, a weight/height index should
level) can be calculated. Multiplying tissue have a high correlation with body fat, but also a
volumes with specifi c densities of these tissues low cor
(determined in vitro) allows a recalculation of the relation with body height, otherwise in short
body weight, a necessary but not suffi cient people body fat would be systematically
exercise for validation of a whole body technique. overestimated or underestimated.
Research in this area has shown that the CT In the literature, a number of weight/height
technique allows the determina tion of total body indices have been proposed. Examples are the
composition, with an error of estimate for fat mass Quetelet index or body mass index (BMI:
of 3–3.5 kg (compared with densitometry). weight/height2), the Broca index [weight/(height –
22 Introduction to Human Nutrition
100)], and the Benn index (weight/height p, in
which the exponent p is popula
CT scanning is expensive and, because of the
 Table 2.5 Classifi cation of weight in adults according to body
mass index
Classifi cation Body mass index (kg/m2) Risk of comorbidities
Underweight <18.5 Low
Normal range 18.5–24.9 Average Overweight
>25.0
Preobese 25.0–29.9 Increased Obese class I 30.0–
34.9 Moderate Obese class II 35.0–39.9 Severe
Obese class III >40 Very severe
Reproduced with permission of the World Health Organization.
tion specifi c). The Quetelet index or BMI is the
most widely used index today. Its correlation with
body fat is high (depending on the age group r =
0.6–0.8) and the correlation with body height is
generally low. The World Health Organization
(WHO) promotes the BMI as a crude indicator for
weight judgment. In Table 2.5 the cut-off points
for underweight, normal weight, overweight, and
obesity according to the WHO are given. These
cut-off values are based on the relation of BMI
with mortality and with risk factors for disease as
found in Caucasian populations. For non-
Caucasian populations other cut-off values may
Box 2.8 Box 2.9
Recent studies have shown that the relationship between
body mass index (BMI) and body fat percentage differs
among ethnic groups. For example, compared with
Caucasian populations some Asian populations have 3–
5% more body fat for the same BMI, age, and gender.
These differences can be explained by differences in body
build or frame size, subjects with a smaller frame having
more body fat at the same BMI.
These differences can have important consequences
for the defi nition of obesity (based on BMI cut-off values)
and the preva lence of obesity in a population. In
Indonesia, obesity has recently been redefi ned as BMI ≥
27 kg/m2. At this BMI, Indonesians have a similar body fat
to Caucasians with a BMI of 30 kg/m2. The lower ing of the
cut-off point for obesity from 30 to 27 kg/m2 increased the
prevalence of obesity from less than 5% to over 10%.
Recently an Expert Consultation of the World Health
Organiza tion (WHO) resulted in new guidelines to redefi
ne “action points” in non-Caucasian populations. For this
not only was the different relationship between BMI and
body fat percentage important, but also the high levels of
cardiovascular risk factors at low BMI values (WHO, 2004).
apply (WHO, 2004).
The cut-off values for BMI as in Table 2.5
cannot be used in children. In younger children,
weight com pared with height is relatively low,
and so is the BMI. During growth, the increase in
weight is larger than the increase in height and,
consequently, the BMI increases with age during
the pubertal phase of life. There are age-related
BMI cut-off values for obesity for children.
The BMI can also be used as a predictor for the
percentage of body fat. Several studies have
been pub lished in which a good relationship
between the BMI and the amount of body fat
(either as fat mass or as body fat percentage)
was demonstrated. The relation ship between
BMI and body fat percentage is age and gender
dependent and is different among certain ethnic
groups (Box 2.8). When using such age- and
gender-specifi c prediction equations, body fat per
centage can be predicted with an error of 3–5%.
This error is similar to the prediction error of other
doubly indirect methods, for example skinfold
thickness or total body bioelectrical impedance
measurements. The disadvantage of these
prediction formulae is that they obviously cannot
be used in certain subjects or
Body Composition 23
groups of subjects such as pregnant women or
body builders. However, other predictive methods
also have their limitations in these subjects.
TBW can also be predicted from weight and
height, and numerous prediction formulae have
been pub lished. The standard error of estimate
of these equa tions is, however, considerable.
Skinfold thickness measurements Body fat is
located both internally and subcutane ously. If
one assumes a constant relationship between
subcutaneous fat and total body fat, then total
body fat can be estimated by measuring the
amount of the subcutaneous adipose tissue. The
amount of subcu taneous tissue can be estimated
by measuring the thickness of the subcutaneous
fat layer at different sites of the body using a
skinfold caliper, infrared interactance, or
ultrasound measurements. In a given age group,
the relation between subcutaneous fat and total
fat is indeed relatively constant. However, the
relationship is different between males and
females, females having relatively more internal
fat (Box 2.9). Thus, it is possible by using age-
and gender-specifi c prediction equations to
assess the total amount of body fat by measuring
skinfolds at different sites of the body.
Skinfolds can be measured all over the body.
The most often measured skinfolds for the
assessment of
From Table 2.6 it can be seen that for the same amount of
subcu taneous fat (identical skinfold thickness) women
have more body fat than men. This is because of the
higher internal (organ) fat content in women. It can also be
seen (in both females and males) that at equal skinfold
thickness older people have more body fat: with age the
amount of internal fat increases.
igure 2.5 Measurement of the biceps skinfold.
total body fat are skinfolds on the upper arm
biceps (Figure 2.5) and triceps, under the scapula
(subscapu lar) and above the iliac crest
(suprailiac). The sum of more skinfolds is
normally used to reduce the error in
measurement and to correct for possible differ
ences in subcutaneous body fat distribution
between subjects within the same age and
gender group. Various prediction formulae for
body fat from skin fold thickness have been
published. For children, in whom the relationship
between skinfold thickness and body fat depends
on biological age, separate for mulae must be
used.
Measuring skinfolds adequately requires a
trained and experienced observer, otherwise
excessively large errors in the assessment of the
body fat can occur. A disadvantage of the method
is that the subject has to be partly undressed.
This hampers the use of the method in
epidemiological studies. In bed-ridden or
seriously ill patients the measurement of the trunk

24 Introduction to Human Nutrition

Table 2.6 Body fat percentage from the sum of four skinfolds (biceps, triceps, subscapular, suprailiac) in males and

females of different agesa Age (Males) Age (Females)

Skinfolds (mm) 17–29 30–39 40–49 ≥50 60–87 17–29 30–39 40–49 ≥50 60–87

20 8 12 12 13 24 14 17 20 22 34 30 13 16 18 19 27 20 22 25 27 37 40 16 19 21 23 29 23 26 28 30 39 50 19 22 25 27 31
27 28 31 33 41 60 21 24 27 29 32 29 31 33 36 42 70 23 25 29 32 33 31 33 35 38 43 80 25 27 31 34 34 33 34 37 40 44
90 26 28 33 36 35 35 36 38 41 45 100 28 29 34 37 36 36 37 40 43 46 110 29 30 36 39 37 38 39 41 44 47 120 30 31 37
40 37 39 40 42 45 48 130 31 32 38 42 38 40 41 43 46 48 140 32 33 39 43 38 41 42 44 47 49 150 33 34 40 44 39 42 43
45 48 49

a
Based on data from Durnin and Womersley (1974) for the age groups 17 to ≥50 and on Visser et al. (1994) for the elderly.

skinfold thicknesses can be diffi cult. This can be thickness at the upper arm, for example the
over come by measuring only the skinfold triceps. However, the error can be large because
the triceps does not neces sarily represent the stature), when comparing stature-adjusted body
total amount of subcutaneous fat. With advancing composition between young and old people.
age, the triceps skinfold becomes less Circumferences of the extremities or the trunk
representative of total body fat. are used to obtain information on body
In elderly subjects, the correlation between skin composition.
fold thickness and total body fat as measured by
densitometry is generally lower than in young and
middle-aged adults. This is due to an increased
amount of internal fat in the elderly. Obese From the mid-arm circumference, in combination
subjects are diffi cult to measure and the error is with the triceps skinfold thickness, information on
large even when measured by trained observers. muscle mass and fat mass of the upper arm can
This is also the case in subjects with edema, in be obtained. Circumferences of the trunk at waist,
whom the thickness of the subcutaneous adipose hip, and upper thigh level are used as indicators
tissue is easily overesti of body fat distribution. The WHO suggests an
mated. In patients with human immunodefi ciency upper limit waist-to-hip circumference ratio above
virus (HIV) lipodystrophy, peripheral 0.85 for females and 1.00 for males for abdominal
subcutaneous fat may be almost absent, while fat distribu tion. Diameters can also be used to
abdominal fat is increased. In this situation, gain insights into body fat distribution. A high
skinfolds can be very mis sagittal diameter com
leading as indicators of total body fat, and should pared with transverse diameter of the abdomen is
be used only to assess regional fat. indicative of an enlarged amount of visceral fat.
The calculation of the body fat percentage once However, it has to be kept in mind that the
the skinfolds have been measured is very simple. relation ship between these anthropometric
For a given skinfold thickness, the amount of parameters of body fat distribution and the intra-
body fat can be read from a table (Table 2.6). abdominal fat determined by CT or MRI scan is
The prediction error in body fat percentage is generally low. Changes in internal fat detected by
3– 5% compared with densitometry, depending MRI are only weakly associated with changes in
on age, these anthropo metric parameters of fat
gender, and level of body fatness. Given the distribution.
possible error in densitometry (3%), this means
Infrared interactance
that in extreme cases body fat from skinfolds can
be as much as 10– 15% off. The principle of infrared interactance is based on
differences in absorbance and refl ection of
Other anthropometric variables infrared light in different tissues. When the
Measurements of widths of skeletal diameters absorbance of near-infrared light (700–1100 nm)
provide an indication of the amount of skeletal is measured at well-defi ned sites of the body,
mass. There are formulae that allow the information on the thickness of the subcutaneous
calculation of the skeletal mass from body height, adipose tissue layer can be obtained. In the
wrist diameter, and knee diameter. The current prediction formulae used, many other parameters
reference data for ideal weight in the USA use the are included, such as weight, height, age, and
elbow diameter to classify people into three types gender, and it has been argued that the prediction
of body build. of body fat depends more on these parameters
In elderly subjects, the measurement of stature than on the measured infrared interaction.
can be diffi cult owing to kyphosis and/or
Ultrasound measurements
shrinkage of the spinal vertebrae. Knee height
can then be used to predict the maximal stature Ultrasound measurements can also be used to
during lifetime. Arm span is also used for that study several aspects of body composition. With
purpose. However, one has to realize that the ultrasound measurements, the thickness of the
current prediction formulae are developed in subcutaneous fat layer can be determined and
younger populations, in which the rela total body fat can be calculated. A good refl ection
tionship between stature and surrogate measure signal depends heavily on the absence of
ments may be different. In addition, the prediction connective tissue between adipose tissue and
error (3–5 cm) is relatively high. Knee height can muscle. The main diffi culty in ultrasound
also be used by itself (without predicting total measurements is the exact application of the ultra
sound transducer perpendicular to the tissue of the FFM, bioelectri cal impedance
without any pressure. In the literature, several measurements can also be used for the
studies report a good correlation between skinfold prediction of the FFM and hence body fat
thicknesses mea percentage. For those prediction equations, the
sured by calipers and those measured by impedance index was related to measures of
ultrasound. The correlation of skinfolds with total FFM, normally obtained by densitometry or by
body fat was higher than when using ultrasound, DXA.
suggesting that Body impedance depends on the frequency of
Body Composition 25 the current used and on body water distribution
between the extracellular and intracellular space
and between the different geometrical body
skinfold thickness measurements have a better compartments (legs, trunk, and arms). This calls
pre dictive value. for extreme caution in the
Internal abdominal body fat can also be 26 Introduction to Human Nutrition
assessed with ultrasound. Studies have shown
that ultrasound measurements provide a better
method than anthro pometry to assess internal
Box 2.10
abdominal fat depots.
The relative validity of impedance prediction formulae can
Bioelectrical impedance be demonstrated by a simple calculation example. A man,
In bioelectrical impedance, a small alternating aged 35 years, of height 170 cm, weight 75 kg, and
current is applied to the body. It is assumed that measured impedance (from foot to hand) 400 Ω, has a
predicted fat-free mass (FFM) of 64.7 kg according to
the body consists of different components, of Lukaski et al. (1986) and a predicted FFM of 60.5 kg
which water and dissolved electrolytes are able to according to Segal et al. (1988). Both prediction formulae
conduct the current. Hence, body impedance is a were developed in US populations and were cross-
measure of body water. The electrical resistance validated. The instrument used was the same and the
method of reference in both studies was underwater
or impedance of an electro
weighing.
lyte solution depends on several factors, of which
the most important are the amount of electrolytes
(pure water does not conduct the current), the
kind of electrolytes, and the temperature of the
solution. If currents of low frequency (<5 kHz) are interpretation of calculated body composition
used, body impedance is a measure of ECW, as values in situations where body water distribution
a low-frequency current cannot penetrate the cell can be disturbed, as is the case, for example, in
membrane, which acts, with its layers of protein, dialysis patients and in ascites. In general,
lipids, and proteins, as an electrical capacitor. prediction formulae based on impedance values
With increasing frequencies the capacitor are strongly population specifi c, and age and
features of the cell membrane diminish and gender are important contribu
gradually ICW also participates in the conduc tors. Differences between populations and
tance of the current, resulting in lower impedance individuals are partly caused by differences in
values at higher frequencies. Hence, at higher fre body build (eg, relatively long legs), which is not
quencies, TBW is measured. TBW and ECW can surprising, as the legs contribute most to total
be predicted from impedance at high and low body impedance relative to other parts of the
frequency, respectively, using empirically derived body (Box 2.10).
prediction for mulae. Other parameters are often Currently available impedance analyzers vary
taken into consid in their electrical features and in their principles.
eration, such as body weight, age, and gender. Many companies have developed impedance
Most prediction equations are based on statistical analyzers for personal use, anticipating
relationships between empirically measured im considerable interest among the public in
2
pedance index values (height /impedance) and determining their body fat per
body water values obtained by dilution techniques centage. There are instruments that measure
such as deuterium oxide dilution (for TBW) and imped ance from foot to foot while standing on a
bromide dilution (for ECW). As body water in weighing scale and provide not only body weight
healthy subjects is an assumed fi xed part (73%) but also body fat percentage. Other instruments
measure imped ance from hand to hand and mass. As the cell cannot recycle creatinine, the
allow the reading of body fat percentage, using a kidneys excrete it. Since metabolized creatine
built-in software program in which weight, height, phosphate is not the only source of urinary
age, and gender have to be entered. creatinine (eg, creatinine in ingested meat is also
Combinations of foot-to-foot and hand-to excreted immediately), the validity of the method
hand impedance analyzers are also marketed. As is dubious. A day-to-day coeffi cient of variation in
for all other impedance analyzers, the incorpo the excretion of creatinine of almost 20% is
rated formulae are population specifi c and have reported, when the subject is “free living” and the
a prediction error of 4–5%. This means that, apart urine is sampled over constant periods of 24
from a systematic error (prediction formula is not hours. The high variation is due to the ingestion of
valid), the value can be as much as 10% off in creatinine with nonvegetarian meals, differences
extreme cases. This kind of error is similar to the in physical activity levels, and variation in
possible error in skinfold thickness creatinine excretion within the phase of the
measurements, and hence impedance is no menstrual cycle. After careful standard ization,
better than skinfold thickness mea surements. which includes a controlled diet, the day-to
The advantage of impedance analyzers is that day variability in excretion can be decreased to
there is no need to undress and measurements about 5%. To obtain a reliable assessment of the
are less prone to observer bias. creatinine excretion, sampling of urine over
multiple 24 hour periods is necessary.
The excretion of 3-methylhistidine has also
been proposed as a measure for muscle mass.
Total body electrical conductivity FFM deter
Total body electrical conductivity (TOBEC) was
devel oped in the 1970s. The principle of the
method is that conductive material (body water
and dissolved elec trolytes) that is placed in an mined by densitometry correlates well with
electromagnetic fi eld will cause an inductive excreted 3-methylhistidine. The chemical
current, which is related to the amount of determination of 3-methylhistidine is, however,
conductive material. In practice, the subject lies more complicated than that of creatinine. A
on a stretcher, which enters the inner space of an unique feature of 3-
electric wire coil, through which a high-frequency methylhistidine is that it gives a measure of
current (2.5–5 MHz) passes. The measurement is muscle protein breakdown. Given the greater
very quick (it takes only seconds), painless, and expense of measuring 3-methylhistidine and the
without any risk to the subject. The reproducibility limited benefi t for muscle mass estimates, it is
of a mea surement is within 2% and the error in probably best to use it primarily for turnover
the predicted FFM was found to be about 3 kg in studies.
a group of adult subjects, which is similar to, for The main disadvantages of creatinine and 3-
example, skinfold thickness measurements or methylhistidine excretion as measures for body
impedance measurements. The TOBEC method com position are the large variability in excretion,
is especially suitable for mea surements in infants the necessity to follow a controlled (meat-free)
and young children, in whom bioelectrical diet for several days before and during the urine
impedance measurements are diffi cult or collections, and the diffi culties associated with
impossible to perform, owing to movement. The collecting 24 hour urine samples.
main disadvantage of the method is the high
Use and misuse of body composition data
price.
Information on the body composition of groups of
subjects or individuals is important, as body
Creatinine excretion and compo sition is an indicator of nutritional status
N-methyl-histidine excretion and also provides information about acute water
In the muscle cell, creatine phosphate, necessary homeostasis. Depending on what information is
for the energy metabolism of the cell, needed, several methods are available. However,
degenerates to cre atinine at a constant daily rate all have their advan tages and limitations. The
of about 2%. It is assumed that 1 g of excreted price of the method (both the instrument and the
creatinine is equivalent to 18–22 kg of muscle required personnel), the eventual stress and
danger (eg, radiation) for the subject, and the
time necessary to obtain the informa tion
determine the choice of the method, as well as
Body Composition 27
the required accuracy. The use in epidemiological
studies is different than that in clinical situations
or in physiological research. Table 2.7 provides a
“buyer's guide” to the several methods discussed
in this chapter. It is diffi cult to generalize as to
which method should be used in a given study.
Apart from the factors mentioned in Table 2.7,
availability plays an important role. Some
situations are discussed below.
For the description of body fatness of a large
general population group, the calculation of the
body fat per centage from the BMI may be as
good as or even better than the more expensive
information obtained from bioelectrical impedance
or the laborious mea surement of skinfold
thicknesses. Whichever method is to be used for
the prediction of body fat percentage in the
population, it is important to remember that the
formulae used should have been validated in the
population under study. The fact that a formula is
cross-validated by the authors who published the
formula does not mean that the formula is valid in
another population. The use of the Durnin and
Womersley (1974) equations for estimating body
fat from the sum of four skinfolds may be correct
if the population is adult but younger than about
60 years. In older subjects, the amount of body
fat is likely to be underestimated with these
formulae. Therefore, their use and thoughtless
interpretation in the elderly would lead to
completely wrong conclusions about

Table 2.7 Buyer's guide to different methods used to determine body composition

Method Accuracy Expenses Radiation Time Convenience for subject

Carcass analysis + + + − −
Neutron activation + + + − − − − + + + + Densitometry + + + + + +/− Dilution method + + +/− (−) + + 40K method +
+ − + + + + DXA + + + +/− − + + + + More-compartment models + + + − − −+ CT scanning + + − − − + + + + MRI
scanning + + − + + + Anthropometry + + + + + + + Infrared interactance + + + + + + + Bioelectrical impedance + +
+ + + + + + + TOBEC + − + + + + Creatinine/N-methylhistidine excretion + + − −

DXA, dual-energy X-ray absorptiometry; CT, computed tomography; MRI, pencitraan resonansi magnetik; TOBEC, total body
electrical conductivity.
+ + +, excellent; + +, very good; +, good; +/–, reasonable; –, bad; – –, very bad.
28 Introduction to Human Nutrition Dutch Asian

Age (years) 21 22 Height (cm) 175.6 158.0 Weight (kg)

body composition in the elderly and changes in
body composition with age. For the same reason, 77.2 43.9 Relative sitting height 0.53 0.55 Wrist (cm) 5.6
the BMI as an indicator of body fatness is only
suitable when corrections for age and gender are 4.4 Knee (cm) 9.1 8.4 Arm span (cm) 177.0 157.5 Waist
made; for example, a BMI of 25 kg/m 2 at the age
of 20 years refl ects a much lower percentage of (cm) 82.0 62.2 Hip (cm) 104.0 87.0 BMI 25.0 17.6 BF
body fat than at the age of 70 years. Body fat in
females is always higher than body fat in males (%) 31.1 29.6
with the same BMI and age. Recent studies have
shown that the relationship between BMI and
body fat percentage also differs among ethnic
groups. Figure 2.6 gives a good insight on how
misleading a low BMI can be. The biodata also among other factors, and there are no universally
show the differences in body build, which is at valid prediction equations based on impedance.
least part of the reason for the paradox. Choice of method to assess body composition in
Prediction equations based on impedance are a population would also take into consideration
dependent on body build, within
and between-observer variability. Thus,
impedance may be better than skinfold thickness
 measurements in a study of 1000 people, where body fat percentage or in BMI will result in large
variability between technicians is important. One differences in the preva
may argue that, for population studies, a slight lence of obesity.
overestimation or under
estimation of body fat is not important. However,
if, for example, obesity is defi ned as a certain
amount of body fat that is achieved at a certain
distinct level of BMI, even minor differences in

Figure 2.6 The difference in the relationship between BMI and

body fat percentage across populations is best demonstrated in this fi gure and the given biodata. Note that the two young
women are the same age and their percentage body fat as determined by a chemical four compartment model (bias free!) is
the same. The Asian woman has relatively shorter legs and a more slender body build (determined as height/(sum of knee
and wrist diameter). Relative leg length and “slenderness” are main determining factors in the BMI/percentage body fat
relationship in addition to physical activity level.

–10
)t p

10
t

gi
a

ydo
Young females (22 years)
e b

i
w

iB

ydo
b
20
%

0
t

e
Elderly females (72 years) Figure 2.7 Individual
Body Composition 29 differences in mea sured body
fat percentage compared with a
chemical four-compartment
model in 20 young and 20
elderly females using various
techniques. y-axis, BF% from
four-compart ment model
minus: , densitometry (Siri); ,
deuterium dilution; , DXA; +,
three compartment model
(Siri); , skinfold thick ness; ,
bioelectrical impedance.

In physiological studies where, for example, model. It is obvious that errors can be
energy metabolism has to be corrected for body considerable, both at a group level and espe
composition, a systematic bias in body cially at an individual level in the elderly. Many of
composition results may lead to completely wrong these errors can be explained by the violation of
conclusions. When a body composition formula or assumptions. This clearly shows that information
method sys on body composition must be used with an
tematically overestimates the FFM in obese awareness of the limitations of the techniques.
subjects, a “normal” resting metabolic rate in
obese subjects would be underestimated after 2.8 Perspectives on the future
“correction” for dif ferences in body composition
(expressed as kJ/kg FFM). This may lead to the Given the importance of body composition to
conclusion that obese subject s have lower evalu ate nutritional status and to gain information
metabolic rates, which might have caused their on certain disease processes, it can be expected
excess adipose tissue stores. This argument that more
does not even imply whether the FFM can be 30 Introduction to Human Nutrition
treated as one entity in different populations
instead of different components such as the
heart, liver, or muscles (Gallagher et al., 1998). easy-to-use methods and instrumentation will be
These con developed and become available in the future.
siderations are important for comparative studies Application and interpretation of data from these
between groups (eg, obese versus nonobese, methods should be used with an awareness of
elderly versus young or middle-aged), but also for pos
longitudi nal studies (eg, weight-loss studies). sible limitations, as mentioned above. There is a
The use of bioelectrical impedance to predict growing perception that, because of differences in
changes in FFM, TBW, or body fat during weight body build, frame size, and possibly also other
loss is questionable. The difference in body water vari ables, there are differences in the relationship
distri bution (anatomically as well as between BMI and body fat percentage among
intracellular/extracel lular) before and after weight ethnic groups. Some Aboriginal and Asian people
loss may be considerable, leading to a different have a higher fat percentage and therefore
and systematic bias of the pre diction equation. In greater risk for several chronic diseases at a
such a study it may be better to given BMI than Caucasian popu lations.
report changes in impedance values instead of Conversely, Africans often have higher bone
changes in predicted body composition. Another density than Caucasians. Much more research is
example is the very low change in impedance needed to defi ne the optimal BMI values that will
after water removal in patients with ascites. correlate with optimal health and the lowest risk of
Despite the fact that a consid disease for different populations.
erable amount of water is removed from the body,
the change in total body impedance is low as the Acknowledgment
trunk hardly contributes to total body impedance.
All methods have their limitations, doubly This chapter has been revised and updated by
indirect more than indirect methods. Figure 2.7 Paul Deurenberg based on the original chapter by
shows the individual errors in body fat percentage Paul Deurenberg and Ronenn Roubenoff. For
from den sitometry, DXA, deuterium dilution, a more infor mation on this topic visit
chemical three-compartment model, skinfold www.nutritiontexts.com
thicknesses, and bioelectrical impedance in
young and elderly women compared with body fat
percentage obtained from a four-compartment
References
Durnin JVGA, Womersley J. Body fat assessed from total Positive balance is when intake exceeds expenditure and
body density and its estimation from skinfold thickness: is associated with increases in body energy stores
measure (weight gain). During negative balance, as in periods of
starvation, body energy stores are depleted.
• Energy intake corresponds to the energy content of macro
nutrients in foods. Carbohydrate provides 16.8 kJ/g,
ments on 481 men and women aged from 17 to 72 years. protein also 16.8 kJ/g, and fat 37.8 kJ/g. In addition,
Br J Nutr 1974; 32: 77–97. alcohol provides 29.4 kJ/g.
Forbes GB. Human Body Composition. Springer, New York, • Total energy expenditure constitutes approximately two-
1987. Gallagher D, Belmonte D, Deurenberg P, Wang ZM, thirds of the energy expended by the body to maintain
Krasnow N, Pi-Sunyer FX, Heymsfi eld SB. Organ-tissue
basic physiological functions plus the thermic effect of a
mass measurement by MRI allows accurate in vivo modeling
of REE and metabolic active tissue mass. Am J Physiol 1998: meal and energy expended during physical movement.
275: E249–258. Heymsfi eld SB, Lohman TG, Wang ZW, The basic physiological functions include heart beat,
Going SB. Human Body Composition, 2nd edn. Human muscle function, and respiration (resting or basal
Kinetics, Champaign, IL, 2005. metabolic rate). The thermic effect of a meal is about 10%
Lukaski HC, Bolonchuk WW, Hall CB, Siders WA. Validity of of the caloric value of the meal needed to digest,
tet rapolar bioelectrical impedance method to assess metabolize, and store ingested macronutrients. The
human body composition. J Appl Physiol 1986; 60: 1327– energy expended during physical activity is energy
1332. expended when skeletal muscles are used for any type of
Segal KR, Van Loan M, Fitzgerald PI, Hodgdon JA, Van Itallie physical movement. In infants and children, the cost of
TB. Lean body mass estimation by bio-electrical impedance
growth is added.
analysis: a four site cross-validation study. Am J Clin Nutr
1988; 47: 7–14. • Energy requirement is the amount of food energy needed
Visser M, Heuvel van den E, Deurenberg P. Prediction to balance energy expenditure in order to maintain body
equations for the estimation of body composition in the size, body
elderly using anthropometric data. Br J Nutr 1994; 71: 823–
833.
WHO Expert Consultation. Appropriate body-mass index for
Asian populations and its implications for policy and
interven tion strategies. Lancet 2004; 363: 157–163.
3.1 Introduction
Further reading Defi nition and conceptualization of
energy balance
Siri WE. Body composition from fl uid spaces and density: The average adult human consumes close to 1
analysis of methods. In: Brozek J, Henschel A, eds.
Techniques for Measuring Body Composition. National 000 000 calories (4000 MJ) per year. Despite this
Academy of Sciences, Washington, DC, 1961: 223–244. huge energy intake, most healthy individuals are
Snijder WS, Cook MJ, Nasset ES, et al. Report of the Task able to strike a remarkable balance between how
Group on Reference Man. Pergamon Press, Oxford, 1984.
Wang ZM, Pierson RN, Heymsfi eld SB. The fi ve-level model; much energy is consumed and how much energy
a new approach to organise body composition rsearch. Am is expended, thus resulting in a state of energy
J Clin Nutr 1992; 56: 19–28. balance in the body. This accurate balance
Organisasi Kesehatan Dunia. Obesity: Preventing and
Managing the Global Epidemic. WHO, Geneva, 1998. between energy intake and energy expenditure is
an example of homeostatic control and

3 © 2009 A Astrup and A Tremblay.

Energy
Metabolism Arne
Astrup and Angelo Tremblay

Key messages
• Energy balance in the body is the balance between how
much energy is consumed and how much is expended. composition, and level of physical activity, consistent with
 long term good health. This includes the energy needs for
optimal growth and development in children, and the
needs of pregnancy
and lactation (deposition of tissue and secretion of milk). •
Body mass index (BMI) classifi es weight relative to height
squared and is the most accepted and widely used crude
index of obesity. A BMI of 18–24.9 kg/m2 is regarded as
normal for adults, between 25 and 29.9 kg/m2 as overweight,
and >30 kg/m2 as obese.
• Energy expenditure can be measured by direct methods
(calor imetry) or indirect methods, in which oxygen
consumption and carbon dioxide production are used to
calculate energy expendi ture. However, the modern gold
standard is measurement by doubly labeled water, which
is a noninvasive method used to measure total energy
expenditure over periods of 7–14 days while subjects are
living in their usual environments.
• Hunger is the physiological need to eat and results in
actions to attempt to obtain food for consumption. Appetite
is a psychologi cal desire to eat and is related to the
pleasant sensations that are often associated with food.
Thus, hunger is more of an intrinsic instinct, whereas
appetite is often a learned response.
now considered one of the major nutritional
disorders.
Components of energy balance
Energy intake
Energy intake is defi ned as the caloric or energy
content of food as provided by the major sources
of dietary energy: carbohydrate (16.8 kJ/g),
protein (16.8 kJ/g), fat (37.8 kJ/g), and alcohol
(29.4 kJ/g).
Energy storage
The energy that is consumed in the form of food
or drinks can either be stored in the body in the
form of fat (the major energy store), glycogen
(short-term energy/carbohydrate reserves), or
protein (rarely used by the body for energy except
in severe cases of starvation and other wasting
conditions, as discussed later in the chapter), or
be used by the body to fuel energy-requiring
events.

Energy expenditure
The energy that is consumed in the form of food
results in maintenance of body weight and body is required by the body for metabolic, cellular, and
energy stores. This regulation of energy balance mechanical work such as breathing, heart beat,
is achieved over the long term despite large fl and muscular work, all of which require energy
uctuations in both energy intake and energy and result in heat production. The body requires
expenditure within and between days. The energy for a variety of functions. The largest use
accuracy and precision by which the body of energy is needed to fuel the basal metabolic
maintains energy balance is high rate (BMR), which is the energy expended by the
lighted by the fact that even a small error in the body to maintain basic physiological functions
system can have detrimental consequences over (eg, heart beat, muscle con traction and function,
time. If energy intake chronically exceeds energy respiration). BMR is the minimum level of energy
expenditure by as little as 105 kJ/day, then, over expended by the body to sustain life in the awake
time, a person will become substantially obese. state. It can be measured after a 12 hour fast
The achievement of energy balance is driven by while the subject is resting physically and
the fi rst law of thermo- mentally, and maintained in a thermoneutral,
32 Introduction to Human Nutrition quiet environment. The BMR is slightly elevated
above the metabolic rate during sleep, because
energy expenditure increases above basal levels
dynamics, which states that energy can be owing to the
neither destroyed nor created. This principle
necessitates that when energy intake equals
energy expenditure, body energy stores must
remain constant. This chapter explains how the energy cost of arousal. Because of the diffi culty
body is able to achieve this state of energy in achieving BMR under most measurement
balance through control of energy intake and situations, resting metabolic rate (RMR) is
energy expenditure. In addition, the various ways frequently measured using the same
that body energy stores can be measured and measurement conditions stated for BMR. Thus,
some exam ples of conditions in which energy the major difference between BMR and RMR is
balance may be disrupted are summarized. the slightly higher energy expended during RMR
Particular emphasis is placed on obesity, which is (~ 3%) owing to less subject arousal and non
the end-result of a positive energy balance and is fasting conditions. Because of this small
 difference, the terms basal and resting metabolic occurs in humans, eg, during the initial months
rate are often used interchangeably. RMR occurs of life and during fever and other pathological
in a continual process throughout the 24 hours of conditions, but also as a contributor to daily
a day and remains relatively constant within energy expenditure.
individuals over time. In the average adult human, ● Thermogenesis is increased by a number of
RMR is approximately 4.2 kJ/min. Thus, basal or agents in the environment, including in foods
resting metabolic rate is the largest component of and bever ages. Nicotine in tobacco is the most
energy expenditure and makes up about two- important one, and heavy smokers may have a
thirds of total energy expenditure. 10% higher energy expenditure than
In addition to RMR, there is an increase in nonsmokers of similar body size and
energy expenditure in response to food intake. composition and physical activity. Caffeine and
This increase in metabolic rate after food derivatives in coffee, tea, and choco late,
consumption is often referred to as the thermic capsaicin in hot chilies, and other substances in
effect of a meal (or meal foods and drinks may possess minor thermo
induced thermogenesis) and is mainly the energy genic effects that affect energy expenditure.
that is expended to digest, metabolize, convert,
and store ingested macronutrients, named Energy balance
obligatory thermo genesis. The measured thermic Energy balance occurs when the energy content
effect of a meal is usually higher than the of food is matched by the total amount of energy
theoretical cost owing to a facultative component that is expended by the body. An example of
caused by an activation of the sympathoadrenal energy balance would be the scenario cited at the
system, which increases energy expenditure outset of this chapter in which, over a year, the
through peripheral β-adrenoceptors. The energy average adult con
cost associated with meal ingestion is pri sumes and expends 1 000 000 calories, resulting
marily infl uenced by the composition of the food in no net change in the energy content of the
that is consumed, and also is relatively stable body. When energy intake exceeds energy
within indi viduals over time. The thermic effect of expenditure, a state of positive energy balance
a meal usually constitutes approximately 10% of occurs. Thus, positive energy balance occurs
the caloric content of the meal that is consumed. when excessive overfeeding relative to energy
The third source of energy expenditure in the needs occurs, and the body increases its overall
body is the increase in metabolic rate that occurs energy stores. Examples of positive energy
during physical activity, which includes exercise balance include periods around major festivals
as well as all forms of physical activity. Thus, when overeat ing and inactivity generally prevail,
physical activity energy expenditure (or the and during preg
thermic effect of exercise) is the term fre nancy and lactation when the body purposefully
quently used to describe the increase in increases its stores of energy. When energy
metabolic rate that is caused by use of skeletal intake is lower than energy expenditure, a state of
muscles for any type of physical movement. negative energy balance occurs, for example
Physical activity energy expen diture is the most during periods of starvation. In this regard,
variable component of daily energy expenditure evidence suggests that, under conditions of
and can vary greatly within and between substantial energy imbalance, be it positive or
individuals owing to the volitional and variable negative, energy expenditure may reach a level
nature of physical activity patterns. that is beyond what could be predicted by body
In addition to the three major components of weight changes. This so-called “adaptive
energy expenditure, there may be a requirement thermogen esis” might contribute to the
for energy for three other minor needs. occurrence of resistance to lose fat in the context
of obesity treatment or the achievement of a new
body weight plateau following overfeeding. It is
important to note that energy balance can occur
● The energy cost of growth occurs in growing indi
regardless of the levels of energy intake and
viduals, but is negligible except within the fi rst expenditure; thus, energy balance can
few months of life. Energy Metabolism 33
● Adaptive thermogenesis is heat production
during exposure to reduced temperatures, and
occur in very inactive individuals as well as in regulated by a number of factors involving
highly active individuals provided that adequate complex inter actions among various hormones,
energy sources are available. It is also important neuroendocrine
to think of energy balance in terms of the major 34 Introduction to Human Nutrition
sources of energy, ie, carbohydrate, protein, and
fat. For example, carbohydrate balance occurs
when the body balances the amount of factors, the central nervous system, and organ
carbohydrate ingested with that expended for systems (eg, brain and liver), and environmental
energy. and exter nal factors.
Appetite is usually defi ned as a psychological
desire to eat and is related to the pleasant
3.2 Energy intake sensations that are often associated with specifi c
foods. Scientifi cally, appetite is used as a general
Sources of dietary energy term of overall sensations related to food intake.
As mentioned above, the sources of energy in the Hunger is usually defi ned as the subjective
food we eat include the major macronutrients: feeling that determines when food consumption is
protein, car bohydrate, and fat, as well as alcohol. initiated and can be described as a nagging,
Carbohydrate and protein provide 16.8 kJ of irritating feeling that signifi es food deprivation to
energy for each gram; alcohol provides 29.4 kJ/g, a degree that the next eating episode should take
whereas fat is the most energy dense, providing place.
37.8 kJ/g. Note that 4.2 kJ is defi ned as the Satiety is considered as the state of inhibition
amount of heat that is required to raise the over eating that leads to the termination of a
temperature of 1 liter of water by 1°C. The energy meal, and is related to the time interval until the
content of food can be measured by bomb calori next eating episode. Thus, hunger and satiety are
metry, which involves combusting a known weight more intrinsic instincts, whereas appetite is often
of food inside a sealed chamber and measuring a learned response.
the amount of heat that is released during this The internal factors that regulate the overall
process. Thus, 1 g of pure fat would release 37.8 feeling of hunger and satiety include the central
kJ during its complete combustion, whereas 1 g nervous system (primarily the hypothalamus and
of pure carbohy the vagus nerve), the major digestive organs such
drate would release 16.8 kJ. Thus, if the gram as the stomach and liver, and various hormones.
quanti ties of any type of food are known, the In addition, envi
energy content can easily be calculated. For ronmental factors (eg, meal pattern and composi
example, if a protein-rich nutrition snack contains tion, food availability, smell and sight of foods,
21 g of carbohydrate, 6 g of fat, and 14 g of climate), emotional factors (eg, stress), and some
protein, then the total energy content is (21 ⋅ 16.8) diseased states (eg, anorexia, trauma, infection)
+ (6 ⋅ 37.8) + (14 ⋅ 16.8) = 814.8 kJ. The may infl uence the feelings of both hunger and
macronutrient composition of food is typically appetite. The factors that infl uence appetite
assessed in the percentage contribution of each include factors external to the individual (eg,
mac climate, weather), specifi c appetite cravings,
ronutrient to the total number of calories. If a food specifi c learned dislikes or avoidance (eg,
has a carbohydrate content of 21 g, which is alcohol), intrinsic properties of food (eg, taste,
352.8 kJ, and the total energy content is 820 kJ palatability, texture), cultural practices or
the proportion of energy derived from preferences, specifi c effects of some drugs and
carbohydrate is 43%; the fat content is 6 g, or dis
226.8 kJ, equivalent to 28% of the energy; and eases, and metabolic factors such as hormones
the protein contributes 14 g, 235.2 kJ and 29% of and neurotransmitters. Some of these factors are
the energy. described in further detail below.
The classic way to describe the complex
Regulation of food intake appetite regulating system is the satiety cascade
put forth by John Blundell. The satiety cascade
Appetite, hunger, and satiety describes four dis tinctly different but overlapping
The quality and quantity of food that is consumed categories of mecha nisms involved in acute
are closely regulated by the body. Food intake is within-meal feeling of satiety (referred to as
satiation) and the inbetween-meal satiety (Figure
3.1).
Factors infl uencing food intake
Digestive factors
Several factors in the digestive system exert a
short term infl uence over food intake. The
presence of food
Sensory
Cognitive Postabsorptive Postingestive
system, resulting in regula
tion of energy balance. In addition, other
gastrointes tinal hormones, such as glucagon-like
peptide-1 and -2 (GLPs), CCK, and glucose-
dependent insulinotro pic polypeptide (GIP) are
likely to play a role in the mediation of gut events
and brain perception of hunger and satiety.
Central nervous system factors
The main contributory factor regulating food
intake in the central nervous system is the
hypothalamus. The hypothalamus is linked to
specifi c parts of the

Early Late
Food brain that are known to modify feeding behavior,
specifi cally the paraventricular nuclei and the
Satiation Satiety nigro striatal tract. These areas of the brain
respond to various neurotransmitters as well as
Figure 3.1 The satiety cascade by John Blundell (1987). The
four categories of mechanisms are sensory, cognitive,
sympathetic nervous system activity. In general,
postingestive, and postabsorptive. The sensoric phase food intake will decrease as sympathetic nervous
includes stimuli mediated via sensory fi bers in the cranial system activity increases, and vice versa.
nerves and relates to the palatability of the ingested meal
including smell, taste, temperature, and texture. The cognitive
Circulating factors
phase of the satiety cascade represents learned, known, and
assumed properties of the ingested food. In the postingestive After consumption of a meal, food is broken down
phase the gastrointestinal tract is involved in a number of into its basic components (ie, carbohydrate is
satiety signals both via chemo- and mechanoreceptors and broken down to glucose, protein to amino acids,
appetite-regulating peptides from the gut and pancreas either and fats or triglycerides to glycerol and fatty
entering circulation and acting directly in the CNS or signaling
acids) and the cir
via the vagus nerve. Important satiety signals in the
postabsorptive phase include circulating nutrients, signals culating levels of some of these breakdown
derived from differences in energy turnover, substrate oxida products increase in the blood. Consequently,
tion, and neurohormonal factors. Reprinted from Blundell et glucose, amino acids, glycerol, and fatty acids are
al. further metabo lized, primarily in the liver, or used
(1987), copyright 1987 with permission of Elsevier.
for immediate energy (eg, in muscle or brain).
There is evidence to suggest that this resultant
metabolism, especially in the liver, may in turn
regulate food intake. After meal consumption, the
and drink in the stomach and intestine and the circulating levels of nutrients fall (within minutes
resul tant pressure that they exert may regulate for glucose, several hours for triglyc
food intake. This effect is known as erides) and the feelings of hunger return. The link
gastrointestinal distension. In addition, the from nutrient metabolism to central control of food
stomach produces a hormone called intake occurs through signals from the liver to the
cholecystokinin (CCK) in response to food intake, brain via the vagus nerve. Thus, circulating
which may, in turn, regulate food intake. factors provide a link between the digestive
Furthermore, when subjects have fat or system and the central nervous system, which
carbohydrate infused directly into the small provides another system for regulating food
intestine, they report feelings of satiety. This intake.
suggests that factors in the intestine regulate food
intake. Indeed, receptors in the intes Signals from the periphery
tine have been identifi ed that recognize the Leptin is a hormone that is produced by fat cells
presence of specifi c macronutrients; these and communicates with the central nervous
receptors are linked to the brain and therefore system through leptin receptors in the
can communicate directly with the central nervous hypothalamus. Reduced production of leptin, or
lack of sensitivity of the hypo cultural infl uences in the envi ronment, such as
thalamus to leptin, may regulate food intake and time of day, social factors, peer infl uence, and
play a key role in the etiology of rare forms of cultural preferences, can also play a role in infl
obesity in humans. Leptin and the other uencing food intake.
peripheral hormones with a central effect on
appetite are divided into two broad categories: (1)
the so-called adiposity signals, which are tonically 3.3 Energy expenditure
active providing information on body fat stores to
Concept of energy expenditure
the CNS and (2) the satiety signals which are
released in response to food intake and are The process of energy expenditure and the
thought to be involved in short-term regulation of oxidation or combustion of food for energy in the
energy intake. Currently known adiposity signals body is anal ogous to a woodstove that burns
are insulin, leptin, and adiponectin, which are wood to release heat in a controlled fashion. In
considered as long-acting signals reducing this analogy, large chunks of wood are fed to the
energy intake. Among the satiety signals are the stove and the wood is gradually combusted in the
hunger hormone ghrelin, which is secreted in the presence of oxygen to release carbon dioxide,
stomach, and the short-acting water vapor, and heat. Similarly, in the body, the
Energy Metabolism 35 food consumed is oxidized or combusted in the
presence of oxygen to release carbon dioxide,
water, and heat. When ingested food is used for
gut- and pancreas-derived satiety signals CCK, energy,
peptide YY (PYY), GLP-1, oxyntomodulin (OXM), 36 Introduction to Human Nutrition
and pan creatic polypeptide (PP). Many of the
peripheral satiety signals have receptors in the
arcuate nucleus (ARC) of the hypothalamus, however, the release and transfer of energy occur
which plays an impor tant role in appetite through a series of tightly regulated metabolic
regulation. The ARC contains neuropeptide Y path ways in which the potential energy from food
(NPY)- and agouti-related peptide (AgRP)- is released slowly and gradually over time. This
expressing neurons acting to stimulate food process ensures that the body is provided with a
intake along with the adjacent pro- gradual and constant energy store, rather than
opiomelanocortin (POMC) and cocaine- and relying on a sudden release of energy from an
amphetamine-regulated transcript (CART)- immediate combustion of ingested food. As a
expressing neurons which inhibit feeding. Besides simple example of how the body uses food for
the ARC, the nucleus of the solitary tract (NTS) energy, consider the combustion of a simple
and the area postrema (AP) receive appe tite- glucose molecule:
regulating inputs from vagal afferents and circu C6H12O6 + 6O2 → 6H2O + 6CO2 + Heat
lating factors and are connected to the
hypothalamic nuclei controlling food intake. Similar chemical reactions can be described for
the combustion of other sources of energy, such
as fat and other types of carbohydrates. These
External factors types of reac tion occur continuously in the body
Various nonphysiological or external factors are and constitute energy expenditure. As discussed
also known to modify food intake, and these previously, the three major sources of energy
effects may be mediated through the intrinsic expenditure in the body are to fuel RMR, the
factors described above. Psychological factors thermic effect of meals, and physical activity. As
such as depression may lead to either increased discussed in more detail below, energy
or decreased food intake, or changes in the expenditure can be measured by assessment of
consumption of specifi c types of foods. total heat production in the body (direct
Environmental factors are also important, the calorimetry) or by assessment of oxygen
most obvious being food availability. Even when consumption and carbon dioxide production
food is available, some of the specifi c properties (indirect calorimetry).
of foods make them more or less appealing,
thereby modifying food intake. Important physical Historical aspects of energy expenditure The
characteristics of food include taste, texture, burning or combustion of food in the body was
color, temperature, and presentation. Other originally described in the classic experiments of
Lavoisier, who worked in France in the late eight
eenth century. Lavoisier discovered that a candle
would burn only in the presence of oxygen. In
addi tion, he was the fi rst to describe how living
organisms produced heat in a similar way, as
they required oxygen for life and combusted food
as they released heat. His experiments were the
fi rst to document the heat production of living
organisms. Working before the invention of
electricity, he built the fi rst calorim eter in which a
small animal was placed in a sealed chamber.
Lavoisier packed ice into a sealed pocket around
the chamber (he could only perform these studies
in the winter when ice was collected from the
ground), and then placed the chamber and ice
layer inside an insulated chamber. Lavoisier then
collected and measured the volume of melting
water. Since the ice layer was insulated from the
outside world, the only way that the ice could melt
was by the increase in heat produced by the living
animal. Lavoisier therefore measured the volume
of melted ice water, and, by so doing, was able to
calculate accurately the
amount of heat that had to be produced by the
animal to melt the measured amount of ice.
typically measured by monitoring the changes in
metabolic rate by indirect calorimetry for 3–6
hours following consumption of a test meal of
known caloric content. The energy expended in
physical activity can be measured under
laboratory conditions, also using indirect
calorimetry during standard activities. In addition,
free-living physical activity-related energy
expenditure over extended periods of up to 2
weeks can be measured by the combination of
doubly labeled water (DLW) to measure total
energy expenditure (see below), and indirect
calorimetry to measure resting energy expen
diture and the thermic effect of a meal. Indirect
calo rimetry has an added advantage in that the
ratio of carbon dioxide production to oxygen
consumption (the respiratory quotient, or RQ) is
indicative of the type of substrate (ie, fat versus
carbohydrate) being oxidized, for example
carbohydrate oxidation has a RQ of 1.0 and fat
oxidation has a RQ close to 0.7.
Energy expenditure can be assessed from
indirect calorimetry in a simple, less accurate way
by ignoring the contribution of protein oxidation or
by collecting urine during the measurement to
analyze the excreted nitrogen. The latter
approach is preferable because it

Measurement of energy expenditure Lavoisier's

device was the fi rst calorimeter that was used to gives a more accurate estimate of energy
measure heat production. This approach is expenditure and RQ.
termed direct calorimetry because heat
production is measured directly. Direct Step 1
calorimeters have been designed for measuring First, the contribution of protein oxidation to
heat production in humans, but this approach is
technically demanding, especially in human oxygen consumption (V.O2) and carbon dioxide
studies, and is now infrequently used. Indirect
calorimetry measures energy production via production
respiratory gas analysis. This approach is based
(V.CO2) is estimated based on the knowledge that
on oxygen consumption and carbon dioxide the nitrogen content of protein is 1/6.25:
production that occurs during the combustion (or
V.O2(prot) = n ⋅ 6.25 ⋅ 0.97
oxidation) of protein, carbohydrate, fat, and
alcohol, as shown in the example of glucose V.CO2(prot) = n ⋅ 6.25 ⋅ 0.77
combustion. Respiratory gas analysis can easily where V is volume, 0.97 and 0.77 are liters of O
2
be achieved in humans either over short con sumed and CO2 produced by the biological
measurement periods at rest or during exercise oxidation of 1 g of protein, respectively, and prot
using a face mask, mouthpiece, or canopy is protein.
system for gas collection, and over longer periods
of 24 hours (and longer) with subjects living in a Step 2
metabolic chamber. BMR is typically measured by
indirect calo rimetry under fasted conditions while Next, nonprotein V.O2 (V.O2(nonprot)) and nonprotein
subjects lie quietly at rest in the early morning for
30–40 min. The thermic effect of a meal is V.CO2 (V.CO2(nonprot)) are calculated:

V.O2(nonprot) = V.O2 − V.O2(prot)
V.CO2(nonprot) = V.CO2 − V.CO2(prot)
V.O2(nonprot) = C ⋅ 0.828 + F ⋅ 2.03
V.CO2(nonprot) = C ⋅ 0.828 + F ⋅ 1.43
where C and F are grams of oxidized
carbohydrate and fat, respectively, and can be
found by solving the two equations with two
unknowns; O2 dan CO2 pro duced by the
combustion of 1 g of carbohydrate is 0.828 liters,
whereas the combustion of 1 g triglycer ide
consumes 2.03 liters O2 and produces 1.43 liters
CO2. The protein oxidation (P) is n ⋅ 6.25 g.
Step 3
The RQ is defi ned as:
V.CO2/V.O2
Nonprotein RQ (RQ(nonprot)) is calculated by the
Energy Metabolism 37
The caloric equivalent for O2 is similar to the
three main substrates: 21 kJ/l O2 for
carbohydrate, 19 kJ/l O2 for fat, and 17.8 kJ/l O2
for protein (which con tributes only modestly to
energy expenditure). Energy expenditure can
therefore be calculated with reason able accuracy
by the equation:
Energy expenditure (kJ/min) = 20 kJ/l ⋅ V.O2 (l/min)
With pure fat oxidation the RQ is 0.707, with pure
carbohydrate oxidation it is 1.0, and with pure
protein oxidation it is approximately 0.8.
Step 5
Oxidation of protein (P), carbohydrate (C), and fat
(F) can be calculated by the following equations,
where n is the unit g/min:
P (g/min) = 6.25 ⋅ n
C (g/min) = 4.55 ⋅ V.CO2 − 3.21 ⋅ V.O2 − 2.87 F
(g/min) = 1.67 ⋅ V.O2 − 1.67 ⋅ V.CO2 − 1.92 ⋅ n

where 17, 17.5, and 38.9 are the heat produced 3.4 Factors that infl uence
(kJ) by the combustion of 1 g of protein, glycogen, energy expenditure
and triglyceride, respectively.
The equations are produced by the insertion of Resting metabolic rate
the heat equivalent for carbohydrate and fat, and Each of the components of energy expenditure is
are valid even though there is a quantitative determined by various factors. RMR is highly
conversion of carbohydrate to lipid (de novo variable between individuals (±25%), but is very
lipogenesis) or glyconeogenesis. consistent
equation: Step 4 the body, the main source of
RQ(nonprot) = V.CO2(nonprot)/V.O2(nonprot)
individual variability in RMR is an
within individuals (<5%). Since individual's amount of organ and
RMR occurs pre dominantly in muscle
muscle and the major organs of
Next, energy expenditure can be calculated: larger the wood
stove (or FFM), the larger the amount of heat pro
Energy expenditure (kJ/min) duction (or the larger the RMR). Since FFM is a
= [19.63 + 4.59 (RQ(nonprot) − het erogeneous mixture of all nonfat body
components, the metabolic rate associated with
0.707] ⋅ V.O2(nonprot) + 18.78 ⋅
each kilogram of
38 Introduction to Human Nutrition
V.O2(nonprot)
or FFM is dependent on the quality of the FFM, in
Energy expenditure (kJ/min) = terms of hydration and relative contribution of the
17 ⋅ P + 17.5 ⋅ C + 38.9 ⋅ F different organs that make up the FFM. For
mass. Thus, fat-free mass (FFM; the total mass of example, skeletal muscle constitutes
the body that is not fat, ie, predominantly organs approximately 43% of total mass in an adult, but
and muscle) explains 60–80% of the variation in contributes only 22–36% of the RMR, whereas
RMR between individuals. This concept can be the brain, which constitutes approximately only
explained using the woodstove analogy; the 2% of mass, contributes 20–24% of the RMR. In
addition, the metabolic cost of each kilogram of
FFM decreases with developmental progression,
probably owing to developmental increases in the
muscle mass to organ mass ratio within FFM.
Thus, the relation ship between RMR and FFM is
not linear across all ages and is estimated to be
331.8 kJ/kg between the ages of 0 and 2.5 years,
151.2 kJ/kg in children aged 4–7 years, 88.2
kJ/kg during adolescence, and 151.2 kJ/ kg in
adulthood.
Although fat mass is generally thought to be meta
bolically inert, it signifi cantly contributes to varia
tions in RMR. This is likely explained, at least in
part, by neurobiological effects (eg, changes in
sympathetic nervous system activity) resulting
from variations in fat mass which affect the
metabolism of other tissues. RMR is also infl
uenced by fat mass, even though fat mass is
generally thought to be meta bolically inert. Fat
mass contributes in the order of 42.0–54.6 kJ/kg
to RMR. This difference is indepen dent of the
gender difference in FFM; in other words, if one
studied a group of males and females of identical
FFM and similar age, RMR would be higher in
males than in females by around 210.0 kJ/ day.
This gender difference is consistent across the
lifespan, and the source of the difference is not
well understood (Table 3.1). More active people
tend to have a higher RMR than inactive
individuals. This difference may be explained in
part by the resid ual effects of chronic exercise on
metabolic rate. In other words, RMR appears to
be elevated because of the long-lasting effects of
the thermic effect of exer cise. However, other
factors are also involved, since the higher RMR in
more active individuals persists long after the last
bout of exercise has been com pleted.
Collectively, FFM, fat mass, age, gender, and
physical activity explain 80–90% of the variance
in RMR. In addition, a portion of the unique
variance in RMR across individuals has been
ascribed to genetic factors, although the specifi c
source of this genetic variation has not yet been
identifi ed. Other factors that have been shown to
infl uence metabolic rate include thyroid
hormones (higher levels increase
Table 3.1 Variation in total energy expenditure (TEE) as a
function of resting metabolic rate (RMR) among various
populations
Study group Average TEE/RMR (range)
5-year-old children in Arizona, USA 1.37 (1.15–1.70)
Obese women in the UK 1.39 (1.20–1.77) Elderly
women in Vermont, USA 1.42 (1.25–1.82) 5-year-old
children in Vermont, USA 1.44 (1.11–1.77) Elderly
men in Vermont, USA 1.50 (1.30–2.11) Obese Pima
Indians 1.56 (1.03–1.99) Adolescents in the UK 1.56
Dutch adults 1.64
Obese women in New York, USA 1.68
Young men in Boston, USA 1.70 (1.38–2.32) Obese
women in New York, USA 1.73
Elderly men in Boston, USA 1.74
Young men in the UK 1.88 (1.44–2.57) Young men in
Boston, USA 1.98 (1.57–2.60) Mount Everest
climbers 2.0
Tour de France cyclists 5.3
Burns patients 1.3
Range of TEE/RMR is given in parentheses for studies in
which the individual data were reported.
metabolic rate) and sympathetic nervous system
activity.
Several prediction equations have been
developed to estimate RMR from other simple
measures. These equations are often useful for
making estimates in clinical situations when
measurement of RMR cannot be achieved, or for
estimating energy needs for other individuals. The
classic equations of Harris and Benedict are
frequently used for this purpose. These equations
were developed from limited measures per
formed in the early 1900s, and predict RMR from
age, height, and weight, and may be of limited
accuracy. More recent equations have been
developed in larger groups of subjects and can
predict RMR from body weight (Table 3.2). These
new equations have been shown to be more
accurate.
Thermic effect of feeding
The thermic effect of meal ingestion is primarily
infl uenced by the quantity and macronutrient
quality of the ingested calories. The thermic effect
of food has also been termed meal-induced
thermogenesis, or the specifi c dynamic action of
food. The increase in meta
bolic rate that occurs after meal ingestion occurs
over an extended period of at least 5 hours; the
cumulative energy cost is equivalent to around
10% of the energy ingested. In other words, if one
consumed a mixed meal of 2.1 MJ, the body
would require 210.0 kJ to
Table 3.2 Simple equations for estimating resting metabolic
rate (RMR) from body weight according to gender and age
RMR (kJ/day)
Age (years) Equation for males Equation for females
0–3 (60.9 ⋅ wt) − 54 (61.0 ⋅ wt) − 51 3–10 (22.7 ⋅ wt) + 495
(22.5 ⋅ wt) + 499 10–18 (17.5 ⋅ wt) + 651 (12.2 ⋅ wt) + 746
18–30 (15.3 ⋅ wt) + 679 (14.7 ⋅ wt) + 496 30–60 (11.6 ⋅ wt)
+ 879 (8.7 ⋅ wt) + 829 >60 (13.5 ⋅ wt) + 487 (10.5 ⋅ wt) +
596
wt, body weight (kg).
digest, process, and metabolize the contents of
the meal. The thermic effect of feeding is higher
for protein and carbohydrate than for fat. This is
because, for fat, the process of energy storage is
very effi cient, whereas, for carbohydrate and
protein, additional energy is required for
metabolic conversion to the appropriate storage
form (ie, excess glucose con verted to glycogen
for storage, and excess amino acids from protein
converted to fat for storage). In addition to the
obligatory energetic cost of processing and
storage of nutrients, a more variable facultative
ther
mogenic component has been described. This
com ponent is mainly pertinent to carbohydrates,
which through increased insulin secretion
produce a dipha sic activation of the
sympathoadrenal system. The initial phase is an
insulin-mediated increase in sym pathetic activity,
which produces a β-adrenoceptor mediated
increase in energy expenditure. The second and
later phase occurs when a counter-regulatory
increase in plasma epinephrine is elicited by the
falling blood glucose. This increase in epinephrine
has a similar slight stimulatory effect on energy
expenditure. As a result of the mediation by β
adrenoceptors the thermic effect of carbohydrate-
rich meals can be slightly reduced by
pharmacological β adrenoceptor antagonists.
Energy expenditure related to physical
activity
Physical activity energy expenditure
encompasses all types of activity, including sports
and leisure, work related activities, general
activities of daily living, and fi dgeting. The
metabolic rate of physical activity is determined
by the amount or duration of activity (ie, time), the
type of physical activity (eg, walking,
Energy Metabolism 39
40 Introduction to Human Nutrition
Table 3.3 Examples of metabolic equivalent
(MET) values for various physical activities
environment. The technique was fi rst introduced
Activity MET
Basketball 8.0
Chopping wood 6.0
Cleaning house 2.0–4.0
Cycling for pleasure 8.0
Gardening 5.0
Kayaking 5.0
Mowing lawn (power mower) 4.5
Painting house 4.0–5.0
Playing musical instrument 2.0–4.0
Running slowly (8–11 km/h) 8.0–10.0
Running quickly (14–16 km/h) 16.0–18.0
Soccer 7.0–10.0
Strength training 6.0
Stretching 4.0
Tennis 6.0–8.0
Skiing 7.0–14.0
Swimming laps 6.0–12.0
Walking 3.0–5.0
Water skiing 6.0
running, typing), and the intensity at which the par
ticular activity is performed. The metabolic cost of
physical activities is frequently expressed as
metabolic equivalents (METs), which represent
multiples of RMR. Thus, by defi nition, sitting
quietly after a 12 hour fast is equivalent to 1 MET.
Table 3.3 provides MET values for other typical
physical activities. The cumulative total daily
energy cost of physical activity is highly variable
both within and between individuals. Therefore,
physical activity provides the greatest source of
plasticity or fl exibility in the energy expenditure
system, and is the component through which
large changes in energy expenditure can be
achieved.
Total energy expenditure: measurement
by doubly labeled water
The integrated sum of all components of energy
expenditure is termed total energy expenditure.
Until recently, there was no good way to measure
total energy expenditure in humans living under
their habitual conditions. Total energy expenditure
can be measured over 24 hours or longer in a
metabolic chamber, but this environment is artifi
cial and is not representative of the normal daily
pattern of physical activity. The DLW technique
can be used to obtain an integrated measure of
all components of daily energy expenditure over
extended periods, typically 7–14 days, while
subjects are living in their usual
in the 1950s as an isotopic technique for
measuring the carbon dioxide production rate in
small animals. Unfortunately, it was not possible
to apply the tech
nique to humans because the dose required was
cost prohibitive given the relatively poor sensitivity
of the required instrumentation at that time. It was
not for another 20 years that the inventors of this
technique described the feasibility of applying the
technique to measure free-living energy
expenditure in humans, and 10 years later this
concept became a reality.
The DLW method requires a person to ingest
small amounts of “heavy” water that is isotopically
labeled with deuterium and oxygen-18 (2H2O and
H218O). These forms of water are naturally
occurring, stable (nonradioactive) isotopes of
water that differ from the most abundant form of
water. In deuterium
labeled water, the hydrogen is replaced with
deute rium, which is an identical form of water
except that deuterium has an extra neutron in its
nucleus com pared with hydrogen, and is thus a
heavier form of water; similarly, 18O-labeled water
contains oxygen with an additional two extra
neutrons. Thus, these stable isotopes act as
molecular tags so that water can be tracked in the
body. After a loading dose, deute rium-labeled
water is washed out of the body as a function of
body water turnover; 18O is also lost as a function
of water turnover, but is lost via carbon dioxide
production as well. Therefore, using a number of
assumptions, the rate of carbon dioxide produc
tion and energy expenditure can be assessed
based on the different rates of loss of these
isotopes from the body.
The major advantages of the DLW method are
that the methodology is truly noninvasive and
nonobtru sive (subjects are entirely unaware that
energy expen diture is being measured), and
measurement is performed under free-living
conditions over extended periods (7–14 days).
Moreover, when used in combi nation with
indirect calorimetry for assessment of resting
metabolic rate, physical activity-related energy
expenditure can be assessed by the difference
(ie, total energy expenditure minus resting
metabolic rate, minus the thermic effect of meals
= physical activity energy expenditure). The
additional power of assessing total energy
expenditure with the DLW method is that this
approach can provide a measure of total energy
intake in subjects who are in energy balance. This
is because, by defi nition, in a state of
energy balance, total energy intake must be
equiva lent to total energy expenditure. This
aspect of the technique has been used as a tool
to validate energy intakes using other methods
such as food records and dietary recall. For
example, it has been known for some time that
obese subjects report a lower than expected
value for energy intake. At one time it was
thought that this was due to low energy
requirements in the obese due to low energy
expenditure and reduced physical activity.
However, using DLW, it has now been
established that obese subjects system
atically underreport their actual energy intake by
30– 40% and actually have a normal energy
expenditure, relative to their larger body size.
The major disadvantages of the technique are
the periodic nonavailability and expense of the
18
O isotope (around —500–600 for a 70 kg adult),
the need for and reliance on expensive
equipment for analysis of samples, and that the
technique is not well suited to large-scale
epidemiological studies. Furthermore, although
the technique can be used to obtain esti
mates of physical activity energy expenditure, it
does not provide any information on physical
activity pat terns (ie, type, duration, and intensity
of physical activity periods during the day).
The DLW technique has been validated in
humans in several laboratories around the world
by compari son with indirect calorimetry in adults
and infants. These studies generally show the
technique to be accurate to within 5–10%, relative
to data derived by indirect calorimetry for subjects
living in metabolic chambers. The theoretical
precision of the DLW tech nique is 3–5%.
However, the experimental variability is ±12%
under free-living conditions, owing to fl uc tuations
in physical activity levels, and ±8% under more
controlled sedentary living conditions. The good
accuracy and reasonable precision of the tech
nique therefore allow the DLW method to be used
as a “gold standard” measure of free-living energy
expenditure in humans against which other
methods can be compared.
3.5 Energy requirements
How much energy do we need to sustain life and
maintain our body energy stores? Why do some
people require more energy and others less? In
other words, what are the energy requirements of
different types of people? Based on our earlier
defi nition of
energy balance, the energy needs or energy
require ments of the body to maintain energy
balance must be equal to total daily energy
expenditure. Total daily energy expenditure is the
sum of the individual com ponents of energy
expenditure as discussed previ ously, and
represents the total energy requirements of an
individual that are required to maintain energy
balance. Until recently, there was no accurate
way to measure total energy expenditure or
energy needs of humans. The DLW technique
has provided a truly noninvasive means to
measure accurately total daily energy
expenditure, and thus energy needs, in free living
humans. Before DLW, energy requirements were
usually assessed by measurement or prediction
of RMR, the largest component of energy require
ments. However, since the relationship between
RMR and total energy expenditure is highly
variable because of differences in physical
activity, the estimation of energy needs from
knowledge of RMR is not that accurate and
requires a crude estimate of physical activity
level. Nevertheless, reasonable estimates can be
made to estimate daily energy budgets for indi
viduals (Table 3.4).
Following the validation of DLW in humans, this
technique has been applied to many different
popula tions. Total energy expenditure is often
compared across groups or individuals using the
ratio of one's total energy expenditure to RMR, or
physical activity level (PAL). Thus, for example, if
the total energy expenditure was 12.6 MJ/day and
the RMR was 6.3 MJ/day, the PAL factor would
be 2.0. This value indicates that total energy
expenditure is twice the
Energy Metabolism 41
RMR. The PAL factor has been assessed in a
variety of types of individual. A low PAL indicates
a sedentary lifestyle, whereas a high PAL
represents a highly active lifestyle. The highest
recorded sustained PAL in humans was recorded
in cyclists participating in the Tour de France road
race. These elite athletes could sustain a daily
energy expenditure that was up to fi ve times their
RMR over extended periods. Smaller animals,
such as migrating birds, have a much higher
ceiling for achieving higher rates of total energy
expen diture, which can reach up to 20 times their
RMR.
Factors such as body weight, FFM, and RMR
account for 40–60% of the variation in total
energy expenditure. Total energy expenditure is
similar between lean and obese individuals after
taking into account differences in FFM. Thus,
fatness has small, but important, additional
effects on total energy expenditure, partly through
RMR, as discussed previ
ously, but also by increasing the energetic cost of
any physical activity.
With regard to age, some studies suggest that
only a limited change in total energy expenditure
(relative to RMR) occurs from childhood to
adulthood, but that a decline occurs in the elderly.
Recent data also suggest a gender-related
difference in total energy expenditure, in addition
to that previously described for RMR. In a meta-
analysis that examined data from a variety of
published studies, absolute total energy
expenditure was signifi cantly higher in males
than in females by 3.1 MJ/day (10.2 ± 2.1 MJ/day
in females, 13.3 ± 3.1 MJ/day in males), and
nonresting energy expenditure remained higher in
men by 1.1 MJ/day.

Table 3.4 Typical daily energy budgets for a sedentary and a physically active
individual of identical occupation, body weight, and resting metabolic rate of 6.0
MJ/day (4.2 kJ/min)

Minutes per day MJ per day

Activity Activity index Sedentary Active Sedentary Active

Sleep 1.0 480 480 2.0 2.0 Daily needs 1.06 120 120 5.3 5.3 Occupational 1.5 480
480 3.0 3.0 Passive recreation 2.0 360 300 3.0 2.5 Exercise 12.0 0 60 0 3.0 Total
1440 1440 8.6 11.1 PAL = 1.4 PAL = 1.8

Thus, the sedentary individual would need to perform 60 min of vigorous activity
each day at an intensity of 12.0 to increase the physical activity level (PAL) from a
sedentary 1.4 to an active and healthy 1.8.
42 Introduction to Human Nutrition Individuals who have sedentary occupations
and do not participate frequently in leisure
pursuits that require physical activity probably year old children total energy expenditure by
have a PAL factor in the region of 1.4. Those who DLW aver
have occupations requiring light activity and
participate in light physi
cal activities in leisure time probably have a PAL
around 1.6 (this is a typical value for sedentary ages 5.1 MJ/day, while the currently
people living in an urban environment). recommended intake for these children is 6.2
Individuals who have physically active MJ/day. Thus, newer estimates of the energy
occupations and lifestyles prob requirements of infants are needed based on
ably have a PAL greater than 1.75. It has been assessment of total energy expendi
sug gested that the optimal PAL that protects ture data.
against the development of obesity is around 1.8 Several laboratories have reported
or higher. Increasing one's physical activity index measurements of total energy expenditure in
from 1.6 to 1.8 requires 30 min of daily vigorous young, healthy, free living children around the
activity, or 60 min of light activity (Table 3.4). world. Despite marked dif ferences in
geographical locations, the data are similar,
although environmental factors such as season
3.6 Energy balance in various and sociocultural infl uences on physical activity
can infl u ence total energy expenditure and thus
conditions energy require ments. In the average 5 year old
child weighing 20 kg, total energy expenditure is
Infancy and childhood
approximately 5.5–5.9 MJ/ day, which is signifi
Changes in energy intake during infancy have cantly lower than the existing rec ommended daily
been well characterized. During the fi rst 12 allowance for energy in children of this age, by
months of life, energy intake falls from almost 525 approximately 1.7–2.1 MJ/day. Thus, as with
kJ/kg per day in the fi rst month of life to a nadir of infants, newer estimates of energy needs in chil
399 kJ/kg per day by the eighth month, then rises dren are needed based on assessment of total
to 441 kJ/kg per day by the 12th month. However, energy expenditure data.
total energy expendi
ture in the fi rst year of life is relatively constant at Aging
around 252–294 kJ/kg per day. In infants, the In the elderly, two different problems related to
large difference between total energy expenditure energy balance can be recognized. In one
and energy intake is explained by a positive segment of the elderly population there is a
energy balance to account for growth. In the fi rst decline in food intake that is associated with
3 months of life it is estimated that the energy dynamic changes in body com
accretion due to growth is 701.4 kJ/day, or position where there is a tendency to lose FFM,
approximately 32% of energy intake, falling to which leads to loss in functionality. In others there
151.2 kJ/day, or 4% of energy intake, by 1 year of is a tendency to gain fat mass, which increases
age. Individual growth rates and early infancy the risk for obesity, cardiovascular disease, and
feeding behavior are at least two known factors noninsulin
that would cause variation in these fi gures. dependent diabetes. These two opposing
There is now substantial evidence to suggest patterns suggest that the ability to self-regulate
that existing recommendations may overestimate whole body energy balance may diminish with
true energy needs, based on measurement of aging. Thus, pre scription of individual energy
total energy expenditure in infants. In the fi rst requirements may serve as a useful tool to
year of life, tradi prevent the age-related deteriora tion of body
tional values of energy requirements overestimate composition. Other special consider ations in the
those derived from measurement of total energy elderly relate to meeting energy needs in special
expenditure and adjusted for growth by 11%. populations, such as those with Alzheimer's and
Between 1 and 3 years of age the discrepancy is Parkinson's disease, which frequently can lead to
more striking, where the traditional values for malnourished states and a diminishing of body
requirements are 20% higher than those derived weight. It was thought that these neurological
from total energy expendi condi tions may lead to body weight loss because
ture and adjusted for growth. For example, in 3 of an associated hypermetabolic condition in
 which meta bolic rate may increase above
normal, thus increasing energy needs. However,
more recent studies have clearly shown that the
wasting or loss of body weight often associated
with these conditions is explained by a reduction
in food intake, probably owing to a loss in
functionality.
Energy requirements in physically
active groups
The DLW technique has been used to assess
energy requirements in highly physically active
groups of people. The most extreme case is a
study that assessed the energy requirements of
cyclists performing in the 3 week long Tour de
France bicycle race. The level of total energy
expenditure recorded (PAL factor of 5.3, or
approximately 35.7 MJ/day) was the highest
recorded sustained level in humans. In another
study involving young male soldiers training for
jungle warfare, energy requirements were 19.9
MJ/day (PAL factor of 2.6). The total energy
expenditure of four mountaineers climbing Mount
Everest was 13.6 MJ/day (PAL 2.0–2.7), which
was similar to energy expenditure during on-site
preparation prior to climbing (14.7 MJ/day). Total
energy expenditure in free-living collegiate
swimmers was almost 16.8 MJ/ day in men and
10.9 MJ/day in women. In elite female runners
previously performed studies of energy intake
suggested unusually low energy requirements.
However, in a study in nine highly trained young
women, free-living energy expenditure was 11.9 ±
1.3 MJ/day, compared with the reported energy
intake of 9.2 ± 1.9 MJ/day. This study suggests
that elite female runners underreport true levels of
energy intake and confi rms the absence of
energy-saving metabolic adaptations in this
population.
Regular participation in exercise is traditionally
thought to elevate energy requirements through
the additional direct cost of the activity, as well as
through an increase in RMR. However, in some
situations energy requirements are not
necessarily altered by participation in regular
physical activity. For example, in a study of an
elderly group of healthy volunteers, there was no
signifi cant change in total energy ex penditure in
the last 2 weeks of an 8 week vigorous
endurance training program. The failure to detect
an increase in total energy expenditure occurred
despite a 10% increase in RMR (6703.2 ± 898.8
to 7404.6 ± 714 kJ/day), as well as an additional
630 kJ/ day associated with the exercise
program. These increases in energy expenditure
were counteracted by a signifi cant reduction in
the energy expenditure of physical activity during
nonexercising time (2.4 ± 1.6 versus 1.4 ± 1.9
MJ/day). The lack of increase in total energy
expenditure in this study is probably explained by
a compensatory energy-conserving adaptation to
Energy Metabolism 43
this vigorous training program leading to a
reduction in spontaneous physical activity and/or
a reduction in voluntary physical activities, similar
to that observed in several animal studies. Thus,
it should not automatically be assumed that
energy require
ments are elevated by participation in activity
programs, and the ultimate change in energy
requirements may be dictated by the intensity of
the training program and the net sum of change in
the individual components of energy expenditure.
An important area of research is to identify the
optimal program of exercise intervention in terms
of exercise mode, type, duration, and intensity
that can have optimal effects on all components
of energy balance.
Energy requirements in pregnancy
and lactation
Pregnancy and lactation are two other examples
in which energy metabolism is altered in order to
achieve positive energy balance. The specifi c
changes in energy requirements during
pregnancy are unclear and the various factors
affecting this change are complex. Traditional
government guidelines suggest that energy
requirements are raised by 1.3 MJ/day during
pregnancy. This fi gure is based on theoretical
calculations based on the energy accumulation
asso ciated with pregnancy. However, these fi
gures do not include potential adaptations in
either metabolic effi -
ciency or PAL during pregnancy. In a study that
per formed measures in 12 women every 6 weeks
during pregnancy the average increase in total
energy expenditure was 1.1 MJ/day. The average
energy cost of pregnancy (change in total energy
expenditure plus change in energy storage) was
1.6 MJ/day. However, there was considerable
variation among the 12 subjects for the increase
in average total energy expenditure (264.6 kJ/day
 to 3.8 MJ/day) and the average energy cost of different than controls (matched for age, gender,
pregnancy (147 kJ/day to 5.2 MJ/day). and height). However, physical activity-related
Metabolic adaptations during lactation have energy expendi
been examined in well-nourished women using ture was 1.3 MJ/day higher in anorexia nervosa
the DLW technique. The energy cost of lactation patients, which was compromised by a 1.3
was calculated to be 3.7 MJ/day. Just over half of MJ/day lower RMR. Thus, energy requirements in
this energy cost was achieved by an increase in anorexia nervosa patients are normal, despite
energy intake, while the remainder was met by a alterations in the individual components of total
decrease in physical activity energy expenditure energy expenditure. In infants with cystic fi brosis,
(3.2 MJ + 873.6 kJ/day at 8 weeks of lactation total energy expendi
compared with 3.9 + 1.1 MJ/day in the same ture was elevated by 25% relative to weight-
women prior to pregnancy). matched controls, although the underlying
44 Introduction to Human Nutrition mechanism for this effect is unknown.
Developmental disabilities appear to be
associated with alterations in energy balance and
Energy requirements in disease and trauma nutritional
The DLW technique has been used in various
studies to assess the energy requirements of
hospitalized patients. Information on energy
requirements during hospitalization for disease or status at opposite ends of the spectrum. For
trauma is important because: example, cerebral palsy is associated with
reduced fat mass and FFM, whereas half of
● energy expenditure can be altered by the
patients with myelodysplasia are obese. It is
disease or injury unclear whether the abnormal body com
● physical activity is often impaired or reduced ●
position associated with these conditions is the
both underfeeding and overfeeding of critically ill end result of inherent alterations in energy
patients can lead to metabolic complications; expenditure and/or food intake, or whether
there fore, correct assessment of energy alterations in body composition are an inherent
requirements during recovery is an important part part of the etiology of the specifi c disability. In
of therapy. addition, it is unclear how early in life total energy
The metabolic response during recovery from a expenditure may be altered and whether reduced
burn injury includes an increase in RMR, although energy expenditure is involved with the
this is not necessarily a function of the extent of associated obese state. Nevertheless, pre
the burn. The widely used formulae to predict scription of appropriate energy requirements may
energy needs in burn patients are not based on be a useful tool in the improvement of nutritional
measurement of energy expenditure and estimate status in developmental disabilities.
that most patients require 2–2.5 times their Total energy expenditure has been shown to be
estimated RMR. However, using the DLW lower in adolescents with both cerebral palsy and
technique, total energy expenditure was 6.7 + 2.9 myelodysplasia, partly owing to reduced RMR but
MJ/day in 8 year old children recovering from primarily to reduced physical activity. Based on
burn injury, which was equivalent to only 1.2 measurements of total energy expenditure,
times the nonfasting RMR. The lower than energy requirements of adolescents with cerebral
expected values for total energy expenditure in palsy and myelodysplasia are not as high as
children recov ering from burns suggest that RMR previously specu
is not as elevated in burn patients as previously lated. In nonambulatory patients with cerebral
speculated, and that RMR is not a function of palsy, energy requirements are estimated to be
burn size or time after the injury, probably owing 1.2 times RMR, and in the normal range of 1.6–
to improvements in wound care which reduce 2.1 times RMR in ambulatory patients with
heat loss. In addition, energy requirements in cerebral palsy.
patients recovering from burn injury are reduced
because of the sedentary nature of their
hospitalization. 3.7 Obesity
In a study of patients with anorexia nervosa,
total energy expenditure was not signifi cantly Basic metabolic principles
 Obesity is the most common form of a disruption of glyco
in energy balance and now constitutes one of the gen that is stored, the body has to store an
major and most prevalent disorders of nutrition. additional 3 g of water. Thus, for each 4 g of
Because of the strong relationship between storage tissue, the body stores only 16.8 kJ,
obesity and health risks, obesity is now generally equivalent to just 4.2 kJ/g, compared with the
considered a disease by health professionals. benefi t of fat which can be stored as 37.8 kJ/g.
Although the body continuously consumes a Thus, a typical adult with 15 kg of fat carries
mixed diet of carbohydrate, protein, and fat, and 567.0 MJ of stored energy. If the adult did not eat
sometimes alcohol, the preferred store of energy and was inactive, he or she might require 8.4
is fat. There is a clearly defi ned hierarchy of MJ/day for survival, and the energy stores would
energy stores that out be suffi cient for almost 70 days. This length is
lines a preferential storage of excess calories as about the limit of human survival without food.
fat. For alcohol, there is no storage capacity in the Given that glycogen stores require 4 g to store
body. Thus, alcohol that is consumed is 4.2 kJ (3 g of water plus 1 g of gly cogen = 16.8
immediately oxidized for energy. For protein, kJ), we can calculate that to carry this much
there is a very limited storage capacity and, under energy in the form of glycogen requires 135 kg of
most situations, protein metabo weight. It is no wonder therefore that the body's
lism is very well regulated. For carbohydrate there metabolism favors fat as the preferred energy
is only a very limited storage capacity, in the form store.
of glycogen, which can be found in the liver and
in Defi nition of obesity
Obesity has traditionally been defi ned as an
excess accumulation of body energy, in the form
of fat or adipose tissue. Thus, obesity is a disease
muscle. Glycogen provides a very small and short of positive energy balance, which arises as a
term energy store, which can easily be depleted result of dysregula
after an overnight fast or after a bout of exercise. Energy Metabolism 45
Most carbohydrate that is consumed is
immediately used for energy. Contrary to popular
belief, humans cannot convert excess tion in the energy balance system – a failure of
carbohydrate intake to fat. Instead, when excess the regulatory systems to make appropriate
carbohydrates are consumed, the body adapts by adjustments between intake and expenditure. It is
preferentially increasing its use of carbohy drate now becoming clear that the increased health
as a fuel, thus, in effect, burning off any excessive risks of obesity may be conferred by the
carbohydrate consumption. Large excesses of distribution of body fat. In addition, the infl uence
carbohydrate may induce de novo lipogenesis, of altered body fat and/or body fat dis
but normally this process is quantitatively minor. tribution on health risk may vary across
However, no such adaptive mechanism for fat individuals. Thus, obesity is best defi ned by
exists. In other words, if excess fat is consumed, indices of body fat accumulation, body fat pattern,
there is no mechanism by which the body can and alterations in health risk profi le.
increase its use of fat as a fuel. Instead, when The body mass index (BMI) is now the most
excess fat calories are con sumed, the only accepted and most widely used crude index of
option is to accumulate the excess fat as an obesity. This index classifi es weight relative to
energy store in the body. This process occurs at a height squared. The BMI is therefore calculated
very low metabolic cost and is therefore an as weight in kilo
extremely effi cient process. To store excess grams divided by height squared in meters, and
carbohydrate as glycogen is much more expressed in the units of kg/m2. Obesity in adults
metabolically expensive and therefore a less effi is defi ned as a BMI above 30.0 kg/m 2, while the
cient option. There is another important reason normal range for BMI in adults is 18.5–24.9
why the body would prefer to store fat rather than kg/m2. A BMI in the range of 25–30 kg/m 2 is
glycogen. Glycogen can only be stored in a considered overweight. In children, it is more diffi
hydrated form that requires 3 g of water for each cult to classify obesity by BMI because height
gram of glycogen, whereas fat does not require varies with age during growth; thus, age-adjusted
any such process. In other words, for each gram BMI percentiles must be used.
 One of the major disadvantages of using the Although there are genetic infl uences on the
BMI to classify obesity is that this index does not various components of body-weight regulation,
distin guish between excess muscle weight and and a major portion of individual differences in
excess fat weight. Thus, although BMI is strongly body weight can be explained by genetic
related to body fatness, at any given BMI in a differences, it seems unlikely that the increased
population, there may be large differences in the global prevalence of obesity has been driven by a
range of body fatness. A classic example of dramatic change in the gene pool. It is more likely
misclassifi cation that may arise from the use of and more reasonable that acute changes in
the BMI is a heavy football player or body-builder behavior and environment have contributed to the
with a large muscle mass who may have a BMI rapid increase in obesity, and genetic factors may
above 30 kg/m2 but is not obese; rather, this man be important in the differing individual
has a high body weight for his height resulting susceptibilities to these changes. The most
from increased FFM. striking behavioral changes that have occurred
Since the health risks of obesity are related to have been an increased reliance on high-fat and
body fat distribution, and in particular to excess energy-dense fast foods, with larger portion sizes,
abdominal fat, other anthropometric indices of coupled with an ever-increasing seden tary
body shape are useful in the defi nition of obesity. lifestyle. The more sedentary lifestyle is due to an
Traditionally, the waist-to-hip ratio has been used increased reliance on technology and labor-
as a marker of upper versus lower body-fat saving devices, which has reduced the need for
distribution. More recent studies suggest that physical activ ity for everyday activities. Examples
waist circumference alone pro of energy-saving devices are:
vides the best index of central body-fat pattern
and increased risk of obesity-related conditions. ● increased use of automated transport rather than
The rec ommended location for the measurement walking or cycling
of waist circumference is at the midpoint between ● central heating and the use of automated equip
the lowest point of the rib cage and the iliac crest. ment in the household, eg, washing machines ●
The risk of obesity-related diseases is increased reduction in physical activity in the workplace due
above a waist cir cumference of 94 cm in men to computers, automated equipment, and elec
and above 80 cm in women. tronic mail, which all reduce the requirement for
46 Introduction to Human Nutrition physical activity at work

Etiology of obesity: excess intake or

decreased physical activity
Stated simply, obesity is the end-result of positive
energy balance, or an increased energy intake
relative to expenditure. It is often stated, or
assumed, that obesity is simply the result of
overeating or lack of physical activity. However,
the etiology of obesity is not as simple as this,
and many complex and interre
lated factors are likely to contribute to the develop
ment of obesity; it is extremely unlikely that any
single factor causes obesity. Many cultural, behav
ioral, and biological factors drive energy intake
and energy expenditure, and contribute to the
homeo static regulation of body energy stores, as
discussed earlier in the chapter. In addition, many
of these factors are infl uenced by individual
susceptibility, which may be driven by genetic,
cultural, and hor monal factors. Obesity may
develop very gradually over time, such that the
actual energy imbalance is negligible and
undetectable.
● increased use of television and computers for
enter tainment and leisure activities
● use of elevators and escalators rather than using
stairs
● increased fear of crime, which has reduced the
like lihood of playing outdoors
● poor urban planning, which does not provide
adequate cycle lanes or even pavements in
some communities.
Thus, the increasing prevalence, numerous
health risks, and astounding economic costs of
obesity clearly justify widespread efforts towards
prevention.
The relationship between obesity and lifestyle
factors refl ects the principle of energy balance.
Weight maintenance is the result of equivalent
levels of energy intake and energy expenditure.
Thus, a discrepancy between energy expenditure
and energy intake de
pends on either food intake or energy
expenditure, and it is becoming clear that physical
 activity provides the main source of plasticity in expenditure). Secondly, physical activity has
energy expenditure. In addition, lifestyle factors benefi cial effects on substrate metabo lism, with
such as dietary and activ an increased reliance on fat relative to
ity patterns are clearly susceptible to behavioral carbohydrate for fuel utilization, and it has been
mod ifi cation and are likely targets for obesity hypothesized that highly active individuals can
prevention programs. A second, yet related, main tain energy balance on a high-fat diet.
reason that control of the obesity epidemic will Cross-sectional studies in children and adults
depend on preventive action is that both the have shown that energy expenditure, including
causes and health consequences of obesity begin physical activity energy expenditure, is similar in
early in life and track into adulthood. For example, lean and obese subjects, especially after
both dietary and activity patterns responsible for controlling for differ
the increasing prevalence of obesity are evident ences in body composition. Children of obese and
in childhood. lean parents have also been compared as a
model of preobesity. Some studies show that
Role of physical activity and energy children of obese parents had a reduced energy
expenditure in the development of obesity expenditure, including physical activity energy
Although it is a popular belief that reduced levels expenditure, whereas another study did not. A
of energy expenditure and physical activity lead to major limitation of the majority of studies that
the development of obesity, this hypothesis have examined the role of energy expen
remains con troversial and has been diffi cult to diture in the etiology of obesity is their cross-
prove. There are certainly good examples of an sectional design. Because growth of individual
inverse relationship between physical activity and components of body composition is likely to be a
continuous process, longitudinal studies are
obesity (eg, athletes are lean and nonobese
necessary to evaluate the rate of body fat change
individuals), as well as good examples of the
during the growing process. Again, some
positive relationship between obesity and physical
longitudinal studies support the idea that reduced
inactivity (obese individuals tend to be less
energy expenditure is a risk factor for the
physically active). However, not all studies
development of obesity, whereas others do not.
provide supporting evidence. For example,
Finally, intervention studies have been conducted
several studies suggest that increased television to determine whether the addition of physical
viewing (as a marker for inactivity) increases the activity can reduce obesity. These studies tend to
risk of obesity, whereas others do not. Similar to support the positive role of physical activity in
the results for physical activ- reducing body fat.
Several possibilities could account for such dis
crepant fi ndings. First, the ambiguous fi ndings in
ity, some studies suggest that a low level of the literature may be explained by the possibility
energy expenditure predicts the development of that
Energy Metabolism 47
obesity, and others do not support this
hypothesis.
Physical activity is hypothesized to protect differences in energy expenditure and physical
people from the development of obesity through activ ity and their impact on the development of
several channels. First, physical activity, by defi obesity are different at the various stages of
nition, results in an increase in energy maturation. This hypothesis is supported by
expenditure owing to the cost of the activity itself, previous longitudinal studies in children, showing
and is also hypothesized to increase RMR. These that a reduced energy expenditure is shown to be
increases in energy expenditure are likely to a risk factor for weight gain in the fi rst 3 months
decrease the likelihood of positive energy of life, but not during the steady period of
balance. However, the entire picture of energy prepubertal growth. Secondly, there could be
balance must be considered, particularly the individual differences in the effect of altered
possibility that increases in one or more energy expenditure on the regulation of energy
components of energy expenditure can result in a balance. Thus, the effect of energy expenditure
compensatory reduction in other components (ie, on the etiology of obesity could vary among
resting energy expenditure and activity energy different sub
groups of the population (eg, boys versus girls, dif Obesity has recently been defi ned as a disease
ferent ethnic groups) and could also have a by the World Health Organization. The growing
differential effect within individuals at different problem of obesity worldwide, and in chil dren
stages of devel opment. It is conceivable that and in people who were previously food inse
susceptible individuals fail to compensate for cure and malnourished, needs to be addressed
periodic fl uctuations in energy expenditure. Third, with
explanations related to the meth odology can also 48 Introduction to Human Nutrition
be offered because of the complexity of the
nature of physical activity and its measure ment.
The success of controlled exercise interven tions better information about the behavioral and
in improving body composition indicates an cultural factors that infl uence energy balance.
extremely promising area for the prevention of This demands a more holistic, integrated
obesity. However, further studies are required to approach to the study of obesity in the future.
elu cidate the specifi c effects of different types of
exercise on the key features of body weight Acknowledgment
regulation.
This chapter has been revised and updated by
3.8 Perspectives on the future Arne Astrup and Angelo Tremblay based on the
original chapter by Michael I Goran and Arne
Much is known about how the body balances Astrup.
energy intake and expenditure. There are,
however, areas that need further research. The
technology to deter mine total energy expenditure
with doubly labeled water has been standardized. Reference
Most of the data from using this method have
been obtained in populations living in Blundell JE, Rogers PJ, Hill AJ. Evaluating the satiating power
of foods: implications for acceptance and consumption. In:
industrialized countries. More studies on infants, Solms J, Booth DA, Pangbourne RM, Raunhardt O, eds.
children, adolescents, adults, pregnant and Food Acceptance and Nutrition. Academic Press, London,
lactating women, and the elderly living in 1987: 205–219.
developing countries are indicated. Doubly
labeled water is an expensive method. There is a Further reading
need to develop more cost-effective methods that
can be used in fi eld studies and to determine the Bray G, Bouchard. D, eds. Handbook of Obesity, 3rd edn.
Informa Healthcare, New York, 2008.
energy cost of specifi c activities of people DeFronzo RA, Ferrannini E, Keen H, Zimmet P. International
throughout the life cycle in developing countries. Textbook of Diabetes Mellitus, 3rd edn. John Wiley & Sons,
Chichester, 2004.

4
Nutrition and Metabolism of
Proteins and Amino Acids
Naomi K Fukagawa and Yong-Ming Yu

Key messages
• Protein is the most abundant nitrogen-containing when L-α-amino acids polymerize via peptide bond
compound in the diet and the body. Proteins are formed formation.
 • Amino acids have similar central structures with different
side chains determining the multiple metabolic and
physiological roles of free amino acids.
• Indispensable (essential) amino acids cannot be
synthesized by humans from materials ordinarily
available to cells at a speed commensurate with the
demands of human growth and maintenance.
• The requirements for indispensable amino acids can be
defi ned as “the lowest level of intake that achieves
nitrogen balance or that balances the irreversible
oxidative loss of the amino acid, without requiring major
changes in normal protein turnover and where there is
energy balance with a modest level of physical activity.”
For infants, children, and pregnant and lactating women,
requirements would include protein deposited and
secretion of milk proteins.
• “Conditionally” indispensable amino acids are those for
which there are measurable limitations to the rate at
which they can be synthesized because their synthesis
requires another amino acid and because only a number
of tissues are able to synthesize them, and probably only
in limited amounts. The metabolic demands for these
amino acids may rise above the biosynthetic capacity of
the organism.
• Protein and amino acid requirements are determined by
the pro cesses of protein synthesis, and maintenance of cell
and organ protein content, as well as the turnover rates of
protein and
4.1 Introduction
Protein is the most abundant nitrogen-containing
compound in the diet and in the body. It is one of
the fi ve classes of complex biomolecules present
in cells and tissues, the others being DNA, RNA,
polysaccha
rides, and lipids. The polymerization of L-α-amino
© 2009 NK Fukagawa and YM Yu.
amino acid metabolism, including synthesis, breakdown,
inter conversions, transformations, oxidation, and
synthesis of other nitrogen-containing compounds and
urea. These processes are infl uenced by genetics,
phase of life cycle, physical activity, dietary intake levels,
how energy needs are met, route of delivery
of nutrients, disease, hormones, and immune system
products. • Protein and amino acid requirements can be
determined by nitro gen excretion and balance, factorial
estimations, and/or tracer techniques.
• Existing recommendations on requirements differ by
various authorities because of a lack of data when some
were formu lated, different interpretations of data, and
different criteria for judging adequate intakes.
• The United Nations plans to publish new
recommendations for protein and amino acids in the near
future. Those made by the Institute of Medicine, US
National Academies of Science, in 2002 are cited in this
chapter.
• Apparent protein digestibility, measured in the past as the
differ ence between nitrogen intake and fecal nitrogen
output, under estimates “true” digestibility because fecal
nitrogen is derived, in part, from endogenous nitrogen
sources.
• Tracer techniques have shown that “true” digestibility of
most dietary proteins is high. The quality of food protein
can be assessed as the protein digestibility-corrected
amino acid score.
• Animal protein foods generally have higher concentrations
of indispensable amino acids than plant foods. Lysine is
often the most limiting amino acid, followed by sulfur
amino acids (methio nine and cystine) and tryptophan
and threonine.
acids through synthesis of peptide bonds
contributes to the formation and structural
framework of pro teins. These may contain two or
more polypeptide chains forming multimeric
proteins, with the indi vidual chains being termed
subunits. Proteins are the workhorses in cells and
organs and their building blocks are the amino
acids, which are joined together
50 Introduction to Human Nutrition
according to a sequence directed by the base
sequence of the DNA (the genome), and so they
serve as the currency of protein nutrition and
metabolism. The Human Genome Project
completed in 2000 revealed that the human
genome consists of only 30 000 genes, whereas
there may be hundreds of thousands of pro
teins that are responsible for giving a human its
par ticular characteristics and uniqueness. A new
fi eld of nutrition research has now opened up
and is referred to as “nutrigenomics,” which is the
study of how nutrition and genomics interact to
infl uence health. Proteins and amino acids fulfi ll
numerous functions, many of which are
summarized in Table 4.1. Some
Table 4.1 Some functions of amino acid and
proteins Function Example
Amino acids
amino acids, such as glutamine (Tables 4.2 and
4.3), play multiple roles. It is not surprising,
therefore, that inappropriate intakes of proteins
and/or of specifi c amino acids can have
important consequences for tissue and organ
function, and the maintenance of health and the
well-being of the individual.
 This chapter begins with a short historical Section 4.8 examines how factors other than
perspec tive and then moves in Sections 4.3 and dietary protein can infl uence the requirements for
4.4 to discuss the structure, chemistry, and proteins and amino acids.
classifi cation of amino acids. Section 4.5 is
concerned with the biology of protein and amino 4.2 A historical perspective
acid requirements, with Sections 4.6 and 4.7
describing how the requirements are established The early history of protein metabolism and
and how they may be met, respectively. Finally, nutrition
Substrates for protein synthesis NAG synthase nature. The reason for this is that
Regulators of protein turnover
Regulators of enzyme activity (allosteric) pro teins, on average, contain
Those for which there is a codon Leucine; is closely tied to the discovery of about 16% nitrogen by weight (to
nitrogen and its distribution in convert nitrogen to protein it is
sistein; arginin; glutamine Glutamate and
necessary
Phenylalanine and PAH activation Edinburgh,
to multiply by 6.25). Daniel Rutherford, in
Precursor of signal transducer can be regarded as the discoverer “phlogisticated air” in his
Arginine and nitric oxide
of nitrogen, which he called Doctorate in
Methylation reactions Methionine
Neurotransmitter Tryptophan (serotonin); glutamine Ion
fl uxes Taurine; glutamate
Precursor of “physiologic” molecules Histidine/β-alanine (carnosine) Substrate of protein synthesis (codons:
Arg (creatinine); Glu-(NH2) purines Cysteine/glycine/glutamate (glutathione) CAA, CAG)
Table 4.2 Multiple functions of an amino Anabolic/trophic substance for muscle;
acid; glutamine as an example intestine (“competence
Transport of nitrogen Alanine; glu-(NH2) factor”)
Regulator of gene transcription Leucine: alters activity of initiation factor Substrate for hepatic/renal
Regulator of mRNA translation 4E-BP and P70 (6SK) via mTOR signaling gluconeogenesis
pathway Fuel for intestinal enteroctyes
Controls acid–base balance (renal Fuel and nucleic acid precursor and
Proteins
ammoniagenesis) Substrate for hepatic important for generation of cytotoxic
Amino acid depletion and asparagine
ureagenesis products in immunocompetent cells
synthase gene activation
Enzymatic catalysis Branched chain ketoacid Substrate for citrulline and arginine synthesis
dehydrogenase Nitrogen donor (nucleotides, amino sugars,
Transport B12 binding proteins; ceruloplasmin; apolipoproteins; coenzymes) Nitrogen transport (1/3 circulating N)
albumin (muscle; lung) Precursor of GABA (via glutamate)
Messengers/signals Insulin; growth hormone; IGF-1 Shuttle for glutamate (CNS)
Movement Kinesin; actin; myosin Structure Collagens; Preferential substrate for GSH production?
elastin; actin Storage/sequestration Ferritin; metallothionein Osmotic signaling mechanism in regulation of protein
Immunity Antibodies; cytokine, chemokines synthesis? Stimulates glycogen synthesis
Ammonia scavenger
Growth; differentiation; gene expression factors (umami)
Peptide growth factors; transcription L-Arginine NO metabolism Taste factor

IGF-1, insulin-like growth factor-1; NAG, N-acetyl glutamate; Alanine Ubiquitous nitrogen donor, extracellular
PAH, phenylalanine hydroxylase; glu-(NH2), glutamine. Transporter of three-carbon units
CNS, central nervous system; GABA, γ-aminobutyric acid; Aspartate Ubiquitous nitrogen donor Transfer form of
GSH, growth stimulating hormone; NO, nitric oxide. nitrogen from
cytoplasmic amino acids to
urea
Single carbon metabolism
Table 4.3 Biochemical roles of amino acids not directly related Methionine Donor and acceptor of methyl groups
to protein metabolism Important role in single-carbon
metabolism
Amino acid Biochemical function Glycine Donor of methylene groups Serine Donor of
hydroxymethylene groups
Integration of carbon and nitrogen metabolism
Neurotransmitter synthesis
Leucine, isoleucine, valine Ubiquitous nitrogen donors and
Histidine Precursor for histamine synthesis Phenylalanine
metabolic fuel
and tyrosine Precursors for tyramine, dopamine,
Ubiquitous nitrogen donor,
epinephrine, and
extracellular
Glutamate Transporter of four-carbon units Glutamine See norepinephrine synthesis
Table 4.2 Tryptophan Precursor for serotonin synthesis Glutamate
 Precursor for γ-aminobutyric acid synthesis
Miscellaneous
Arginine Immediate precursor for urea Precursor for nitric
oxide synthesis
Cysteine Potential intracellular thiol buffer Precursor for
glutathione and
taurine synthesis
Glycine Nitrogen donor for heme synthesis
Histidine/β-alanine Precursors for carnosine synthesis
Medicine thesis in 1792. The fi rst amino acid to
be discovered was cystine, which was extracted
from a urinary calculus by Wallaston in England in
1810. It was not until 1935 that threonine, the last
of the so
called nutritionally indispensable (essential)
amino acids for mammals, including man, was
discovered by WC Rose at the University of
Illinois. Finally, the term “protein” was invented by
the Swedish chemist Jons Jakob Berzelius
(1779–1848) and this was later accepted and
promoted by the infl uential Dutch chemist
Gerhardus Mulder in 1838.
Nutrition and Metabolism of Proteins 51
The nutritional importance of nitrogenous com
ponents in the diet was fi rst recognized in 1816
by Magendie. He described experiments in dogs
that received only sugar and olive oil until they
died within a few weeks. It was concluded that a
nitrogen source was an essential component of
the diet. Magendie's insightful views on nitrogen
metabolism and nutri tion were followed by
studies carried out by the French scientifi c
school, including Justus von Leibig, who
investigated the chemical basis of protein meta
bolism and discovered that urea was an end-
product of protein breakdown in the body. Later,
Leibig founded a school of biochemical studies in
Gissen and later in Munich, Germany, from which
52
Introduction
to Human
Nutrition
HO CH2
Histidine
Carl Voit emerged as a distinguished scientist
and laid the foundations of modern studies of
body nitrogen balance. He, in turn, trained many
famous scientifi c celebrities, including Max
Rubner, from Germany, who studied the specifi c
dynamic action of proteins and their effects on
energy metabolism, and Wilbur Atwater and
Graham Lusk, from the USA, who studied food
composition, protein requirements, and energy
metabolism. Through their work, and that of
others, theories of protein metabolism were
proposed and challenged, leading to the more or
less contemporary view which was established
through the seminal work of Rudolf
Schoenheimer, conducted at Columbia
University, New York, in the mid-1930s and early
1940s. He applied the new tracer tool of stable
isotope
enriched compounds, especially amino acids, in
the study of dynamic aspects of protein turnover
and amino acid metabolism. Stable isotopes
(such as 13C, 18O, and 15N) are naturally present in
our environ
ment, including the foods we eat, and they are
safe to use in human metabolic studies. Using
this approach, Schoenheimer established the
fundamental biological principle of a continued
tissue and organ protein loss and renewal, which
forms the basis for the dietary need for protein or
supply of amino acids and a utiliz
able form of nitrogen.
4.3 Structure and chemistry of
amino acids
With the exception of proline, the amino acids that
make up peptides and proteins have the same
central structure (Figure 4.1; the A in this fi gure
and
(a) (b)
Tyrosine Proline
A
COOH
N
H
Tryptophan
A
CH2CH2 A
H N H
N N H Phenylalanine

CH2
A

(c)

Figure 4.1 Structures of some of the nutritionally important amino acids. All are components of proteins and they are coded
by DNA. (a) Nutritionally indispensable (essential) includes also tryptophan and histidine; (b) nutritionally conditionally
indispensable; (c) nutritionally dispensable.

H amino acids within the linear peptide structure,

| while the side-chains distinguish the physical and
subsequent fi gures represent the – C – COOH chemical properties of each chemical class of
moiety). | amino acid. In addition, some features of the
NH2 amino acid side-chains are critical to the
metabolic and physiological roles of free, as
The carboxylic acid and amino nitrogen groups
opposed to protein-bound, amino acids (Table
are the components of the peptide bond that links
4.3; Figures 4.1 and 4.2). These roles are refl
the
ections of

Amino butyric acid N

Nutrition and Metabolism of Proteins 53

● dopamine is formed from tyrosine and fulfi lls a

neurotransmitter function ing molecules modulating the process of protein
● ornithine can be formed from glutamate and synthesis. The translation of mRNA into protein in
serves as both an intermediate in the urea cycle skeletal muscle is initiated from (1) the binding of
and a precursor of the polyamines spermine and met-tRNA to the 40S ribosomal subunit to form the
spermi dine, which are used in DNA packaging. 43S preinitiation complex; (2) the subsequent
binding of this complex to mRNA and its localization
Finally, other amino acids (Figure 4.3) appear in pro to the AUG start codon; and (3) the release of the
teins via a post-translational modifi cation of a initiation factors from the 40S ribosomal complex to
specifi c amino acid residue in the polypeptide allow the formation of the 80S ribosomal complex
chain that is being formed during protein synthesis. via the joining of the 60S ribosomal subunit. Then
In addition to serving the function as precursors for the 80S ribosomal complex proceeds to the
protein synthesis, amino acids also serve as signal elongation stage
heterotrimeric complex of eIF–4F proteins. The
Figure 4.2 Physiologically important amino acid metabolites.
Both the metabolic relationship between alanine and glutamic
signaling pathway regulating mRNA translation
acid and their transamination partners, the keto acids pyruvate
involves the protein kinase termed the
and α ketoglutarate, and the similarity between the catabolic
mammalian target of rapamycin (mTOR). mTOR
oxidation pathway of the branched-chain amino acids and the
regulates the formation of the eIF–4F complex via
β-oxidation pathway of saturated fatty acids are shown.
a series of phosphorylation–dephosphorylation
either their specifi c chemical properties or specifi pro
c metabolic interrelationships. Examples of the cesses of the downstream targets. The mTOR
former are the facility of methionine to donate a signal ing pathway is traditionally considered to
methyl group in one-carbon metabolism, the be solely involved in mediating the action of
propensity for the amide group of glutamine to hormones. Recent studies revealed that the
serve as a nitrogen source for pyrimidine branched-chain amino acids, especially leucine,
synthesis, or the sulfhydryl group of cysteine serve a unique role in regulating mRNA
forming disulfi de bonds for cross-linking. The translation via the same mTOR-signaling
former metabolic relationship allows alanine and pathway. Increased availability of leucine
glutamate (and glutamine) to provide a link activates the mTOR and its downstream targets.
between carbohydrate and protein metabolism; However, inhibi tion of the mTOR pathway by
the latter enables the branched amino acids to rapamycine partially inhibits the stimulatory effect
function when required, as a “universal” fuel of leucine on protein synthesis, indicating the
throughout the body. involvement of an mTOR independent signaling
Some of these amino acid and nitrogen pathway by leucine in the reg ulation of protein
compounds are derivatives of other amino acids: synthesis. The detailed mechanisms involved in
these regulations, especially those of the mTOR-
● creatine is formed from glycine, arginine, and
independent pathways, remain an active fi eld of
methionine and serves in intracellular energy research.
transduction Furthermore, individual amino acids play
of translation. The formation of the 43S multiple regulatory roles in health and diseased
preinitiation complex is mediated by a conditions.
54 Introduction to Human Nutrition

NG,NG-dimethyl-L-arginine (ADMA) CH3 active area of investigation. In

has been well established. The general, these nonprotein func
CH3
physiology of the argi nine–nitric tions of amino acids serve
NG,NG-dimethyl-L-arginine (SDMA)
oxide pathway has also been an important functions in the
HN CH2 N CH2 and other and functions are
N protective neuromuscula primarily
CH2 C CH2
C functions; (2) r function. It is exerted by
A NH A digestive also worth nutri
NH CH3 maintenance function; and noting that tionally
CH2
CH2
H 3C of (1) immune (3) cognitive these dispensable
HN
amino acids. pathways supply of important in physiological 4.4 Classifi acids
Hence, the de and/or the these amino modulating theconditions. cation of
novo amount of acids or their physiological
synthesis exogenous precursors are and patho amino
Ornithine animal organism out of materials ordinarily
available to the cells at a speed
commensurate with the demands for normal
growth.
There are three important phrases in this defi
nition: ordinarily available, at a speed and for
normal growth.
The phrase “ordinarily available” is an important
qualifi er within this defi nition because a number
of nutritionally essential amino acids, for example
the branched-chain amino acids, phenylalanine
and methionine, can be synthesized by
transamination of their analogous α-keto acids.
However, these keto acids are not normally part
of the diet and so are not “ordinarily available to
the cells.” They may be used in special situations
Figure 4.3 Some amino acids that arise via a post-
translational modi fi cation of a polypeptide-bound amino acid. such as in nitrogen-accumulating diseases,
These amino acids are not coded by DNA but are important including renal failure, where they may assist in
determinants of the structural and functional characteristics of maintaining a better status of body nitrogen
proteins. Shown are (1) the formation of hydroxyproline, from metabolism.
proline, involved in the maturation of the differ ent types of The phrase “at a speed” is equally important
collagens in cells; (2) the methylation of a specifi c histidine in
the muscle protein actin (it could be that this modifi cation because there are circumstances in which the
gives this protein its ability to function effectively in the rate of synthesis of an amino acid may be
contractile activities of the skeletal muscles that help us to constrained, such
move about); and (3) the methyla tion of arginine to form
asymmetric and symmetric dimethylarginine, which serve as
an endogenous nitric oxide synthase inhibitor and play
important roles in modulating nitric oxide production and organ
as by the availability of appropriate quantities of
blood fl ow in health and diseased conditions.
“nonessential” nitrogen. Further, the rate of
synthesis becomes of particular importance when
For example, glycine is an important anti-infl considering a group of amino acids, exemplifi ed
amma tory, immunomodulatory, and by arginine, cysteine, proline, and probably
cytoprotective agent through the glycine receptor glycine. These amino acids are frequently
on the cell surface. The role of cysteine in described as being conditionally indispensable.
regulating glutathione synthesis and its role in That is, their indispensability is depen dent upon
protection against oxidative damage the physiological or pathophysiological condition
of the individual.
“Indispensability” as a basis of Finally, the phrase “normal growth” is critical in
classifi cation two respects. First, it serves to emphasize that
the defi - nitions were originally constructed in the
For most of the past 65 years amino acids have context of growth. For example, for the growing
been divided into two general, nutritional rat arginine is an indispensable amino acid, but
categories: indis pensable (essential) and the adult rat does not require the presence of
dispensable (nonessential). This categorization arginine in the diet and so it becomes a
provided a convenient and gener ally useful way dispensable amino acid at that later stage of the
of viewing amino acid nutrition at the time. The life cycle. Of course, if the capacity to synthesize
original defi nition of an indispensable amino acid arginine is compromised by removing a signifi
was: cant part of the intestine which produces citrulline,
One which cannot be synthesized by the a precursor of arginine, then the adult rat once
again requires arginine as part of an adequate
diet. Second, by confi ning the defi nition to
growth, this fails to consider the importance of
amino acids to pathways of disposal other than
protein deposition. This aspect of amino acid
utilization will be considered below.
Chemical and metabolic characteristics as
bases of classifi cation
It is also possible to classify amino acids
according to their chemical and metabolic
characteristics rather than on the basis of their
need for growth. Examination of the amino acids
that are generally considered to be nutritionally
indispensable for humans and most other
mammals indicates that each has a specifi c
structural feature, the synthesis of which cannot
be accomplished owing to the absence of the
necessary mammalian enzyme(s) (Table 4.4).
Indeed, in obliga
a nonamino acid source of nitrogen (eg,
ammonium ion) and a carbon source (eg,
glucose). Accordingly, and from a knowledge of
biochemical pathways, the only true metabolically
indispensable amino acid is glutamic acid, and
possibly also glycine. This is because they can be
synthesized from glucose and ammonium ions, in
the case of glutamate, and from carbon dioxide
and ammonium ions, in the case of glycine.
However, the in vivo conditions may differ in both
qualitative and quantitative terms from studies in
test-tubes or in isolated cells in culture; amino
acid metabolism in vivo is inherently more
complex than is immediately evident from a
simple consideration of biochemical pathways
alone.
Table 4.5 The dietary amino acids of nutritional signifi cance
in humans

Indispensable Conditionally indispensable Dispensable

Table 4.4 Structural features that render amino acids
indispensable components of the diet of mammals Valine Glycine Glutamic acid (?) Isoleucine Arginine Alanine
Leucine Glutamine Serine Lysine Proline Aspartic acid
Methionine Cystine Asparagine Phenylalanine Tyrosine
Amino acid Structural feature Threonine (Taurine)a
Tryptophan (Ornithine)a
Leucine, isoleucine, valine Branched aliphatic side-chain
Lysine Primary amine Threonine Secondary alcohol Histidine (Citrulline)a
Methionine Secondary thiol Tryptophan Indole ring Sebuah
Phenylalanine Aromatic ring Histidine Imidazole Nonproteinogenic amino acids, which have nutritional
ring value in special cases.
Nutrition and Metabolism of Proteins 55 56 Introduction to Human Nutrition

tory carnivores, such as cats, the further loss of Sources of nonspecifi c nitrogen
some critical enzyme(s) renders these animals for humans
particularly dependent on dietary sources of In earlier texts it would have been stated that,
specifi c amino acids, such as arginine. The lack given a suffi cient intake of the indispensable
of arginine in a single meal when given to a cat amino acids, all that is then additionally needed to
can be fatal. However, even within this view, the support body protein and nitrogen metabolism
important term is “de novo synthesis” because would be a source of “nonspecifi c” nitrogen
some amino acids can be synthesized from (NSN) and that this could be in the form of a
precursors that are structurally very similar. For simple nitrogen-containing mixture, such as urea
example, methionine can be synthesized both by and diammonium citrate. However, this is no
transamination of its keto acid analogue and by longer a suffi cient description of what is actually
remethylation of homocysteine. According to this required to sustain an adequate state of protein
metabolic assessment of amino acids, threonine nutriture in the human. This can be illus trated by
and lysine are the only amino acids that cannot considering the nitrogen cycle, on which all life
be formed via transamination or via conversion ultimately depends (Figure 4.4). From this it can
from another carbon precursor. In this narrower be seen that some organisms are capable of fi
metabolic view, they are truly indispensable xing atmospheric nitrogen into ammonia, and
amino acids. A contem porary nutritional classifi plants are able to use either the ammonia or
cation of amino acids in human nutrition is given soluble nitrates (which are reduced to ammonia)
in Table 4.5. produced by nitrifying bacteria. However,
Strictly speaking, a truly dispensable amino vertebrates, including humans, must obtain
acid is one that can be synthesized de novo from dietary nitrogen in the form of amino acids or
other organic compounds, possibly → Glutamine + ADP + Pi + H+ (4.1) and
(2) via the glutamate dehydrogenase reaction:
α-Ketoglutarate + NH+4 + NADPH
Reduction by ∫L-Glutamate + NADP + H2O (4.2)
some anaerobic However, because Km for NH+4 in this reaction is
bacteria
high (>1 mM), this reaction is thought to make
only a modest contribution to net ammonia
assimilation in the mammal.
as urea and purine and pyrimidines. Glutamate
and glutamine provide a critical entry of the
ammonia from the nitrogen cycle into other amino
acids. It is, therefore, important to examine briefl y
the way in which the human body may obtain this Amino acids
NSN so as to maintain the nitrogen economy of and other
reduced
the individual. nitrogen–carbon
Ammonia can be introduced into amino acids compounds
by ubiquitous glutamate ammonia ligase
Synthesis in plants
(glutamine synthetase) that catalyzes the and microorganisms
following reaction:
Glutamate + NH+4 + ATP Degradation by
Nitrate NO–3 by some bacteria Ammonia NH4
animals and (eg, Klebsiella,
microorganisms Azotobacter, Rhizobium)
Denitrification Nitrogen fixation

Nitrification Nitrite NO–2 by soil bacteria

by soil bacteria (eg, Nitrobacter) Nitrification (eg, Nitrosomonas)

Figure 4.4 The nitrogen cycle. The most abundant form of nitrogen is present in air, which is four-fi fths molecular nitrogen
(N2). The total amount of nitrogen that is fi xed in the biosphere exceeds 10 11 kg annually. Reproduced from Lehninger AL,
Nelson DL, Cox MM. Principles of Biochemistry, 2nd edn. New York: Worth, 1993.

In bacteria and plant chloroplasts, glutamate is Hence, the two reactions combined (eqn 4.4)
produced by the action of glutamate synthase, give a net synthesis of one molecule of
accord ing to the reaction: glutamate. However, because glutamate
synthetase is not present in animal tissues, a net
α-Ketoglutarate + glutamine + NADPH + H+ → 2
incorporation of ammonia nitrogen via this
Glutamate + NADP (4.3) nitrogen cycle arises primarily from glutamate
The sum of the glutamate synthase (eqn 4.3) and rather than from glutamine. A net accumulation of
glutamine synthetase (eqn 4.1) reactions is, glutamine would be achieved via the glutamine
therefore: syn thetase reaction that uses ammonia, which
would be derived from various sources including
α-Ketoglutarate + NH+4 + NADPH + ATP → glutamate or other amino acids or via hydrolysis
Glutamate + NADP + ADP + Pi (4.4)
of urea by the microfl ora on the intestinal lumen.
 A net incorporation of ammonia into glycine determine the minimum E/T ratios for
might also be achieved via the glycine synthase different physiological states … . Finally, the
(glycine cleavage) reaction, as follows: question arises whether there is an optimal
pattern of nonessential amino acids.
CO2 + NH+4H+ + NAD + N5,N10-
Methylenetetrahydrofolate This statement can just as well be repeated
∫ Glycine + NAD +
+ Tetrahydrofolate (4.5) today, but clearly recent studies are beginning to
provide deeper metabolic insights into the nature
The glycine could then be incorporated into of the NSN needs of the human body.
proteins and into such compounds as glutathione,
creatine, and the porphyrins, as well as being “Conditional” indispensability
converted to serine. The nitrogen of serine would A contemporary nutritional classifi cation of amino
then either be available for cysteine (and taurine) acids in human nutrition is given in Table 4.5 and
synthesis or be released as ammonia via the some points should be made here about the
serine dehydratase reac “condi tionally” indispensable amino acids, a term
tion. However, the glycine cleavage reaction that is used to indicate that there are measurable
appears to be more important in glycine limitations to the rate at which they can be
catabolism than for its synthesis. Therefore, the synthesized. There are several important
glycine–serine pathway of ammonia incorporation determinants. First, their synthesis requires the
into the amino acid economy of the organism provision of another amino acid, either as the
would appear to have only a limited effect on a carbon donor (eg, citrulline in the case of argi
net nitrogen input into the amino acid economy of nine synthesis or serine in the case of glycine
the body. Serine can be formed from glucose via synthe sis) or as a donor of an accessory group
3-phosphoglycerate, which comes from (eg, the sulfur group of methionine for cysteine
carbohydrate metabolism, and its nitrogen synthesis). The ability of the organism to
obtained from glutamic acid synthesis via synthesize a conditionally essential amino acid is,
transamination with 2-ketoglutarate. therefore, set by the availabil ity of its amino acid
This suggests, therefore, the possibility that precursor. Second, some of these amino acids
gluta mate is a key amino acid in making net are synthesized in only a limited number of
amino tissues. The best example of this is the crucial
Nutrition and Metabolism of Proteins 57 dependence of the synthesis of proline and
arginine on intestinal metabolism. Third, most
evidence sug-
nitrogen available to the mammalian organism;
58 Introduction to Human Nutrition
this glutamate would be derived ultimately from
plant protein. In this sense, glutamate or its lower
homo logue, aspartic acid, which could supply the gests that, even in the presence of abundant
α-amino nitrogen for glutamate, or its derivative, quantities of the appropriate precursors, the
glutamine, would be required as a source of α- quantities of condi tionally essential amino acids
amino nitrogen. While additional research is that can be synthesized may be quite limited.
necessary to determine whether glutamate, or Thus, there are circumstances, for example in
one of these metabolically related amino acids, immaturity and during stress, under which the
would be the most effi cient source of α-amino metabolic demands for the amino acids rise to
nitrogen, these considerations potentially offer a values that are beyond the biosynthetic capacity
new perspective on the NSN com of the organism. This appears to be the case with
ponent of the total protein requirement. In 1965, a regard to the proline and arginine nutrition of
United Nations expert group stated: severely burned individuals, and cysteine and
The proportion of nonessential amino acid perhaps glycine in the nutrition of prematurely
nitrogen, and hence the E/T [total essential delivered infants.
or indispensable amino acids to total
nitrogen] ratio of the diet, has an obvious infl
uence on essential amino acid requirements
4.5 Biology of protein and
amino acid requirements
… . To make the best use of the available
food supplies there is an obvious need to
Body protein mass
A major and fundamental quantitative function of
the dietary α-amino acid nitrogen and of the indis
pensable amino acids is to furnish substrate
required for the support of organ protein synthesis
and the maintenance of cell and organ protein
content. Therefore, in the fi rst instance the body
protein mass is a factor that will infl uence the
total daily require ment for protein. Adult
individuals of differing size but who are otherwise
similar in age, body composi tion, gender, and
physiological state would be expected to require
proportionately differing amounts of nitrogen and
indispensable amino acids. Changes in the
distribution and amount of body protein that occur
during growth and development and later on
during aging may be considered, therefore, as an
initial approach for understanding the metabolic
basis of the dietary protein and amino acid needs.
(For more detailed considerations of body
continually synthesized and degraded in an
overall process referred to as turnover. The rate
of turnover and the balance of synthesis and
degradation of proteins, in addition to the mass of
protein, are also important determinants of the
requirements for nitrogen and amino acids, and
these aspects will be discussed in the following
section.
Turnover of proteins and amino
acid metabolism
Protein synthesis, degradation, and turnover The
principal metabolic systems responsible for the
maintenance of body protein and amino acid
homeo stasis are shown in Figure 4.5. They are:
● protein synthesis
● protein breakdown or degradation
● amino acid interconversions, transformation, and

composi tion please refer to Chapter 2.) eventually oxidation, with elimination of carbon
Direct measures of total body protein cannot dioxide and urea production
yet be made in living subjects, although there are ● amino acid synthesis, in the case of the
various indirect measures from which it is nutritionally dispensable or conditionally
possible to obtain a picture of the body nitrogen indispensable amino acids.
(protein) content at various stages of life. From
Dietary and nutritional factors determine, in
these approaches it is clear that body nitrogen
part, the dynamic status of these systems; such
increases rapidly from birth during childhood and
factors include the dietary intake levels relative to
early maturity, reaching a maximum by about the
the host's protein and amino acid requirements,
third decade. Thereafter, body nitrogen
the form and route of delivery of nutrients, ie,
decreases gradually during the later years, with
parenteral (venous) and enteral (oral) nutritional
the decline occurring more rapidly in men than in
support, and timing of intake during the day,
women. A major contributor to this age-related
especially in relation to the intake of the major
erosion of body nitrogen is the skeletal
energy-yielding substrates, which are the
musculature. Strength training during later life can
carbohydrates and fats in foods. Other factors,
attenuate or
including hormones and immune system
products, also regulate these systems. This will
be a topic for discussion in the following volume.
partially reverse this decline in the amount of Changes in the
protein in skeletal muscles and improve overall
function. The protein requirement of adults is
usually con sidered to be the continuing dietary
intake that is just suffi cient to achieve a f
“maintenance” of body nitro gen, often measured
only over relatively short experi mental periods.
For infants and growing children and pregnant
women an additional requirement is needed for
protein deposition in tissues. However, this
concept is oversimplifi ed since the chemical com
position of the body is in a dynamic state and
changes occur in the nitrogen content of
individual tissues and organs in response to
factors such as diet, hor monal balance, activity b
patterns, and disease. Thus, proteins are being
A Figure 4.5 The major systems in
amino acid uptake, utilization, and
catabolism, with an indication of the
processes involved and some factors
that can affect them. TNF, tumor necro
sis factor, IL, interleukin.

rates and effi ciencies of one or more of these

systems lead to an adjustment in whole body
nitrogen (protein) balance and retention, with the
net direc tion and the extent of the balance
depending upon the sum of the interactions
occurring among the pre vailing factor(s).
In effect, there are two endogenous nitrogen Figure 4.6 The two endogenous nitrogen cycles that
cycles that determine the status of balance in determine the status of body protein (nitrogen) balance.
(Adapted from Waterlow JC. The mysteries of nitrogen
body protein: balance. Nutr Res Rev 1999; 12: 25–54, with permission of
●the balance between intake and excretion ● the Cambridge University Press.)
balance between protein synthesis and break
down (Figure 4.6).
4–5 g protein/kg per day, respectively. Three
In the adult these two cycles operate so that they points relevant to nutritional requirements may be
are effectively in balance (nitrogen intake = drawn from these data. First, the higher rate of
nitrogen excretion and protein synthesis = protein protein syn thesis in the very young, compared
break down), but the intensity of the two cycles with that in the adult, is related not only to the fact
differs, the fl ow of nitrogen (and amino acids) that a net deposi tion of protein occurs during
being about three times greater for the protein growth, which may account for about 30% of the
synthesis/breakdown component than for nitrogen total amount of protein synthesized in the 6 month
intake/excretion cycle. old infant, but also to a high rate of protein
Protein synthesis rates are high in the turnover (synthesis and break down) associated
premature newborn, possibly about 11–14 g with tissue remodeling and repair, as well as to
protein synthesized per kilogram of body weight removal of abnormal proteins. In the adult the
per day, and these rates decline with growth and protein turnover is associated with cell and
development so that in term babies and young 60 Introduction to Human Nutrition
adults these rates are about 7 g and
Nutrition and Metabolism of Proteins 59
organ protein maintenance since there is no net
tissue growth under most circumstances. Second,
as will be seen later, at all ages in healthy subjects
the rates of whole body protein synthesis and
breakdown are considerably greater than usual
intakes (the latter are about 1–1.5 g protein/kg per
day in adults) or those

c
levels of dietary protein thought to be just within the body of the amino acids liberated during N

necessary
the course of protein breakdown. If this were not
the case it might be predicted that we would be
obligate carnivores and this, undoubtedly, would
have changed the course of human evolution.
Third, although not evident from this discussion
alone, there is a general as well as functional
relationship between the basal

to meet the body's needs for nitrogen and amino

Diet adequate
in protein

Diet low
in protein

acids (about 0.8 g protein/kg per day). It follows,

AB
Energy intake
energy metabolism or resting metabolic rate and
the rate of whole body protein turnover. Protein
synthe sis and protein degradation are energy-
requiring pro cesses, as will be described
elsewhere in these volumes, and from various
studies, including interspecies com ponents, it
can be estimated that about 15–20 kJ (4– 5 kcal)
of basal energy expenditure is expended in
association with the formation of each gram of
e
new protein synthesis and turnover. In other
words, protein and amino acid metabolism may
g

therefore, that there is an extensive reutilization

be respon sible for about 20% of total basal
energy metabolism. Because basal metabolic rate
accounts for a signifi cant proportion of total daily
energy expenditure, it should be clear from this
discussion that there are signifi cant, quantitative
interrelationships between energy and protein
metabolism and their nutritional require ments.
For these reasons it would not be diffi cult to
appreciate that both the level of dietary protein
and the level of dietary energy can infl uence the
balance between rates of protein synthesis and
protein break down and so affect body nitrogen
o
balance. Their effects are interdependent and
their interactions can be complex. This can be
r

i
illustrated by the changes in body nitrogen
balance that occur for different protein and energy
intakes (Figure 4.7); as seen here, the level of
energy intake, whether above or below require
ments, determines the degree of change in the
nitro gen balance that occurs in response to a
change in nitrogen intake. Conversely, the level of
nitrogen intake determines the quantitative effect
of energy intake on nitrogen balance. Therefore,
optimum body protein nutrition is achieved when
protein and
Figure 4.7 Relationship between nitrogen balance and energy involves a reutilization of endog enous cysteine, a
intake with diets of different protein levels. Between energy low intake of dietary methionine and cyst(e)ine
intake A (low) and B (higher) the two lines are parallel. would be expected to have an unfa vorable infl
(Reproduced from Munro HN, Allison JB, eds. Mammalian
Protein Metabolism, vol. I. New York: Academic Press, 1964:
381 with permission.)
energy intakes (from sources such as
carbohydrates and lipids) are suffi cient to meet
or balance the needs for amino acids, nitrogen,
and the daily energy expen diture or, in the case
of growth, the additional energy deposited in new
tissues.
Amino acids as precursors of
physiologically important
nitrogen compounds
As already pointed out, amino acids are also used
for the synthesis of important nitrogen-containing
com pounds that, in turn, play critical roles in cell,
organ, and system function. In carrying out these
particular roles the amino acid-derived
metabolites also turn over and they need to be
replaced ultimately by the nitrogen and
indispensable amino acids supplied by protein
intake. Estimates on the quantitative utiliza tion of
these precursor and nonproteinogenic roles of
amino acids in human subjects are limited but it is
possible to give some examples.
● Arginine is the precursor of nitric oxide (NO); the
total amount of NO synthesized (and degraded)
per day represents less than 1% of whole body
arginine fl ux and less than 1% of the daily
arginine intake.
● In contrast, the rate of synthesis and degradation
of creatinine is relatively high and accounts for
10% of the whole body fl ux of arginine and for
70% of the daily intake of arginine.
● Similarly, the synthesis and turnover of
glutathione (a major intracellular thiol and
important antioxi dant, formed from glutamate,
glycine, and cyste ine) accounts for a high rate of
cysteine utilization such that it greatly exceeds
the equivalent of the usual daily intake of
cysteine. Since continued glu tathione synthesis
uence on glutathione status and synthe sis. This
has been shown experimentally to be the case,
especially in trauma patients and those suffer ing
from acquired immunodefi ciency syndrome
(AIDS). Because glutathione is the most
important intracellular antioxidant that protects
cells against damage by reactive oxygen species,
this would mean
Nutrition and Metabolism of Proteins 61
that particular attention should be paid to such
amino acids in nutritional therapy in these
groups of patients.
Urea cycle enzymes and urea production
Finally, with reference to the major processes
shown in Figure 4.5, the urea cycle enzymes,
which are dis tributed both within the
mitochondrion and in the cytosol (Figure 4.8), are
of importance. The produc tion of urea may be
viewed largely, but not entirely, as a pathway
involved in the removal of amino nitro gen and
contributing to an adjustment of nitrogen loss to
nitrogen intake under various conditions. The fi ve
enzymes of urea biosynthesis associate as a
tightly connected metabolic pathway, called a
metabalon, for conversion of potentially toxic
ammonia as well as removal of excess amino
acids via their oxidation with transfer of the
nitrogen to arginine and ulti mately urea. This is
especially important when the supply of protein or
amino acids is high owing to
 – Mitochondrion CPS Carbamyl
+ HCO + 2ATP
+
NH 43+ 2ADP + Pi

i
N-Acetyl glutamate Acetyl CoA + Glutamate Citrulline
h

ti
phosphate OTC Ornithine Pi
n

Arginine
r

2+ +
Mg , K

Fumarate Aspartate Citrulline

Arginine Argininosuccinate

Urea
Figure 4.8 The urea cycle enzymes and their distribution in the liver. CPS, carbamoyl phosphate synthetase; OTC, ornithine
transcarbamylase; Asy, argininosuccinic synthetase; AS, argininosuccinate; Arg, arginase.
62 Introduction to Human Nutrition
variations in the exogenous intake or when there
is a high rate of whole body protein breakdown in
cata bolic states, as occurs in severe trauma and
following overwhelming infection.
Altered intakes of indispensable amino acids
and of total nitrogen result in changes in rates of
amino acid oxidation and the output of urea
nitrogen in urine. There is a roughly parallel
change in urea pro
duction and excretion throughout a relatively wide
range of change in the level of dietary nitrogen
intake above and below physiological requirement
levels. Part of this urea enters the intestinal
lumen, where there is some salvaging of urea
nitrogen, via intestinal hydrolysis of urea to form
ammonia. This ammo
AS ASy
Arg Ornithine
nium nitrogen can be made available to the host
for the net synthesis of dispensable or
conditionally indispensable amino acids.
However, the quantitative extent to which this
pathway of nitrogen fl ow serves to maintain
whole body N homeostasis and retention under
normal conditions is a matter of uncertainty. The
ammonia from urea could also enter the nitrogen
moiety of the indispensable amino acids, but this
would be essentially by an exchange mechanism
and so would not contribute to a net gain of these
amino acids in the body.
The reutilization of urea nitrogen starts from the
hydrolysis of the intact urea molecule. By
constantly infusing the [15N2]-urea tracer, the
appearance of the singly labeled [ 15N]-urea
should represent the extent of urea hydrolysis. A
24 hour constant infusion of [ 15N2]-urea revealed a
minimal amount of [15N]-urea appearance in the
plasma, and a linear relationship over a wide requirement can usefully be divided: fi rst, into
range of protein intake versus total urea those needs directly associated with protein
production and urea hydrolysis. Furthermore, the deposition, a critical issue in infants, early
possible metabolic pathways involved in the childhood nutrition, and during recovery from prior
assimila depletion due to disease or malnutrition; and,
second, into those needs associ ated with the
maintenance of body protein balance, which
accounts for almost all of the amino acid
tion of ammonia generated from urea nitrogen
requirement in the healthy adult, except for that
include (1) citrulline synthesis, (2) l-glutamate
dehy drogenase pathway in the mitochondria, and due to the turnover and loss of the various
(3) glycine synthase. The net formation of amino physiologi cally important nitrogen-containing
nitro gen from these pathways is quantitatively products, some of which were mentioned above.
minimal compared with the metabolic fl uxes of Quantifying the minimum needs for nitrogen and
these amino acids through their major pathways, for indispensable amino acids to support growth
such as protein turnover, dietary intake, and de should be relatively easy, in principle, because
novo synthesis (of the nutritionally dispensable these needs are simply the product of the rate of
amino acids only). protein nitrogen deposition and the amino acid
composition of the proteins that are deposited.
Summary of the metabolic basis for protein Here, it may be pointed out that the gross amino
and amino acid requirements It should be acid composition of whole body proteins shows
evident from this account of the underly ing essentially no difference among a variety of
aspects of the needs for α-amino nitrogen and mammals, including humans (Table 4.6). Thus, at
indispensable amino acids, that the “metabolic” the

Table 4.6 Essential amino acid composition of mixed body protein of immature mammals

Amino acid composition (mg/g protein)

Lysine Phenylalanine Methionine Histidine Valine Isoleucine Leucine Threonine

Rat 77 43 20 30 52 39 85 43 Human 72 41 20 26 47 35 75 41 Pig 75 42 20 28 52 38 72 37 Sheep 75 42 17 23 53 33

79 47 Calf 69 39 18 27 42 30 74 43

From Reeds PJ. Dispensable and indispensable amino acids for humans. J Nutr 2000; 130: 1835S–1840S. Reprinted with
permission of The American Society for Nutrition.
Nutrition and Metabolism of Proteins 63

Table 4.7 The involvement of amino acids in physiological systems and

metabolic function System Function Product Precursor

Intestine Energy generation ATP Glu, Asp, Glutamine Proliferation Nucleic acids
Glutamine, Gly, Asp
Protection Glutathione Cys, Glu, Gly
Nitric oxide Arg
Mucins Thr, Cys, Ser, Pro
Skeletal muscle Energy generation Creatine Gly, Arg, Met
Peroxidative protection Taurine (?) Cys
Nervous system Transmitter synthesis Adrenergic Phe
Serotinergic Try
Glutaminergic Glu
Glycinergic Gly
Nitric oxide Arg
Peroxidative protection Taurine (?) Cys
Immune system Lymphocyte proliferation (?) Glutamine, Arg, Asp Peroxidative
protection Glutathione Cys, Glu, Gly
 Cardiovascular Blood pressure regulation Nitric oxide Arg
Peroxidative protection (?) Red cell glutathione Cys, Glu, Gly

major biochemical level the qualitative pattern of as well as the branched-chain amino acids as
the needs of individual amino acids to support nitrogen precur sors for cerebral glutamate
protein deposition would be expected to be synthesis), the necessary precursors shown here
generally similar. are the dispensable and con ditionally
In humans, in contrast to rapidly growing indispensable amino acids.
mammals such as the rat and pig, the obligatory
amino acid needs for the purposes of net protein 4.6 Estimation of protein and
deposition are for most stages in life a relatively amino acid requirements
minor portion of the total amino acid requirement.
Hence, most of the requirement for nitrogen and Having considered the biology of protein and
amino acids is associ protein requirements, this section now considers
ated with the maintenance of body protein stores how these factors may be used to estimate the
(or body nitrogen equilibrium). A major portion of requirement for protein and for amino acids. The
the maintenance nitrogen and amino acids needs fi rst section dis
is directly associated with protein metabolism and cusses nitrogen balance and the defi nition of
refl ects two related factors. protein requirements, before discussing how
these vary with
●Amino acids released from tissue protein
64 Introduction to Human Nutrition
degrada tion are not recycled with 100% effi
ciency. ● Amino acid catabolism is a close function
of the free amino acid concentration in tissues, age and for various physiological groups.
and so the presence of the fi nite concentrations Subsequent sections cover the estimation of the
of free amino acids necessary to promote protein requirements for the indispensable amino acids.
synthesis inevi tably leads to some degree of
amino acid catabolism and irreversible loss.
Nitrogen balance and defi nition
The other metabolic component of the require of requirement
ment for nitrogen and amino acids, as mentioned The starting point for estimating total protein
above, is due to the turnover of functionally impor needs has been, in most studies, the
tant products of amino acid metabolism, which measurement of the amount of dietary nitrogen
are also necessary to maintain health. Although needed for zero nitrogen balance, or equilibrium,
this may not necessarily be a major quantitative in adults. In the growing infant and child and in
component of women during pregnancy and lactation, or when
the daily requirement, it is qualitatively and repletion is necessary following trauma and
function ally of considerable importance; health infection, for example, there will be an additional
depends on the maintenance of this component requirement associated with the net depo sition of
of the protein need. protein in new tissue and that due to secre
Finally, four physiological systems appear to be tion of milk. Thus, a United Nations (UN) Expert
critical for health: the intestine, to maintain absorp Consultation in 1985 defi ned the dietary need for
tive and protective function; the immune and protein as follows.
repair system and other aspects of defense; the
skeletal musculature system; and the central The protein requirement of an individual is
nervous system. Within each system it is possible defi ned as the lowest level of dietary protein
to identify critical metabolic roles for some specifi intake that will balance the losses from the
c amino acids (Table 4.7). Also of note is that, body in persons maintaining energy balance
with certain exceptions (the involvement of at modest levels of physical activity. In
phenylalanine and tryptophan in the maintenance children and pregnant or lactating women,
of the adrenergic and serotinergic neu the protein requirement is taken to also
rotransmitter systems, and methionine as a include the needs associated with the
methyl group donor for the synthesis of creatine, deposition of tissues or the secretion of milk
 at rates consistent with good health. including those via sweat and the integument. For
children, estimates of nitrogen deposition or
Most estimates of human protein requirements
retention are also included. In the case of very
have been obtained directly, or indirectly, from young infants the recommendations for meeting
measurements of nitrogen excretion and balance
protein requirements are usually based on
(Nitrogen balance = Nitrogen intake – Nitrogen estimated protein intakes by fully breast-fed
excretion via urine, feces, skin, and other minor infants.
routes of nitrogen loss). It must be recognized
that the nitro
gen balance technique has serious technical and Protein requirements for various age and
inter pretative limitations and so it cannot serve physiological groups
as an entirely secure or suffi cient basis for The protein requirements for young adult men
establishing the protein and amino acid needs for and women have been based on both short- and
human subjects. Thus, there are: long term nitrogen balance studies. This also
applies to healthy elderly people, whose protein
● a number of inherent sources of error in nitrogen
requirements have been judged not to be different
balance measurements that should be considered
from those of younger adults. In order to make
● a number of experimental requirements that
practical recom mendations to cover the
must
requirements for most indi viduals, it is necessary
be met if reliable nitrogen balance data are to
to adjust the average or mean requirement for a
be obtained.
group by a factor that accounts for the variation in
These include protein requirements among appar ently similar
● the need to match closely energy intake with
individuals in that group. This factor is usually
energy need, for the various reasons discussed taken to be the coeffi cient of variation (CV)
earlier around the mean requirement and traditionally a
value of 2 ⋅ CV (SD/mean) is added to the mean
physiological requirement, so that the needs of all
● an appropriate stabilization period to the experi but 2.5% of individuals within the population
mental diet and periods long enough to would be covered. This adjusted requirement
establish reliably the full response to a dietary value is taken to be the safe practical protein
change intake for the healthy adult (Table 4.8). Most
● timing and completeness of urine collections ●
individuals would require less
absence of mild infections and of other sources of
stress.
Reference to detailed reviews of the concepts Table 4.8 The United Nations (1985 FAO/WHO/UNU) and
behind and techniques involved in the nitrogen Institute of Medicine (2002/2005) recommendations for a safe
practical protein intake for selected age groups and
balance approach is given in the reading list at physiological states. Reproduced with permission from WHO
the end of this chapter.
When direct nitrogen balance determinations of Safe protein
the protein requirement data are lacking, as is the level (g/kg/day)
case for a number of age groups, an interpolation Nutrition and Metabolism of Proteins 65
of requirements between two age groups is
usually made simply on the basis of body weight
. . . . the lowest level of intake of an
considerations. A factorial approach may also be
indispensable amino acid that achieves
applied; here, the so
nitrogen balance or that balances the
called obligatory urine and fecal nitrogen losses
irreversible oxidative loss of the amino acid,
are determined (after about 4–6 days of
without requiring major changes in normal
adaptation to a protein-free diet in adults),
protein turnover and where there is energy
summated together with other obligatory losses,
balance with a modest level of
Group Age (years) physical activity. For infants, lactating women, the requirements
UNU IOM children and preg nant and for
Infants 0.3–0.5 1.47 1.5 0.75–1.0 1.15 1.1 Children 3–4 1.09 0.95 9–10 0.99 0.95
Adolescent 13–14 (girls) 0.94 0.85 13–14 (boys) 0.97 0.85
Young adults 19+ 0.75 0.80 Elderly 0.75 0.80 Women:
pregnant 2nd trimester +6 g daily ~1.1 3rd trimester +11 g
daily ~1.1
lactating 0.6 months ~+16 g daily ~1.1 6–12 months 12 g
daily ~1.1
Values are for proteins such as those of quality equal to a
hen's egg, cow's milk, meat, or fi sh.
than this intake to maintain an adequate protein
nutritional status.
It is worth emphasizing two points. First, the
current UN recommendations shown in Table 4.8
apply to healthy individuals of all ages. However,
it is highly likely that the needs of sick or less
healthy patients would differ from and usually
exceed those of healthy subjects. In this case, the
values given in this table can be regarded only as
a basis from which to begin an evaluation of how
disease and stress, including surgery, affect the
needs for dietary protein. Unfortunately, the
quantitative needs for protein (total nitrogen) in
sick, hospitalized patients can be only very
crudely approximated at this time.
Second, the values shown in Table 4.8 apply to
high-quality food proteins, such as eggs, milk,
Determination
In general, the approaches and methods that
have been most often used to determine specifi c
indis pensable amino acid requirements are
similar to those used for estimation of total protein
needs, ie, nitrogen excretion and balance and
factorial estimation. Thus, amino acid
requirements have been assessed by nitrogen
balance in adults, and by determining the
amounts needed for normal growth and nitrogen
balance in infants, preschool children, and
school-aged children. For infants, they have also
been approached by assessment of the intakes
pro
vided by breast milk or those supplied from
intakes of good-quality proteins. In addition,
factorial pre dictions of the amino acid
requirements of infants and adults have been
made. One such factorial approach for use in
adults includes the following assumptions.
● The total obligatory nitrogen losses (those losses
occurring after about 4–6 days of adjustment to
a protein-free diet) are taken to be
approximately 54 mg/kg nitrogen per day in an
adult, or equiva
lent to 0.36 g protein/kg/day.
● The average amino acid composition of body pro

meat, and fi sh. The differing nutritional value of teins can be used to estimate the contribution
food pro teins will be considered below. made

Defi nition and determination of

indispensable amino acid requirements
Defi nition
It is possible to modify slightly the earlier defi
nition for the requirements for protein (nitrogen)
for a spe cifi c, indispensable amino acid, which
can be stated, therefore, as:
the amino acid will include the additional
amount of the amino acid needed for net
protein deposition by the infant, child or fetus
and con ceptus and for the synthesis and
secretion of milk proteins.
The foregoing is an operational defi nition of
requirement, as in the case of protein. Ideally, a
func tional defi nition and determination of these
require ments inherently would be preferable.
However, the choice and nature of the functional
index or (indices) (such as maximum resistance
to disease or enhanced physical performance)
and its quantitative defi nition remain a challenge
for future nutrition and health related research.