017 Bone Korver

Bone mineralization and development: broiler
performance
D. R. KORVER1* and C. A. TORRES1,2
1
Department of Agricultural, Food & Nutritional Science, University of Alberta, Edmonton, AB,
Canada T6G 2P5.
2
Current address: Nutreco Poultry Research Centre, Ctra. Cm. 4004, Km. 10,5 45950,
Casarrubios del Monte (Toledo) Spain.
*Corresponding author: doug.korver@ualberta.ca
Introduction.
Bone development and metabolism is essential to the health and productivity of poultry.
Although most research in poultry bone metabolism is understandably focused on calcium
and phosphorus, trace elements are essential for proper bone formation and maintenance as
well. Bone formation starts in the egg at approximately 3.5 days of incubation with the
formation of a cartilage model (Osdoby and Caplan, 1981), with mineralization beginning at
approximately 10 days of incubation (Pechak et al., 1986). As the embryo develops,
ossification begins at the mid-region along the length of the developing bone. The
mineralized region begins to extend towards either end of the bone, and the cartilaginous
growth plates form a new organic framework for the linear growth of the bone. Skeletal
growth continues through the life of the broiler, and broilers are normally slaughtered well
before skeletal development is completed.
Trace elements and bone metabolism. Bone tissue consists of a cartilage matrix,
containing about 95% collagen (Knott and Bailey, 1999; Rath et al., 2000) which gives the
bone tensile strength (Rath et al., 2000; Williams et al., 2004; Saito and Marumo, 2010), and
mineral, most of which is in the form of hydroxyapatite (Ca10(PO4)6(OH)2; Rath et al., 2000).
The main role of trace elements in bone formation is the formation of the organic matrix
which forms a scaffold which is subsequently mineralized.
A number of trace elements play important roles in bone metabolism, but copper, zinc and
manganese are among the most important. The copper-dependent enzyme lysyl oxidase is
involved in the cross-linking of elastin and collagen in the organic matrix of bone (Marturano
et al., 2014), which gives tensile strength and elasticity to bones. Manganese is a cofactor for
polymerase and galactotransferase, which are involved in the biosynthesis of chondroitin
sulphate (Leach et al., 1969), which is a major component of the bone hyaline cartilage
Seminario Internacional de Manejo y Sistemas Operativos en Pollo de Engorde - AMEVEA, Bogot D.C. Junio 17-19, 2014
International Seminar in Management and Operative Systems in Broilers AMEVEA, Bogot D.C. June 17-19, 2014
structure (Eyre, 2004). A deficiency in Mn reduced chick bone size (Caskey et al., 1939),
likely due to reduced chondroitin sulphate content of the bone organic matrix (Leach et al.,
1969). Zinc is a cofactor for collagenase (Starcher et al., 1980), which cleaves pro-collagen
into collagen, and for bone alkaline phosphatase (Seo et al., 2010), which releases phosphorus
from phosphates at the site of bone calcification for formation of hydroxyapatite.
Additionally, zinc stimulates osteoblast proliferation and osteoprotegerin activity (Liang et
al., 2012). Zinc also influences gene transcription at the growth plate during long bone
growth (Oviedo-Rondn et al., 2006). Zinc deficiency reduced collagenase activity and
subsequently collagen synthesis in chicks (Starcher et al., 1980). Other trace elements
involved in bone metabolism include fluorine, iron, selenium and boron (Zofkova et al.,
2013).
Dietary trace element supplementation. The nutritional requirements of poultry for each
of the main trace elements involved in bone metabolism are quite well understood, and little
refinement to the recommended feeding levels has been made in recent years. Of much
greater interest to poultry nutritionists is the role of chelated trace elements (trace elements
chelated to specific organic ligands; organic trace elements). The basic rationale for feeding
organic trace elements is the protection of the trace element atom from dietary compounds
that might bind and reduce digestibility or absorption by the gut (Yu et al., 2010). In general,
commercially-available organic trace element products are intended to have increased
bioavailability relative to the more traditional inorganic element supplements such as sulfates
or oxides (Guo et al., 2001; Bao et al., 2007; Huang et al., 2009). The increased
bioavailability of organic trace elements may be useful in situations where the diet contains
factors which might bind the mineral and reduce digestibility, such as phytate (Tahir et al.,
2012). Increasing trace element bioavailability through the use of organic trace elements may
to increase bone mineralization in broilers when supplemented in the chicks diet (Kidd et al.,
1992).
Maternal trace element nutrition. Genetic selection of broilers for more rapid
growth rate has come almost exclusively as a result of post-hatch growth; the incubation
period for rapidly-growing commercial broilers and unselected lines has remained relatively
constant. Since bone development begins in ovo, maternal transfer of trace elements, and the
influence of trace element form in the hen diet has received increased research attention
recently (Hudson et al., 2004; Sun et al., 2012; Torres 2012). Because rapid post-hatch
growth can compromise bone integrity (Williams et al., 2004), increasing the development of
bone during incubation may prove to be an important means of maintaining skeletal health in
fast-growing broilers.
Bone development and mineralization. The organic matrix of the bone initially forms a
bone-shaped structure in the developing embryo, which is progressively mineralized, starting
at the metaphysis. After hatch, the mineralization continues. The growth plates at either end
of long bones deposit new organic matrix, resulting in long bone growth. Advancing behind
the growth plate is the calcifying zone, at which the newly deposited organic matrix is
mineralized by the deposition of bone mineral, primarily hydroxyapatite (Ca10(PO4)6(OH)2;
Rath et al., 2000). Bone mineralization occurs by the deposition of new mineral obtained
from the diet, but also by resorption of previously deposited bone. The larger diameter of the
epiphyses therefore are a result of re-distribution of tissue as the bone grows in length, rather
than the epiphyses being pushed out by growth occurring at the diaphysis.
Image courtesy of Creative Commons

Bones not only grow in length, but also width as the broiler ages (appositional growth).
Osteoblasts on the outer surface of the bone (periosteum) form new bone, while existing bone
from the inner surface (endosteum) is mobilized by osteoclasts. In this way, the bones do not
become excessively thick and heavy as the bird ages. Osteons (Haversian canals) are
channels within the bone that are the basic structure of bone growth. The osteons generally
are parallel to the length of the bone, and form the compact bone. Osteons start off as tubes,
which are gradually mineralized on the inner surface by osteoblasts, leaving only a narrow
channel in mature bone. Because broiler growth rate is so rapid, the osteons do not fill in
completely, leaving a porous bone structure that is not fully mineralized by processing age in
broilers. More rapid growth of broilers leads to greater porosity (Williams et al., 2000;
Leterrier and Nys, 1992; Leterrier and Constantin, 1999.
Effects of bone defects on performance. Rapid growth of broilers can lead to skeletal
defects including tibial dischondroplasia (Havenstein et al., 1994; Farquharson and Jeffries,
2000), angular bone deformity (Classen and Riddell, 1989; Yu and Robinson, 1992),
spondylolisthesis (kinky back; Julian, 2004), bacterial chondronecrosis with osteomyelitis
(femoral head necrosis; McNamee et al., 1999) . Within the same population, birds in the top
quartile growth had lower measures of bone quality (corrected for 6-week body weight) than
birds in the lowest quartile for growth (Shim et al., 2012). Before weight correction, the fastgrowing broilers had lower tibia ash than slow-growing broilers. Therefore, although larger
broilers tend to have larger and stronger bones than smaller broilers, this is accomplished by
increasing the diameter of the bones, rather than increasing mineralization. The ability of
rapidly-growing broilers to deposit bone is limited, and strength is increased by increasing
bone diameter, at the expense of creating porous bones.
Rapid growth rate of broiler chickens has, in the past, led to reductions in skeletal quality
(reviewed by Bessei, 2006). Skeletal defects can lead to pain (Caplen et al., 2013) and
therefore reductions in bird welfare (Bradshaw et al., 2002). Lameness associated with
skeletal defects can prevent locomotion, and birds may not be able to reach feed and water,
resulting culling or death. Primary breeders have made skeletal health a priority in their
selection programs, in addition to growth, carcass conformation and efficiency (Kapell et al.,
2012). Anecdotal evidence suggests that the incidence of most skeletal deformities in broilers
has decreased since the mid-1990s, even as growth rate and breast meat yield continues to
increase as a result of genetic selection.
Conclusion. The formation of a sound skeletal system is dependent upon the proper
formation and maintenance of the organic component of bones, and the subsequent
mineralization of the cartilage matrix. Both the embryonic cartilage model and the cartilage
growth plate are each dependent on the activity of enzymes containing trace elements or
requiring these elements as co-factors. Improvements in nutrition and selection for skeletal
development as well as growth have reduced skeletal defects since the 1990s. Nevertheless, as
modern broilers are increasingly selected for increased growth rate, all aspects of bone
metabolism must be fully understood to allow for continued bird health and productivity.
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Bone mineralization and development: broiler

Image courtesy of Creative Commons

JULIAN, R. J. (2004) Evaluating the impact of metabolic disorders on the welfare of

OVIEDO-RONDN, E.O., FERKET, P.R. and HAVESTEIN, G.B. (2006) Understanding

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