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      Broiler genetics and the future outlook

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      Frontiers in Physiology
      Frontiers Media S.A.
      broiler, chicken, genetics, breeding, history

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          Abstract

          While preparing this essay, the quote by Abner Kovner kept recurring. “To Remember the Past. To Live the Present. To Trust the Future” Hanging on the wall of my study is a plaque, dated 1948, that reads “The Chicken-of-Tomorrow Committee presents this Certificate of Quality to Paul Siegel for outstanding achievement in breeding and development of superior meat-type chickens.” That was 75 years ago, and it is only during the past 100 years that the production of chickens for human meat consumption was no longer a by-product of the commercial egg industry. This comment may be surprising unless we recognize that the domestication of the chicken from its wild ancestry is recent in the context of human history (Smith and Daniel, 1975). Moreover, among domesticated farm animals, the chicken increased in size while mammals became smaller (Diamond, 1995). In an evolutionary context, the domestication of the chicken had not been great, as Jungle Fowl cross fully with domestic chickens (Sutherland et al., 2018). There is a wealth of literature on the domestication of the chicken for religious, cultural, and sport reasons. Its origins and roles as a food source too was beautifully discussed essentially a century ago in the National Geographic magazine (Jull, 1927, 1930; Lewis, 1927). For broilers, examples of anthologies include Gordy (1974), Watts (1996), Cahaner and Siegel (1986), and Siegel (2014, 2018). These publications and others reveal that it is only during the past 100 years that the broiler ceased to be a by-product of the commercial egg industry, fostered by Cecile Steele, with a subsequent focus on meat (broilers and broiler genetics). Initially, the process of producing broilers via broiler genetics involved the development of the brown egg “dual purpose” chicken. Males were still reared for meat and females for egg production. It was post World War II when the “Chicken-of-Tomorrow” program (Gordy, 1974) provided the impetus for the development of breeding programs explicitly genetically designed for the production of a commercial meat-type (broiler) chicken. The initial stocks, which consisted of line crosses, were distinct from that of dual purpose chickens. Thus, although the chicken was domesticated during Neolithic times, the development of genetic programs designed for broiler performance (meat) was a 20th century event. The rapid development of a broiler per se was based on available stocks and sound breeding principles based on development of qualitative and quantitative genetics, which were first demonstrated in animals early in the first decade of the 20th century [e.g., Bateson and Punnett, 1959 (1905–1908)]. Broiler genetics, although conceptually new, was founded on a solid biological background. The plethora of literature on the reduction in time and feed required to produce today’s “broiler” is a story well documented and beyond the scope of this essay. Yet, it is instructive to review the numerous traits that favored domestication of the chicken. Although some of these are no longer relevant in current broiler production due to human intervention, they are necessary to our understanding of why domestication of the chicken was not complicated. They were small and did not migrate, there were social groupings of males and females, and they possessed behavior traits such as promiscuity and broodiness. Precocial young, with well-developed motor ability and auditory and tactile senses, contributed to an adaptation to a range of environments (Hale, 1989). The advent of electricity facilitated further human intervention on a larger scale via artificial incubation and brooding, which provided humans with tools to manipulate the photoperiod and thus maintain persistent egg production. The gasoline engine and railroad for transportation allowed for more interactions among geneticists, facilitating exchange of ideas. These, plus the emergence of vaccines and understanding of nutritional requirements for growth and reproduction, allowed for year round production and marketing of broilers. Thus, broiler genetics was becoming a specialty area per se. As stated previously, with the rediscovery of Mendelism at the turn of the 20th century, the chicken, because of traits described earlier, became a model animal for genetic research. This fundamental knowledge led to an understanding for the development and application of breeding programs for meat traits that were quantitatively inherited. Broiler breeders had a fountain for broiler genetics research from publicly supported research as its basis, as well as a range of stocks developed by fanciers, many developed before the rediscovery of Mendelism. These, plus an appreciation for quantitatively inherited traits, genotype-environment interactions, genetic correlations, heterosis, and the concept of resource allocations facilitated development of the broiler per se, not as a spin-off from the genetics of egg production. Expansion of mass transportation and development of computer technology contributed to specialized breeding programs that capitalized on a short generation interval with mini-generations. The short generation interval (which is often overlooked), plus a moderate to high heritability for body weight, facilitated reduction in broilers reaching market weights at younger ages, which also improved feed efficiency. These are items that should not be ignored when discussing broiler genetics and improvements in broiler performance during the last 70 years. Husbandry practices and high energy diets were contributing factors, but they were secondary to the dynamics of selection and crossing of specific male and female lines, i.e., breeding and genetics were the primary contributors (e.g., Havenstein et al., 2003). The financial investment was considerable, and thus it was essential for broiler breeders to have control of their parental lines. Basically, they were utilizing Mendelian genetics per segregation and recombination to protect their investments. Thus, while broiler genetics did not precede the founding of the broiler industry, without the genetic paradigm, the global industry would not be where it is today. Development of sophisticated breeding programs capitalized on the availability of science and technology. As stated previously, during the early phases of commercial broiler breeding, there was reliance on readily available science and technology and a broad gene pool. With a short generation interval, capital investments were necessary and considerable. The result was that only a handful of international groups survived. By producing a 4-way cross, they are able to protect their investments. That a baby chick could survive for a few days on nourishment from the yolk, coupled with development of the fixed wing aircraft, allowed for global distribution of broiler stocks throughout the world. Broiler production is based on breeding programs (i.e., sound broiler genetics). Its shape is a V, where final product has a narrow base for the source of elite stocks. An analogy is the limited number of sources for long distance aircraft for the international airlines. Broiler genetics has capitalized on a storehouse of genetic material coupled with science and technology developed over decades. It has allowed for application, which has allowed for an industry to provide an inexpensive meat product derived mainly from plant sources to a global consumer. The basic germplasm and research that allowed for the development of the broiler was derived mainly from public funding with little return to the science per se from which the programs were based. The caveat is that the broiler industry (not unlike some other industries) is dependent on a few multinational groups for their basic product. Their main biological tool is the genetics of the broiler. Their goal is to provide a food product—the broiler—to a growing public. Yes, they should support and conduct fundamental research, but, that is, not their function. The timeline from pedigree to broiler covers several generations and considerable resources. It is important that elite broiler breeding programs rightfully are located at multiple sites. This is essential not only in the event of disease outbreaks, but also, for example, climatic disparities and geopolitical issues. Thus, technological advances in network security, cloud, cybersecurity, redundancy, big data, and business continuity have become ever more relevant to successful broiler genetics. Globally, an ever emerging human population, with serious climate issues, suggests that there will be numerous challenges in the conversion of plant sources to broiler in the years to come (the production of laboratory meat is not within the realm of this essay). The major genetic changes in broiler breeding (e.g., Havenstein et al., 2003; Siegel, 2014) have been “cherry picked” from the availability of base populations, moderate to high heritability for important traits, and a short generation interval. Credit is given to those who took advantage of these items and realized that broiler breeding should be specific unto itself. Namely, broiler genetics is a subset of genetics per se and the broiler is the result of a complex biological system involving the life cycle of its genetic history. The plateau in broiler genetics will not be for body weight and accompanying positively correlated traits. Body weight is a trait influenced by many genes with small effects (Lillie et al., 2018). It is multifaceted and thus an issue with its genetic variation (as we know from Darwin, the lifeblood of a breeding program is genetic variation) and how to use it. The challenge is from biological and economic constraints of allocation of resources. Because the broiler as the final product is immature when marketed, reproduction cannot be ignored. Broiler breeders have to produce fertile eggs. Biologically, there is competition for mesodermal, endodermal, and ectodermal branches of development. This balancing among resources and allocations are seen in neural and metabolic factors associated with skeletal (e.g., Siegel et al., 2019), muscular (e.g., Petracci et al., 2019), cardiovascular (e.g., Wideman et al., 2013), food consumption (e.g., te Pas et al., 2020), and additional (perhaps unseen) issues. An ironic example is the replacement of plant sources in broiler diets with insects, once considered a pest (van Huis and Gasco, 2023). In this context, not to be dismissed is the coevolution of the microbiome and the hologenome concept (Yang et al., 2017; Zhou et al., 2022a; Zhou et al., 2022b). Such recent discoveries and technological advances provide new tools and challenges for the broiler breeder in the application of broiler genetics. To address this dynamic for competition for biological resources will require greater interactions, recognizing the sensitivity of proprietary rights and access of information to the scientific community and general public. This interface will not occur “overnight”, because public funding for broiler genetic research has declined. This has contributed to there being just a limited number of public institutions with the capability to train the next-generation of broiler geneticists, i.e., a basic understanding of the interface of the biology of avian species (poultry per se) with the technical skills necessary for the application of genetics in broiler breeding. Broiler genetics is the V of broiler breeding. Just as the distance from the primary breeder to the broiler per se is great, so is the distance from genotype to phenotype. This biological process is multifaceted, complex, and challenging. Be it broiler breeding or broiler genetics, the “kettle” is far from full. Thus, in concluding this essay, the quote from Eric Hoffer may be appropriate—“The only way to predict the future is to have the power to shape it.”

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          Most cited references23

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          Wooden-Breast, White Striping, and Spaghetti Meat: Causes, Consequences and Consumer Perception of Emerging Broiler Meat Abnormalities: Emerging broiler meat abnormalities…

          Ten years ago, the occurrence of macroscopic defects in breasts muscles from fast-growing broilers challenged producers and animal scientists to label and characterize myopathies wholly unknown. The distinctive white striations in breasts affected by white striping disorder, the presence of out-bulging and pale areas of hardened consistency in the so-called wooden breast, and the separation of the fiber bundles in breasts labelled as spaghetti meat, made these myopathies easily identified in chicken carcasses. Yet, the high incidence of these myopathies and the increasing concern by producers and retailers led to an unprecedented flood of questions on the causes and consequences of these abnormal chicken breasts. This review comprehensively collects the most relevant information from studies aimed to understand the pathological mechanisms of these myopathies, their physicochemical and histological characterization and their impact on meat quality and consumer's preferences. Today, it is known that the occurrence is linked to fast-growth rates of the birds and their large breast muscles. The muscle hypertrophy along with an unbalanced growth of supportive connective tissue leads to a compromised blood supply and hypoxia. The occurrence of oxidative stress and mitochondrial dysfunction leads to lipidosis, fibrosis, and overall myodegeneration. Along with the altered appearance, breast muscles affected by the myopathies display poor technological properties, impaired texture properties, and reduced nutritional value. As consumer's awareness on the occurrence of these abnormalities and the concerns on animal welfare arise, efforts are made to inhibit the onset of the myopathies or alleviate the severity of the symptoms. The lack of fully effective dietary strategies leads scientists to propose whether "slow" production systems may alternatively provide with poultry meat free of these myopathies.
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            Growth, livability, and feed conversion of 1957 versus 2001 broilers when fed representative 1957 and 2001 broiler diets.

            Body weight, feed consumption, and mortality were measured in the 1957 Athens-Canadian Randombred Control (ACRBC) strain and in the 2001 Ross 308 strain of broilers when fed representative 1957 and 2001 diets. The dietary regimens were chosen to be representative of those used in the industry in 1957 vs. 2001. The 1957 diets were fed as mash, the 2001 starter was as crumbles, and the grower and finisher diets were pellets. Feed consumption and BW were recorded at 21, 42, 56, 70, and 84 d of age to cover the two broiler strains normal span of marketing ages. Mortality was low, and the mortality of the ACRBC was approximately half that of the modem strain. Average BW for the ACRBC on the 1957 diets were 176, 539,809, 1,117, and 1,430 g vs. 743, 2,672, 3,946, 4,808, and 5,520 g for the Ross 308 on the 2001 diets at 21, 42, 56, 70, and 84 d of age, respectively. The 42-d feed conversion (FC) on the 2001 and 1957 feeds for the Ross 308 were 1.62 and 1.92 with average BW of 2,672 and 2,126 g and for the ACRBC were 2.14 and 2.34 with average BW of 578 and 539 g, respectively. The Ross 308 broiler on the 2001 feed was estimated to have reached 1,815 g BW at 32 d of age with a FC of 1.47, whereas the ACRBC on the 1957 feed would not have reached that BW until 101 d of age with a FC of 4.42.
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              Evolution of the modern broiler and feed efficiency.

              Although the chicken was domesticated during the Neolithic period, the development of the modern broiler is a recent event that has occurred within the past 100 years. The chicken's adaptability has allowed it to be grown globally under a range of husbandry conditions. That is, the same genetic stock may be found in a range of environments, where it is noted for rapid growth to market weight and efficiency of feed use, which has increased dramatically, mainly through genetic selection. Under good husbandry and a high-energy diet, at 35 days of age a 1.40-kg broiler required 3.22 kg of feed in 1985. Twenty-five years later, we have a 2.44-kg broiler produced on 3.66 kg of feed. This review attempts to address the history of factors contributing to these changes, obstacles that have had to be overcome, and future limitations.
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                Author and article information

                Contributors
                Journal
                Front Physiol
                Front Physiol
                Front. Physiol.
                Frontiers in Physiology
                Frontiers Media S.A.
                1664-042X
                08 March 2023
                2023
                : 14
                : 1150620
                Affiliations
                School of Animal Sciences , Virginia Tech , Blacksburg, VA, United States
                Author notes

                Edited by: Sandra G. Velleman, The Ohio State University, United States

                Reviewed by: Colin Guy Scanes, University of Arkansas, United States

                Francesca Soglia, University of Bologna, Italy

                *Correspondence: Paul B. Siegel, pbsiegel@ 123456vt.edu

                This article was submitted to Avian Physiology, a section of the journal Frontiers in Physiology

                Article
                1150620
                10.3389/fphys.2023.1150620
                10031763
                36969607
                13027a47-7042-4b16-9b65-2ce0145db3d8
                Copyright © 2023 Siegel.

                This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

                History
                : 24 January 2023
                : 23 February 2023
                Categories
                Physiology
                Opinion

                Anatomy & Physiology
                broiler,chicken,genetics,breeding,history
                Anatomy & Physiology
                broiler, chicken, genetics, breeding, history

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