FITOHORMON

Fajar R. D. N. Sianipar, S.Si, M.Sc. Fisiologi Tumbuhan – Jurusan Biologi. Universitas Papua
 • Any condition that affects plant growth is called a stimulus.
What do plants
• What are three types of stimuli that plants respond to?
respond to? • Which stimulus will have the strongest effect on the plant?

light water gravity

 (2) How does a seed
become a seedling?
(1) How is a seed
formed?
Perkembangan
Fruiting
Seedling
Tumbuhan
Plant Life Cycle
Faktor internal = hormon
(3) How is
shoot formed?
Faktor eksternal =
cahaya/suhu/panjang
hari/gravitasi (4) How is root
(5) How is a flower
formed?
formed?
 • Karya Yunani “hormon” = to set in motion (untuk
menggerakkan)
• Molekul sinyal yang dihasilkan di lokasi tertentu.
Fitohormon • Aktif dalam konsentrasi rendah.
• Menyebabkan proses diubah dalam sel target di lokasi
lain.
• Secara umum dalam biokimia, hormon berada di dalam
sel dengan konsentrasi yang berbeda-beda.
• Fitohormon mengontrol aktivitas seluler (pembelahan,
pemanjangan dan diferensiasi) dan proses-proses lain
termasuk pembetukan pola, penentuan jenis kelamin,
organogenesis, serta tanggapan terhadap cekaman
biotik maupun abiotik.
 Classes of Hormones

Sumber: Wani et al. 2016. Phytohormones and their metabolic engineering for abiotic stress tolerance in crop plants. The cropjournal 162-176
Tabel 1. Gambaran umum Hormon Tumbuhan
Hormon Tempat Hormon Diproduksi/
Ditemukan pada Tumbuhan
Auksin Meristem apikal tunas, daun-daun
muda dan embrio pada biji
Sitokinin Sitokinin disintesis di akar dan
ditranspor ke organ-organ lain.
Giberelin Meristem kuncup apikal, daun
muda, dan embrio
Brasinosteroid Senyawa-senyawa ini terdapat
disemua jaringan tumbuhan
Fungsi Utama
Merangsang pemanjangan batang (hanya dalam konsentrasi rendah); mendorong
pembentukan akar-akar lateral dan adventisia; meregulasi perkembangan buah;
meningkatkan dominansi apikal; berfungsi dalam fototropisme dan gravitropisme;
mendorong diverensiasi vaskuler, memperlambat absisi daun.
Meregulasi pembelahan sel pada tunas dan akar; memodifikasi dominansi apikal
dan mendorong pertumbuhan kuncup lateral; merangsang perkecambahan biji;
menunda senensia daun.
Merangsang pemanjangan batang; perkembangan polen; pertumbuhan buah,
serta perkembangan dan perkecambahan biji.
Mendorong ekspansi sel dan pembelahan sel pada tunas; mendorong
pertumbuhan akar dalam konsentrasi rendah, menghambat apertumbuhan akar
dalam konsentrasi tinggi; mendorong diferensiasi xilem dan menghambat
diferensiasi floem; mendorong perkecambahan biji dan pemanjangan tabung
polen.
Hormon Tempat Hormon Diproduksi/ Fungsi Utama
Ditemukan pada Tumbuhan

Asam absisat Akar, batang, daun dan buah muda Menghambat pertumbuhan; mendorong penutupan stomata selama stres
kekeringan; mendorong dormansi biji dan menghambat perkecambahan dini;
mendorong senensia daun.

Etilen Hormon diproduksi oleh hampir Mendorong pematangan buah, absisi daun dan respon rangkap tiga pada semaian
semua bagian tumbuhan. (menghambat pemanjangan batang, mendorong ekspansi lateral dan
pertumbuhan horosontal); meningkatkan laju senesensia; mendorong
pembentukan akar dan rambut akar.

Strigolactone sinyal ekstraseluler diproduksi di Mendorong perkecambahan biji, mengontrol dominasi apikal, dan adanya daya
akar. tarik untuk jamur mikoriza pada akar.

Sumber:
 • Charles Darwin mempelajari mekanisme kecambah yang tumbuh ke arah
datangnya cahaya, yang merupakan efek langsung aksi auksin.
• Darwin menyimpulkan bahwa ada sinyal yang bergerak dalam tumbuhan dan
1 mengontrol pertumbuhan.

Auksin
• Perbedaan kecepatan pertumbuhan sel merupakan akibat auksin yang pindah
ke bagian yang ternaungi.

Akumulasi
Auksin pada
Panjang bagian yang
sel ternaungi
Sumber : menstimulasi
Coleoptile drawing from Darwin, C., and Darwin, F. (1881) The power of pemanjangan
movement in plants. Available online.
Esmon, C.A. et al. (2006) A gradient of auxin and auxin-dependent Konsentrasi sel dan
transcription precedes tropic growth responses. Proc. Natl. Acad. Sci. Auksin pembelokan
USA 103: 236–241. Friml, J., et al. (2002) Lateral relocation of auxin arah tumbuhan
efflux regulator PIN3 mediates tropism in Arabidopsis. Nature 415:
806-809.
Biosintesis Auksin

IAA diproduksi dari

triptofan (Trp) melalui
beberapa jalur semi- v
dependent dan satu jalur
Trp-independent .

Faktor lingkungan dan fase

perkembangan
menentukan gen-gen yang
mengontrol sintesis auksin,
konjugasi dan degradasi
yang mengakibatkan
adanya homeostasis
auksin.
Mano, Y. et al (2010) The AMI1 gene family: indole-3-acetamide hydrolase functions in auxin biosynthesis in plants J Exp Bot 61:
25-32, by permission of Oxford University Press.
Auxin: Apical dominance

http://users.rcn.com/jkimball.ma.ultranet/BiologyPages/A/Auxin.html
 • Sitokinin ditemukan dalam penelitian identifikasi senyawa yang meningkatkan
pertumbuhan sel tanaman.

2 Control With kinetin

In the 1950s, Folke
Skoog’s research group
identified a synthetic
Sitokinin cytokinin, kinetin.

• Sitokinin merupakan sekelompok senyawa yang mirip adenin

• Senyawa adenin yang terkait N6- diganti/disubtitusikan.

Isopentenyl adenine trans-zeatin dihydrozeatin cis-zeatin

Sumber :
Amasino, R. (2005) 1955: Kinetin Arrives. The 50th Anniversary of a
newplant hormone. Plant Physiol. 138: 1177 – 1184.

Hirose, N., Takei, K., Kuroha, T., Kamada-Nobusada, T., Hayashi, H., and
Sakakibara, H. (2008). Regulation of cytokinin biosynthesis,
compartmentalization and translocation. J. Exp. Bot. 59: 75–83.
 iPRMP
isopentenyladenine (iP)
Biosintesis sitokinin riboside phosphate

tZDP/ tZTP
transZeatin (tZ)
Regulated by Isopentenyl-
riboside
phosphate
CK and transferase
nitrogen Tissue specific; auxin,
CK and ABA sensitive
Biosintesis dan inaktivasi
CK sangat diatur oleh CK,
Meristem hormon lain dan faktor
specific
dimethylallyl- lingkungan.
diphosphate

Inactive form

CKX
Upregulated by CK
and ABA

Hirose, N., Takei, K., Kuroha, T., Kamada-Nobusada, T., Hayashi, H., and Sakakibara, H. (2008). Regulation of cytokinin biosynthesis, compartmentalization and translocation. J. Exp. Bot. 59: 75–83.
Sitokinin beraksi secara antagonis dengan auksin

CK Auxin
Promote
Promote stem cell Promote lateral branching in the
fate at the shoot organ initiation at the shoot
apical meristem shoot apical Inhibit branching
meristem in the shoot

Inhibit branching
Promote Maintain stem cell Promote in the shoot
differentiation at the fate at the root apical branching in
root apical meristem meristem the root

Reprinted by permission from Macmillan Publishers, Ltd: NATURE Wolters, H., and Jürgens, G. (2009). Survival of the
flexible: Hormonal growth control and adaptation in plant development. Nat. Rev. Genet. 10: 305–317. Copyright 2009.
Auksin dan sitokinin saling
mengatur fungsinya pada
ujung akar Cytokinin
(Cell differentiation)
Cell
Auxin transport Cytokinin
differentiation and response biosynthesis
In the root meristem, cytokinin
promotes cell differentiation, Cell division
whereas auxin sustains root Auxin
meristem activity by promoting (Cell division)
cell division.

Adapted from Dello Ioio, R., et al. (2008). A genetic framework for the control of cell division and differentiation in the root
meristem. Science 322: 1380 - 1384.
Auksin, sitokinin dan strigolactones mengontrol percabangan

Cabang-cabang akar atau

akar lateral dipacu oleh
auksin tetapi dihambat
oleh sitokinin

Percabangan mengontrol
semua aspek proruktivitas
tanaman dari penyerapan
Percabangan pada tunas nutrient sampai ke hasil
pucuk dipacu oleh sitokinin tanaman
dan strigolakton tetapi
dihambat oleh auksin

Coleus shoot image by Judy Jernstedt, BSA ; lateral root image from Casimiro, I., et al. (2001) Auxin
transport promotes Arabidopsis lateral root initiation. Plant Cell 13: 843-852.
Sitokinin mempengaruhi produksi padi dan berperan pada mekanisme toleransi
cekaman kering

Tanaman padi yang

mengakumulasi CK dapat
memproduksi lebih banyak
bulir padi karena adanya
perubanhan asitektur malai

Wild-type Elevated CK
Tanaman tembakau yang memproduksi
lebih banyak CK lebih toleran kekeringan
karena adanya penundaan penuaan
daun.

Ashikari, M. et al. (2005) Cytokinin oxidase regulates rice grain production. Science 309: 741 – 745, with permission from AAAS; Rivero, R. M. et al.
(2007) PNAS 104: 19631-19636.
 Sitokinin menunda
penuaan daun

Senescence is dramatically
reduced in kinetin-treated
leaves or leaves of plants
producing higher levels of
cytokinins. Plants
overproducing
Control plants
cytokinin

Smart, C.M., Scofield, S.R., Bevan, M.W., and Dyer, T.A. (1991). Delayed leaf senescence in tobacco plants transformed with tmr, a gene for cytokinin production
in Agrobacterium. Plant Cell 3: 647-656.
 • Strigolactones, disintesis dari karotenoid, diproduksi pada akar.
Strigolakton akan menjadi attractan bagi mikoriza dan memacu

3 perkecambahan tanaman parasit dari genus Striga.

Strigolactone

Sumber :
Reprinted from Tsuchiya, Y., and McCourt, P. (2009). Strigolactones: A
new hormone with a past. Curr. Opin. Plant Biol. 12: 556–561 with
permission from Elsevier.
 Biosintesis Strigolactone Strigolactones menghambat
pertumbuhan cabang
WT Mutant
Auxin yang ditransport
dari pucuk ke akar akan
Apex menginduksi sintesis
strigolactone, yang
menghambat
Bud
pertumbuhan tunas
samping.

Pada tanaman padi

mutan yang tidak
memproduksi
strigolactones,
Auxin Strigolactone tillers (tunas
samping/anakan)
tumbuh seperti
gambar disamping.

Lin, H., et al. (2009) DWARF27, an iron-containing protein required for the
Reprinted from Tsuchiya, Y., and McCourt, P. (2009). Strigolactones: A new
biosynthesis of strigolactones, regulates rice tiller bud outgrowth. Plant Cell 21:
hormone with a past. Curr. Opin. Plant Biol. 12: 556–561 with permission from
1512-1525.
Elsevier.
 • Giberelin terdiri dari sekelompok senyawa yang mirip

4 GA4 merupakan GA
aktif pada Arabidopsis

Giberelin Hanya beberapa GA yang

memiliki aktivitas biologis.
GA aktif yang utama seperti
tertera pada gambar
disamping.

• Giberelin didefinisikan sebagai penghambat dari penghambat.

A car doesn’t roll down
a hill when the brake is
on to inhibit it.
Releasing the brake
“inhibits the inhibitor”,
allowing another force
(gravity) to move it.

GA doesn’t promote
growth on its own, but
Sumber :
Reprinted from Tsuchiya, Y., and McCourt, P. (2009). Strigolactones: A instead inhibits growth
new hormone with a past. Curr. Opin. Plant Biol. 12: 556–561 with inhibitors (i.e. releases
permission from Elsevier. the brakes).
 Perception and signaling

DELLAs GA

Tumbuhan tidak Della

Protein
dapat dirombak Sel tambah
penghambat
memanjangkan oleh panjang
Giberelin
sel Giberelin
 GA responses

Biosynthesis mutant Response mutant Response mutant

No GA signaling Constitutive GA signaling

Harberd, N.P., Belfield, E., and Yasumura, Y. (2009). The angiosperm gibberellin-GID1-DELLA growth regulatory mechanism: How an
“inhibitor of an inhibitor" enables flexible response to fluctuating environments. Plant Cell 21: 1328-1339.
Biosintesis giberelin

Active

Inactivation

Olszewski, N., Sun, T.p., and Gubler, F. (2002) Gibberellin Signaling: Biosynthesis, Catabolism, and Response Pathways Plant Cell 14: S61-S80
 The manipulation of GA levels is tremendously
important for agriculture
One of the most significant
accomplishments of 20th century
science was the development of semi- - GA
dwarf grain varieties which are
deficient in GA synthesis or response.

+ GA

Fewer flowers and Delayed harvesting Increased fruit size

larger fruits

GAs are commercially used for increased fruit

size in table grapes and to regulate citrus
flowering and rind maturation
Distinguished plant breeder and
Nobel Laureate Norman Borlaug
1914-2009
 • Etilen berupa gas.
• Perlakuan etilen pada kecambah menimbulkan “triple response”
• hipokotil pendek
5 • hipokotil menebal
• hipokotil membengkok seperti kait.
Air C2H4

Etilen

Increasing ethylene Pea

• Respirasi dan produksi etilen meningkat seiring dengan pemasakan buah

Sumber :
Neljubov, D.N. (1901) Uber die horizontale Nutation der Stengel von Pisum
sativum und einiger anderen Pflanzen. Beih. Bot. Centralbh. 10: 129–139.
 Respon terhadap etilen
Air (control) 7 days ethylene pada Arabidopsis

Ethylene-induced gene expression

Cotton plants

Ethylene memacu senescence

Inhibition of leaf
daun cell expansion
Acceleration of leaf senescence

Lorenzo, O., Piqueras, R., Sanchez-Serrano, J.J., and Solano, R. (2003). ETHYLENE
RESPONSE FACTOR1 integrates signals from ethylene and jasmonate pathways in plant
defense. Plant Cell 15: 165-178; Rüžička, K., Ljung, K., Vanneste, S., Podhorská, R., Beeckman,
T., Friml, J., and Benková, E. (2007). Ethylene regulates root growth through effects on auxin
biosynthesis and transport-dependent auxin eistribution. Plant Cell 19: 2197-2212.
Ethylene promotes petal senescence

Azad, A.K., Ishikawa, T., Ishikawa, T., Sawa, Y., and Shibata, H. (2008). Intracellular energy depletion triggers programmed cell
death during petal senescence in tulip. J. Exp. Bot. 59: 2085-2095, by permission of Oxford University Press.
 Sintesis Etilen

ACC ACC
synthase oxidase

SAM ACC Ethylene

(1-aminocyclopropane-1-
S-adenosyl methionine
carboxylic acid)
 • Asam absisat mengontrol beberapa proses termasuk tanggapan
terhadap cekaman, perkembangan dan reproduksi.

6 Seed quality
Dormancy Germination

Asam absisat

Abscisic Acid
Development Biotic stress
response

Gene expression Stomatal aperture

Stress tolerance
Interaksi hormonal selama pengaturan dormansi
perkecambahan pada biji Nicotiana
 ABA disintesis di dalam plastida dan
sitoplasma serta merupakan derivat dari
zeaxanthin (pigmen tumbuhan).

Zeaxanthin

Zeaxanthin berlimpah dalam

jaringan hijau tetapi terbatas
pada tempat sintesisnya (Akar).
ABA
ABA2

Reprinted from Nambara, E., and Marion-Pol, A. (2003) ABA action and interactions in seeds.
Trends Plant Sci. 8: 213-217 with permission from Elsevier.
 • Discovered as a growth stimulator from pollen extracted from rape
plant (Brassica napus L.) -> Mitchell et al. 1970 -> called substances
Brassins.
7
Brasinosteroid
• Merupakan sekelompok senyawa steroids (ada lebih dari 60 macam).
• Brassinolide and castasterone are steroids like some animal hormones.
Plant Mammal

Reprinted with permission from from Mandava, N., Kozempel, M.,

Worley, J.F., Matthees, D., Warthen, J.D., Jacobson, M., Steffens,
G.L., Kenney, H., and Grove, M.D. (1978). Isolation of brassins by Insect
extraction of rape (Brassica napus L.) pollen. Ind. Eng. Chem. Prod.
Res. Dev. 17: 351-354, copyright 1978 American Chemical Society.
Control Gibberellin Brassinosteroid

In 1970 Mitchell
and colleagues
showed that their
newly purified
compound
promotes stem
elongation.

Reprinted by permission from Macmillan Publishers Limited, Nature: Mitchell, J.W.,

Mandava, N., Worley, J.F., Plimmer, J.R., and Smith, M.V. (1970). Brassins - a new
family of plant hormones from rape pollen. Nature 225: 1065-1066, copyright 1970.
Brassinosteroids are necessary for stamen
and pollen development

Filament Pollen Pollen tube

elongation development elongation

Image credits: Safro; Graham Matthews; Szumlanski, A.L., and Nielsen, E. (2009). The Rab GTPase RabA4d
regulates pollen tube tip growth in Arabidopsis thaliana. Plant Cell 21: 526-544.
BRs contribute to fruit ripening
Endogenous BR levels rise when
berries ripen

Exogenous BR application
hastens ripening. Application
of a BR-synthesis inhibitor
slows ripening.

Symons, G.M., Davies, C., Shavrukov, Y., Dry, I.B., Reid, J.B., and Thomas, M.R. (2006). Grapes on steroids.
Brassinosteroids are involved in grape berry ripening. Plant Physiology 140: 150-158.
Simplified pathway of BL
biosynthesis

Single reactions
Multiple reactions

37
 • Jasmonates dan salicylates merupakan hormon yang berpartisipasi
dalam respon tanaman terhadap cekaman biotik

8 European corn borer

Ostrinia nubilalis
in its host Zea mays

Jasmonates
CH2COOCH3
• Like other esters, methyl jasmonate smells good – it is the dominant
scent from jasmine flowers
O

Images courtesy Clemson University - USDA Cooperative Extension Slide Series,

Bugwood.org; Charles Averre, North Carolina State University, Bugwood.org
Demole, E. Lederer, E., and Mercier, D. (1962) Isolement et détermination de la structure du
jasmonate de méthyle, constituant odorant charactéristique de l’èssence de jasmin. Helv.
Chim. Acta 45: 675 -685.
 Jasmonates disintesis pada bunga dan sebagai tanggapan pertahanan tumbuhan
terhadap cekaman biotik

Wounding, pathogens or herbivores (or

molecules derived from them) trigger
very rapid accumulation of jasmonates

JA

MeJA

Untreated control

Gundlach, H., Müller, M.J., Kutchan, T.M., and Zenk, M.H. (1992). Jasmonic acid is a signal transducer in elicitor-induced plant cell cultures. Proc. Natl.
Acad. Sci. 89: 2389-2393.
 Plants respond to oral secretions as well as
mechanical damage

Proteinase inhibitor gene induction

is much faster when insect saliva or
reguritant is present
Plants recognize insect-derived
compounds as well as mechanical
damage

Korth, K.L., and Dixon, R.A. (1997). Evidence for chewing insect-specific molecular events distinct from a general wound response in leaves. Plant Physiol. 115: 1299-1305.
Herbivore-induced volatiles are recognized by
carnivorous and parasitoid insects

Tim Haye, Universität Kiel, Germany Bugwood.org; R.J. Reynolds Tobacco Company Slide Set and R.J. Reynolds Tobacco Company, Bugworld.org
JA biosynthesis
Cytoplasm

JAR1
conjugation

JA-ILE

JA conjugated to
isoleucine (JA-ILE) is the
active compound.

From Acosta, I., et al. (2009) tasselseed1 is a lipoxygenase affecting jasmonic acid signaling in sex determination of maize. Science 323: 262 – 265. Reprinted
with permission from AAAS.
 • Named for white willow Salix alba

9
• History of salicylate research
Salicylic Acid Untreate In 1990 SA was
Hippocrates In 1987
d side shown to be an
wrote about the endogenous
use of willow to SA was endogenous
relieve pain ~ shown to be signal in
2400 years ago responsible defense
SA responses
treated for heat
production in
In 1979 White Arum lily
showed that pre- flowers
treatment of a leaf
with aspirin or SA
conferred
resistance to
National Library of Medicine; Raskin, I.
(1992). Salicylate, a new plant hormone. Plant tobacco mosaic
Physiol. 99: 799-803. virus (TMV)
 MeSA accumulates in response to pathogens

LEAF 3 MeSA
SA

CH3

SA MeSA 3

Methyl Leaf 3 was

salicylate
inoculated with
tobacco mosaic
virus at time 0 Levels of MeSA increase
dramatically upon pathogen
infection

Seskar, M., Shulaev, V., and Raskin, I. (1998). Endogenous methyl salicylate in pathogen-inoculated tobacco plants. Plant Physiol. 116: 387-392.; Shulaev, V., Leon,
J., and Raskin, I. (1995). Is salicylic acid a translocated signal of systemic acquired resistance in tobacco? Plant Cell 7: 1691-1701.
 Most SA is synthesized from chorismic acid

Isochorismate synthase (ICS)

converts chorismic acid to
isochorismic acid. The enzyme that
converts isochormic acid to salicylic
acid hasn’t been identified in plants.

Flow through the shikimate pathway

increases with stress, stimulating SA
production. A small amount of SA is
derived from cinnamic acid.

Reprinted from Métraux, J.-P. (2002). Recent breakthroughs in the study of salicylic acid biosynthesis. Trends in Plant Science 7: 332-334 with
permission from Elsevier.