Avian Genetic Diversity: Domesticated Species
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2 Avian genetic diversity: Domesticated species GENETIC DIVERSITY IS CONSIDERED crucial to the bird stocks (ring-necked pheasant and bobwhite continued survival of a species, be it wild or do- quail) have been developed and will be noted in mestic. Such within-species diversity has been this report. Noted below are salient features of the raw material of agriculturists over millennia. the most widely used domesticated species that In response to selective breeding and the differ- have the greatest need for conservation of ge- ential survival of less fit animals, preferred traits netic stocks. have been accentuated and clustered to produce distinct breeds and varieties of the modern do- Chicken mesticated species (NRC 1993). In more recent times, researchers have deliberately isolated First domesticated over 6,000 years ago, the various mutations in specialized stocks, permit- chicken (Gallus gallus or G. domesticus) presents ting the systematic study of such mutations and by far the greatest amount of genetic diversity of promoting a better understanding of the normal the domesticated avian species, with over 400 function of the affected genes. identified genetic variations (SOMES 1988). Many The totality of wild and domesticated species are showcased in the more-than 100 recognized form the gene pool or genetic resources base chicken breeds and commercial varieties, which necessary for the survival of the species. The variously integrate most of the naturally occur- genes and genotypes present in this pool repre- ring mutations affecting size, body type, produc- sent genetic resources which are accessible and tion characteristics, posture, color, feather can be exploited by biologists and breeders. In structure and location, comb shape, and behav- this report, we emphasize “genetic stocks” which ior (see Figures 1 and 2 for wild- and domestic- have been bred for specific traits and genes in type chickens). Some of the most extreme vari- contrast to breeds in which the individual birds ants include: the tiny, short-legged Japanese have many traits in common and can generally Bantam; the tall, aggressive Old English Game be maintained with randomly breeding popula- tions. Genetic stocks are typically selected for traits of special interest to breeders and geneti- cists. Many of them are reproductively, physi- cally, or physiologically compromised, and re- quire special care in breeding and management, even for maintenance or conservation purposes. Target species While the AGRTF recognizes the need for conser- vation of undomesticated avian species, this re- port primarily addresses the need for conserva- tion of specialty stocks of domesticated species, particularly chicken, turkey, and Japanese quail. A limited number of waterfowl (duck and Figure 1. Red Jungle Fowl rooster from UCD 001 goose) genetic stocks and semi-domestic game- (Photo courtesy of J. Clark, University of California–Davis). 5 Fowl; the light-weight Single- Box 5. Parthenogenetic turkeys comb White Leghorn hen that can lay more than 300 eggs in PERHAPS THE MOST SPECTACULAR use of tur- hatch, and grow into fully functional her first year of production; and keys in experimental biology was the males (OLSEN 1965). The existence of the large Rock-Cornish commer- study of meiosis, fertilization, and early this line has given rise to the notion embryonic development with a strain that genetic imprinting does not exist cial meat chicken, with its phe- of parthenogenetic turkeys. In the in birds, although this conclusion must nomenal rate of growth and 1960s and 1970s, M.W. Olsen of the be tentative in the absence of any for- well-fleshed carcass. These are United States Department of Agricul- mal investigation of imprinting in the all thought to share a common ture Agricultural Research Center at unique parthenogenetic stock. How- ancestor in the Red Jungle Fowl Beltsville developed a line of turkeys ever, this parthenogenetic stock exists in which an embryo would form in 30 precariously at only two research sta- (G. gallus gallus) (Figure 1) to 50% of the unfertilized eggs (par- tions in the world (the University of which is still found wild in parts thenogenesis). Most of these embryos Guelph (Ontario, Canada) and the Uni- of India and Southeast Asia die, but a small proportion of them versity of Oman). (CRAWFORD 1990; FUMIHITO et al. (about 0.5%) continue to develop, 1994), although some poultry specialists believe that several other jungle fowl species (G. sonnerati, G. lafay- ettei, and G. varius) also contributed to the an- cestral gene pool (CRAWFORD 1990). Turkey The one commercially important avian species originating in North America, the domestic tur- key of commerce, is the product of hybridization between two subspecies of turkey: the domesti- cated Meleagris gallopavo gallopavo from Central America and the wild M. g. sylvestris from the eastern United States (CRAWFORD 1990). From these hybrids, birds were selected for size, tame- ness, carcass yield, and rapid growth, resulting in several distinct breeds and varieties (Figure 3). The modern commercial or exhibition turkeys are large, slow-maturing birds with a much lower reproductive potential than chicken or Figure 3. Flock with different turkey breeds (Photo Japanese quail (at one generation per year for courtesy of F.A. Bradley, University of California–Davis). the turkey). Although this species is far less popular with researchers or hobbyists than the chicken or the Japanese quail, at least six breeds are still kept for exhibition and a few unique research stocks have been developed (Box 5), including several commercial-type long- term selected and randombred-control lines kept at Ohio State University (see survey results, Ap- pendix 2, Tables 2.1 and 2.2). Perhaps as a con- sequence of the few researchers studying the turkey, relatively few mutations (49) have been reported in the turkey compared to the chicken and Japanese quail (SOMES 1988). Japanese quail Gaining in popularity as an experimental animal in both research and education, the Japanese quail (Coturnix japonica) is a small, early matur- Figure 2. White Leghorn rooster from UCD 003 (Photo ing, highly efficient egg and meat producer. Until courtesy of J. Clark, University of California–Davis). recently, the Japanese quail was classified as a 6 subspecies of the common European quail (C. productive life of an individual bird is quite coturnix). It is now classified as a distinct species short, frequently less than one year, and, unlike because of the nonhybridization of the two in the the chicken, close inbreeding is not tolerated. wild or in captivity (CHENG and KIMURA 1990). Thus, only one moderately inbred line exists (at According to all available documentation, the the University of British Columbia). domestic Japanese quail strains used in meat and egg production (even in Europe) are descen- Duck ded from C. japonica, which is still found in small wild populations in Japan. While gaining Almost all of the 15 or so domestic duck breeds popularity as a food animal in the US, its small recognized today are descended from the wild size has limited its use as a meat- or egg-pro- mallard duck (Anas platyrynchus platyrynchus), ducing animal to specialty markets. However, the exception being the Muscovy duck (Cairina the Japanese quail has other qualities that make moschata) (LANCASTER 1990). In addition to the it ideally suited for research. Usually reaching different plumage patterns and colors, a variety sexual maturity by six weeks of age, the females of body types and behavioral traits are found often lay an egg a day for several months. The among the duck breeds, ranging from the boat- males are aggressive breeders and maintain high shaped, vocal Call ducks to the cane-shaped In- fertility even when housed with four or more dian Runner ducks. Only 22 mutations have hens. The early maturity and short incubation been described in the domestic duck, most in- interval (16 to 17 days) permit as many as five volving feather color or pattern (LANCASTER generations in a single year, in contrast to the 1990). As such, these traits have been used in slower-maturing chicken (one to two generations defining breed and variety standards, especially per year) or the even slower maturing and less among the more ornamental duck breeds, such productive turkey (one generation per year). The as the Call, Indian Runner, Crested, Cayuga, quail is sometimes called the mouse of the bird and Swedish. While such breeds are usually world, since it has become extremely popular as only kept by hobbyists, a few are important in a model species for biological research in several commercial meat production, particularly White fields, including toxicology, cell biology, nutri- Pekins, Rouens, and, in some areas, Muscovy or tion, and selective animal breeding. Although Muscovy-domestic duck hybrids. most researchers use unselected or randombred birds, over 100 mutations are known in this spe- Goose cies, including many affecting feather color and shape (Figures 4 and 5), and several causing Six recognized domestic goose breeds were de- embryo-lethal deformities (SOMES 1988; CHENG rived from the western Greylag goose of Europe and KIMURA 1990). At present, most of these mu- (Anser anser anser). Several other breeds are tant strains are only maintained at the Univer- thought to have descended from the smaller sity of British Columbia or by hobbyists. Two Swan goose of central Asia (A. cygnoides). The drawbacks with this species are that the usual African breed is believed to be derived from a Figure 4. Japanese quail silver mutation from UBC SI Figure 5. Japanese quail porcupine mutation from (Photo courtesy of K. Cheng, University of British Co- UBC PC-WB (Photo courtesy of K. Cheng, University lumbia). of British Columbia). 7 hybrid between these two species (HAWES 1990). and pheasant blood-type variants currently kept Strict herbivores, geese have a long history of at Northern Illinois University (JARVI et al.