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Part 1 book "Advances in animal nutrition and metabolism" includes content: Nutrition and metabolism - foundations for animal growth, development, reproduction, and health; insights into the regulation of implantation and placentation in humans, rodents, sheep, and pigs; a role for fructose metabolism in development of sheep and pig conceptuses; nutritional regulation of embryonic survival, growth, and development,.... and other contents.

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Advances in Experimental Medicine and Biology 1354 Guoyao Wu Editor Recent Advances in Animal Nutrition and Metabolism
Advances in Experimental Medicine and Biology Volume 1354 Series Editors Wim E. Crusio, Institut de Neurosciences Cognitives et Intégratives d’Aquitaine, CNRS and University of Bordeaux, Pessac Cedex, France Haidong Dong, Departments of Urology and Immunology, Mayo Clinic, Rochester, MN, USA Heinfried H. Radeke, Institute of Pharmacology & Toxicology, Clinic of the Goethe University Frankfurt Main, Frankfurt am Main, Hessen, Germany Nima Rezaei, Research Center for Immunodeﬁciencies, Children’s Medical Center, Tehran University of Medical Sciences, Tehran, Iran Ortrud Steinlein, Institute of Human Genetics, LMU University Hospital, Munich, Germany Junjie Xiao, Cardiac Regeneration and Ageing Lab, Institute of Cardiovascular Science, School of Life Science, Shanghai University, Shanghai, China
Advances in Experimental Medicine and Biology provides a platform for scientiﬁc contributions in the main disciplines of the biomedicine and the life sciences. This series publishes thematic volumes on contemporary research in the areas of microbiology, immunology, neurosciences, biochemistry, biomedical engineering, genetics, physiology, and cancer research. Covering emerging topics and techniques in basic and clinical science, it brings together clinicians and researchers from various ﬁelds. Advances in Experimental Medicine and Biology has been publishing exceptional works in the ﬁeld for over 40 years, and is indexed in SCOPUS, Medline (PubMed), Journal Citation Reports/Science Edition, Science Citation Index Expanded (SciSearch, Web of Science), EMBASE, BIOSIS, Reaxys, EMBiology, the Chemical Abstracts Service (CAS), and Pathway Studio. 2020 Impact Factor: 2.622 More information about this series at https://link.springer.com/bookseries/5584
Guoyao Wu Editor Recent Advances in Animal Nutrition and Metabolism 123
Editor Guoyao Wu Department of Animal Science Texas A&M University College Station, TX, USA ISSN 0065-2598 ISSN 2214-8019 (electronic) Advances in Experimental Medicine and Biology ISBN 978-3-030-85685-4 ISBN 978-3-030-85686-1 (eBook) https://doi.org/10.1007/978-3-030-85686-1 © Springer Nature Switzerland AG 2022 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, speciﬁcally the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microﬁlms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a speciﬁc statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, expressed or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional afﬁliations. This Springer imprint is published by the registered company Springer Nature Switzerland AG The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
Contents 1 Nutrition and Metabolism: Foundations for Animal Growth, Development, Reproduction, and Health . . . . . . . . . . 1 Guoyao Wu 2 Insights into the Regulation of Implantation and Placentation in Humans, Rodents, Sheep, and Pigs. . . . . . . . . 25 Claire Stenhouse, Heewon Seo, Guoyao Wu, Gregory A. Johnson, and Fuller W. Bazer 3 A Role for Fructose Metabolism in Development of Sheep and Pig Conceptuses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49 Robyn M. Moses, Avery C. Kramer, Heewon Seo, Guoyao Wu, Gregory A. Johnson, and Fuller W. Bazer 4 Nutritional Regulation of Embryonic Survival, Growth, and Development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63 Lawrence P. Reynolds, Kyle J. McLean, Kacie L. McCarthy, Wellison J. S. Diniz, Ana Clara B. Menezes, J. Chris Forcherio, Ronald R. Scott, Pawel P. Borowicz, Alison K. Ward, Carl R. Dahlen, and Joel S. Caton 5 Phosphate, Calcium, and Vitamin D: Key Regulators of Fetal and Placental Development in Mammals . . . . . . . . . . 77 Claire Stenhouse, Larry J. Suva, Dana Gaddy, Guoyao Wu, and Fuller W. Bazer 6 Nutritional and Physiological Regulation of Water Transport in the Conceptus . . . . . . . . . . . . . . . . . . . . . . . . . . . 109 Cui Zhu, Zongyong Jiang, Gregory A. Johnson, Robert C. Burghardt, Fuller W. Bazer, and Guoyao Wu 7 Amino Acids in Microbial Metabolism and Function . . . . . . . 127 Zhaolai Dai, Zhenlong Wu, Weiyun Zhu, and Guoyao Wu v
vi Contents 8 Potential Replacements for Antibiotic Growth Promoters in Poultry: Interactions at the Gut Level and Their Impact on Host Immunity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145 Christina L. Swaggerty, Cristiano Bortoluzzi, Annah Lee, Cinthia Eyng, Gabriela Dal Pont, and Michael H. Kogut 9 Microbiomes in the Intestine of Developing Pigs: Implications for Nutrition and Health . . . . . . . . . . . . . . . . . . . 161 Chunlong Mu, Yu Pi, Chuanjian Zhang, and Weiyun Zhu 10 L-Arginine Nutrition and Metabolism in Ruminants . . . . . . . 177 Guoyao Wu, Fuller W. Bazer, M. Carey Satterﬁeld, Kyler R. Gilbreath, Erin A. Posey, and Yuxiang Sun 11 Hepatic Glucose Metabolism and Its Disorders in Fish . . . . . 207 Xinyu Li, Tao Han, Shixuan Zheng, and Guoyao Wu 12 Protein-Sourced Feedstuffs for Aquatic Animals in Nutrition Research and Aquaculture . . . . . . . . . . . . . . . . . . 237 Sichao Jia, Xinyu Li, Wenliang He, and Guoyao Wu 13 Functional Molecules of Intestinal Mucosal Products and Peptones in Animal Nutrition and Health . . . . . . . . . . . . 263 Peng Li and Guoyao Wu 14 Use of Genome Editing Techniques to Produce Transgenic Farm Animals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 279 Alayna N. Hay, Kayla Farrell, Caroline M. Leeth, and Kiho Lee 15 Cows as Bioreactors for the Production of Nutritionally and Biomedically Signiﬁcant Proteins . . . . . . . . . . . . . . . . . . . 299 P. S. Monzani, P. R. Adona, S. A. Long, and M. B. Wheeler 16 Use of Agriculturally Important Animals as Models in Biomedical Research . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 315 Brandon I. Smith and Kristen E. Govoni 17 Pigs (Sus Scrofa) in Biomedical Research . . . . . . . . . . . . . . . . 335 Werner G. Bergen Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 345
Nutrition and Metabolism: Foundations for Animal Growth, 1 Development, Reproduction, and Health Guoyao Wu Abstract intimately interacts with a diverse community of intestinal antigens and bacteria to inﬂuence gut Consumption of high-quality animal protein and whole-body health. Understanding the plays an important role in improving human species and metabolism of intestinal microbes, nutrition, growth, development, and health. With as well as their interactions with the intestinal an exponential growth of the global population, immune systems and the host intestinal epithe- demands for animal-sourced protein are expected lium can help to mitigate antimicrobial resis- to increase by 60% between 2021 and 2050. In tance and develop prebiotic and probiotic addition to the production of food protein and alternatives to in-feed antibiotics in animal pro- ﬁber (wool), animals are useful models for duction. As abundant sources of amino acids, biomedical research to prevent and treat human bioactive peptides, energy, and highly bioavail- diseases and serve as bioreactors to produce able minerals and vitamins, animal by-product therapeutic proteins. For a high efﬁciency to feedstuffs are effective for improving the growth, transform low-quality feedstuffs and forages into development, health, feed efﬁciency, and survival high-quality protein and highly bioavailable of livestock and poultry, as well as companion essential minerals in diets of humans, farm and aquatic animals. The new knowledge cov- animals have dietary requirements for energy, ered in this and related volumes of Adv Exp Med amino acids, lipids, carbohydrates, minerals, Biol is essential to ensure sufﬁcient provision of vitamins, and water in their life cycles. All animal protein for humans, while helping reduce nutrients interact with each other to inﬂuence the greenhouse gas emissions, minimize the urinary growth, development, and health of mammals, and fecal excretion of nitrogenous and other birds, ﬁsh, and crustaceans, and adequate nutri- wastes to the environment, and sustain animal tion is crucial for preventing and treating their agriculture (including aquaculture). metabolic disorders (including metabolic dis- eases) and infectious diseases. At the organ Keywords level, the small intestine is not only the terminal site for nutrient digestion and absorption, but also Á Animal protein Biomedicine Á Health Á Á Á Disease Intestine Diet Abbreviations G. Wu (&) AAs Amino acids Department of Animal Science, Texas A&M AEMB Adv Exp Med Biol University, College Station, TX, USA IUGR Intrauterine growth restriction e-mail: g-wu@tamu.edu © Springer Nature Switzerland AG 2022 1 G. Wu (ed.), Recent Advances in Animal Nutrition and Metabolism, Advances in Experimental Medicine and Biology 1354, https://doi.org/10.1007/978-3-030-85686-1_1
2 G. Wu NRC National Research Council humans and other animals. Review articles of SDEP Spray-dried egg product select topics on nutrition and metabolism in animals of both agricultural and biomedical importance are highlighted in this volume of Advances in Experimental Medicine and Biology (AEMB) to beneﬁt our readers (Bergen 2022, Dai et al. 2022; Hay et al. 2022; Jia et al. 2022; Li et al. 2022; Li and Wu 2022; Monzani et al. 2022; Moses et al. 2022; Mu et al. 2022; Rey- nolds et al. 2022; Smith and Govoni 2022; 1.1 Introduction Stenhouse et al. 2022a, b; Swaggerty et al. 2022; Wu et al. 2022; Zhu et al. 2022). There are approximately 1.032 million animal species in nature, which include mammals, 4,000; birds, 9,000; ﬁsh and lower chordates, 1.2 Production of High-Quality 18,800; crustaceans, 45,000; reptiles, 6,300; Food Protein and Fiber (Wool) amphibians, 4,200; and insects, 0.9 (Wilson by Animal Agriculture 1992). As important parts of the ecosystem, for Human Consumption, animals ingest nutrients for survival, growth, Health and Well-Being development, reproduction, and health. Despite their vast diversities as two broad groups (ver- Animal agriculture (including aquaculture) plays tebrates and invertebrates), animals exhibit an important role in providing high-quality food greater similarities in physiology, metabolism, protein (e.g., milk, eggs, and meat) for human and nutrition than differences (Wu 2018). In consumption to optimize human growth, devel- human civilization, there has been a rich history opment, health, and well-being, although a mix of studies investigating the dietary requirements of complementary plant proteins through an of farm (e.g., livestock, poultry, ﬁsh, shrimp, and adequate understanding of protein nutrition may crabs), companion (e.g., cats, dogs, and horses), also result in a healthy nutritional state (Grillen- and laboratory (e.g., rats and mice) animals for berger et al. 2003; Murphy and Allen 2003; Wu energy, amino acids (AAs), lipids, vitamins, et al. 2022). Animal proteins generally contain minerals, and water during their life cycles under adequate and balanced amounts of all proteino- various physiological and pathological condi- genic (protein-creating) AAs for human con- tions (Baker 2008; Baldwin 1995; Bauman et al. sumption. In addition, animal-sourced foods 2011; Bazer et al. 2011, 2015, 2018, 2021; Beitz provide taurine, carnosine (b-alanyl-L-histidine), 1985; Bergen 2007, 2021; Burrin and Mersmann anserine (b-alanyl-L-1-methylhistidine), and 2005; Halloran et al. 2021; Matthews et al. 2016; creatine; they are nonproteinogenic nutrients that NRC 2002; Webb et al. 1992; Wu et al. 2022; possess antioxidative properties and are crucial Zhang et al. 2015; Zhu et al. 2022). This is for energy metabolism in tissues, but are absent because animals contribute to (a) the production from plant-sourced foods (Wu 2020b). As a part of high-quality foods (e.g., meats, eggs, and of healthy diets for humans, animal-sourced milk) for human consumption, as well as raw foods also contain highly bioavailable essential materials such as wool and leather for clothing minerals (e.g., iron, zinc, copper, manganese, and and accessories for humans; (b) the companion- selenium) and vitamins (Murphy and Allen ship and well-being of humans; (c) the develop- 2003). This is in contrast to a myth that there are ment of new biotechniques to efﬁciently produce virtually no nutrients in animal-based foods that proteins and other biomolecules; and (d) advanc- are not better provided by plants. Furthermore, ing biomedical research to prevent and treat animal by-products from the rendering industry inborn, metabolic, and infectious diseases of are major sources of protein, AAs, bioactive
1 Nutrition and Metabolism: Foundations for Animal Growth … 3 peptides, lipids, minerals, and vitamins in According to the Food and Agriculture the diets of livestock, poultry, ﬁsh, and crus- Organization of the United Nations (FAO 2021), tacean, and in pet foods (Li and Wu 2022; Li the global numbers of livestock and poultry have et al. 2021e; Wilkinson and Meeker 2021). increased by 3.6-fold over the past 60 years to Improved animal nutrition can enhance the about 33 billion head in 2019 (Table 1.1). Fur- quality of foods for human consumption, thermore, the global aquaculture production has whereas healthy companion animals contribute increased over the past decade at an average rate to the well-being of their owners. Finally, wool of about 3.5% per year to be approximately 120 (textile ﬁber; cysteine-rich a-keratin proteins) billion kg in 2019 and provides more than 50% produced by sheep and other animals (including of ﬁsh ﬁlets for human consumption (FAO cashmere and mohair from goats) is used to 2020). Both extensive (e.g., grazing pasture) and manufacture cloths and related daily life products intensive (e.g., indoor housing) systems are cur- (Cao et al. 2021). Production of proteins by rently used to raise ruminants (e.g., cattle, sheep, animals requires sufﬁcient provision and opti- and goats) and nonruminants (e.g., swine and mum utilization of dietary AAs and other nutri- poultry) worldwide (Bazer et al. 2020). Aquatic ents (Bergen 2021; Gilbreath et al. 2021; He animals are raised through both the outdoor et al. 2021a; Li et al. 2021a, b; Zhang et al. ponds and the indoor recirculating aquaculture 2021). This indicates a close link between animal systems (Ebeling and Timmons 2012). Farm and human nutrition. animals are biological transformers that convert Table 1.1 Global stocks of livestock species and poultry in 1961 and 2019a Species Year World Africa Australia Brazil Canada China Europe India Mexico USA Cattle 1961 942 123 17.3 56.0 10.7 49.5 192 176 16.5 97.7 2019 1510 361 24.7 215 11.5 63.5 117 193 35.2 94.8 Chickens 1961 3906 274 19.9 132 70.0 541 1329 108 62.6 751 2019 25,915 2043 112 1467 171 5247 2020 808 581 1972 Ducks 1961 193 6.23 0.178 2.68 0.398 100 25.8 6.70 1.00 3.50 2019 1,177 16.3 1.31 3.42 1.53 720 77.1 33.5 8.53 7.37 Geese + GF 1961 36.6 3.88 – – 0.314 16.5 13.5 – – – 2019 362 26.0 – – 0.326 312 15.0 – – – Goats 1961 349 94.2 0.04 4.90 0.012 51.3 22.5 60.9 8.93 3.47 2019 1090 459 3.90 11.3 0.301 137 16.1 149 8.79 2.62 Horses 1961 62.2 3.49 0.598 4.41 0.555 6.59 22.0 1.33 4.05 2.37 2019 59.0 7.40 0.222 5.85 0.399 3.67 4.70 0.342 6.38 10.7 Pigs 1961 406 5.67 1.61 25.6 5.00 85.6 168 5.18 5.99 55.6 2019 850 42.7 2.32 40.6 14.4 316 187 9.06 18.4 78.7 Rabbits + hares 1961 98.0 2.78 – 0.550 – 16.1 76.6 – 0.095 – 2019 300 16.0 – 0.161 – 233 9.63 – 1.40 – Sheep 1961 994 135 153 14.0 0.757 61.6 267 40.2 5.85 32.7 2019 1240 408 65.8 19.7 0.828 163 128 74.3 8.71 5.23 Turkeys 1961 204 1.21 0.17 0.698 3.50 0.313 80.5 – 6.00 108 2019 428 33.3 1.11 34.6 5.70 0.090 68.0 – 3.79 229 Adapted from FAO (2021). Values are Â 106 head a GF = guinea fowl; USA = United States of America; “–” = Data are not available
4 G. Wu materials not consumed by humans (e.g., forages; 1.3 Animals as Models by-products of plants such as pasture grasses, for Biomedical Research alfalfa, clovers, hays, straw, and silages; and and as Bioreactors rendered animal by-products) into high-quality for Producing Therapeutic foods (Wilkinson and Meeker 2021; Wu 2018). Proteins As documented in this volume of AEMB, recent research on dietary requirements for Biologically, humans are members of the animal nutrients (particularly AAs) is expected to kingdom. As noted previously, pigs, sheep, cat- enhance the efﬁciency of animal agriculture tle, chickens, and ﬁsh are agriculturally important globally and alleviate its potential adverse effects domestic animal species. There is growing on the environment. The signiﬁcance of animal interest in their use as animal models for nutrition agriculture is indicated by the fact that this research worldwide. This is because the nature of enterprise accounts for 50–75% and 25–40% of medical research often involves invasive tissue the total amount of agricultural output in indus- collections and surgical procedures and may trialized and developing countries, respectively result in potential harmful effects (Bergen 2021; (Wu et al. 2014c). Animal-sourced foods can Govoni et al. 2019; Ireland et al. 2008; Jia et al. prevent protein deﬁciency in children and adults 2021; Odle et al. 2017; Reynold et al. 2019, (including the elderly and hospitalized patients) 2022; Smith and Govoni 2022; Webb et al. 1992; worldwide, particularly those living in underde- Wu and Knabe 1994). Thus, it is neither ethical veloped nations (Wu 2021). In 2016, about 815 nor practical to conduct such studies with million people (10.7% of the world population) humans in the fetal, infant, or adult stages of life. had chronic deﬁciencies of nutrients, particularly Similarities, major differences, as well as protein, vitamins, and microminerals (FAO advantages and disadvantages of porcine, ovine, 2018), and globally 150 million children under bovine, avian, and ﬁsh models are summarized in ﬁve years of age were stunted in their growth Table 1.2. Examples of using animal models to (UNICEF 2018). Maternal malnutrition during pursue nutrition research are highlighted in gestational and neonatal periods affects not only Table 1.3. Of particular note, results of those the ﬁrst generation of offspring, but also subse- studies have aided in delineating the mechanisms quent generations through epigenetic-mediated for the following physiological or pathological mechanisms (Del Curto et al. 2013; Wang et al. features in humans: (1) intrauterine growth 2012). Preventing hunger and malnutrition will restriction (IUGR); (2) arginine deﬁciency and be an even greater challenge as the global pop- hyperammonemia in preterm infants; (3) abun- ulation is expected to grow exponentially from dance of polyamines as well as glutamine and 7.9 billion people in 2021 to 9.6 billion by 2050 proline in milk; (4) the obligatory role of dietary (United Nations 2021). The demands for animal- AAs in intestinal integrity; (5) hyperglycemia- sourced protein are expected to increase by 60% induced endothelial dysfunction; (6) ammonia between 2021 and 2050 for supporting optimal toxicity in patients with N-acetylglutamate deﬁ- human growth and mitigating sarcopenia in the ciency; (7) extrahepatic urea synthesis in the elderly. Thus, animal agriculture plays an small intestine of post-weaning mammals; (8) the important role in providing animal-sourced food susceptibility of neonates to intestinal disease; as part of a healthy diet for humans, while con- and (9) biomarkers for intestinal adaptation in tributing to scientiﬁc, economic, and social preterm neonates (Rhoads et al. 2005; Rhoads developments worldwide. and Wu 2009; Wu and Morris 1998; Wu et al.
1 Nutrition and Metabolism: Foundations for Animal Growth … 5 Table 1.2 Similarities, major differences, as well as advantages and disadvantages of pig, sheep, cattle, chicken, and ﬁsh models for biomedical research on human nutrition, metabolism, and health Animal Similarities to humans Major differences than Advantages Disadvantages model humans Pig Anatomy, physiology, Pregnancy: Time of Provides adequate tissue The growth of fetal pigs digestion, food intake, implantation, type of samples for various is not highly sensitive to and the metabolism of placentation, fetal ﬂuid biochemical assays; maternal deﬁciency of nutrients (including compartments, convenient to work with protein; high risk for AAs, glucose, and gestational length, and both fetal and postnatal abortion in pregnant minerals); monogastric number of offspring; pigs; sensitive to dietary pigs after surgical omnivores; mechanisms Nutrition: Amounts of intakes of AAs and catheterization of blood for nutrient transport; nutrients (fat and iron) other nutrients; a large vessels; no BAT; litter- the regulation of blood stored in the body at database in the literature bearing ﬂow birth; Metabolism: on pigs Synthesis of ascorbic acid and the site of FA synthesis Sheep Metabolic pathways for Pregnancy: Time of Well-established animal Extensive fermentation nutrient utilization, the implantation, type of model for studying of dietary nutrients in regulation of blood ﬂow placentation, fetal ﬂuid human pregnancy, the rumen, most dietary and thermogenesis, compartments, Convenient to work nutrients must be mechanisms for nutrient pregnancy recognition with sheep, pregnant protected from rumen transport, the number of signal, and gestational ewes are well adaptable degradation, sensitive to offspring, maternal size, length; Digestion: to surgical procedures ketosis and copper embryogenesis, BAT Fermentation of of placing catheters into toxicity, a high rate of and thermogenesis, and nutrients in the rumen; maternal and fetal blood mortality during late birth weight Metabolism: Intestinal vessels, and a large pregnancy for ewes catabolism of BCAAs; database in the literature with ! 3 fetuses the metabolism of on sheep glucose, ammonia, and SCFAs; gestational length; the site and substrates for FA synthesis Cattle Metabolic pathways for Pregnancy: Time of Well-established animal Extensive fermentation nutrient utilization, the implantation, type of model for studying of dietary nutrients in regulation of blood ﬂow placentation, fetal ﬂuid human citrullinemiaa, the rumen, most dietary and thermogenesis, compartments, and easiness for in vitro and nutrients must be mechanisms for nutrient pregnancy recognition in vivo gene transfer protected from rumen transport, number of signal; Digestion: studies, historically used degradation, sensitive to offspring, Fermentation of for research on ketosis and rumen bloat embryogenesis, nutrients in the rumen; infectious infectious (excessive accumulation gestational length, as Metabolism: Intestinal diseases (e.g., of gases in the rumen), well as BAT and catabolism of BCAAs, tuberculosis, cowpoxb), inconvenient to work thermogenesis the metabolism of and production of with adult cattle, and glucose, ammonia, and interferon c high costs of doing SCFAs; the site and experiments substrates for FA synthesis Chicken Digestion and Embryo: Hatching of Provides adequate tissue Different mechanisms absorption of nutrients eggs in birds versus samples for various for ammonia detoxiﬁ- in the small intestine, mammalian embryos in biochemical assays; cation, no placenta for the metabolism of most the uterus, different convenient to work with embryonic growth and nutrients, monogastric length of embryonic both pre- and post- development, different omnivores, the development; hatching birds, sensitive paths for the absorption regulation of blood Digestion: to dietary intakes of of lipids and lipid- ﬂow, angiogenesis, Proventriculus and AAs and other nutrients; soluble vitamins, the mechanisms for nutrient gizzard, the expression unique to study gout, lack of muscle GLUT4, transport, the site and of digestive enzymes in retinal degenerationd, and mammalian (continued)
6 G. Wu Table 1.2 (continued) Animal Similarities to humans Major differences than Advantages Disadvantages model humans substrates for FA the small intestine, and and angiogenesis, antibodies are generally synthesis, and BATc lipid absorption via the naturally “diabetic”, and not applicable to birds portal vein Metabolism: spontaneously develops Uric acid synthesis, the ovarian cancer lack of hepatic gluconeogenesis from AAs, and high metabolic rate Rodente Anatomy, physiology, Pregnancy: Time of Provides adequate tissue A short period of digestion, and nutrition; implantation, type of samples for various gestation (21 days), metabolic pathways for placentationf, fetal ﬂuid biochemical assays; ability to synthesize nutrient utilization, compartments, convenient to work with vitamin C, high food monogastric omnivores; gestational length, and both fetal and postnatal intake per kg body mechanisms for nutrient number of offspring; rodents; sensitive to weight, rapid aging transport; the regulation Digestion: Fast gastric dietary intakes of AAs processing, difﬁculties of blood ﬂow; as well as emptying and short GI and other nutrients; a in performing surgeries BAT and thermogenesis transit time (reaching large database in the on rodents, litter- post-absorptive state at literature on rodents; bearing, small size, and 6 h after feeding versus widely available; low very different metabolic 12 h in humans) cost rates between rodents Metabolism: High and humans metabolic rate (6–10 times that in humans) Fish Digestion and Embryo: Hatching of Big ﬁsh provide Different mechanisms absorption of nutrients eggs in ﬁsh vs. adequate tissue samples for ammonia removal, in the small intestine, mammalian embryos in for various biochemical no placenta for metabolic pathways for the uterus, different assays; convenient to embryonic growth and nutrient utilization, length of embryonic work with both pre- and development, different mechanisms for nutrient development; post-hatching ﬁsh, metabolic patterns for transport, intestinal AA Digestion: Some ﬁsh sensitive to dietary glucose and fatty acids metabolism, and the lack a stomach; a short intakes of starch and in ﬁsh than in mammals, secretion of digestive gut; requires a long time other nutrients; unique requires intensive labor, enzymes for nutrient digestion to study starch- mammalian antibodies and absorption; intolerable disease in are generally not Metabolism: Most ﬁsh the liver; useful for applicable to ﬁsh, lack the urea cycle and studying the usual difﬁcult to measure food have high dietary absence of tumors in the intake, and small size requirements for AAs; small intestine; for juvenile ﬁsh intolerable to dietary zebraﬁsh is widely used starch; limited oxidation in biomedical research of glucose and lipids in muscle; unique roles of the head kidneysg a A disease due to the deﬁciency of argininosuccinate synthase (a urea cycle enzyme) b The ﬁrst vaccine against the cowpox virus c Deveopment of brown adipose tissue in response to cold challenge d Mutation in the photoreceptor guanylate cyclase e Rats and mice f Labyrinthine-type in rats (an intricate structure of interconnecting passages) versus the villous-type in humans g A tissue that contains cytokine-producing lymphoid cells of the immune system; endocrine cells that secret cortisol, catecholamines, and thyroid hormones; and hematopoietic stem cells that are capable of hematopoiesis (the production of new blood cells) AAs = amino acids; BAT = brown adipose tissue, BCAAs = branched-chain amino acids; FA = fatty acid; GI = gastrointestinal; GLUT4 = glucose transporter 4; SCFAs = short-chain fatty acids
1 Nutrition and Metabolism: Foundations for Animal Growth … 7 Table 1.3 Use of animal models for nutrition and biomedical research Animal Nutrition and biomedical research References Pig Fetal and postnatal development of intestine and Buddington et al. (2012), Dekaney et al. (2001) other tissues Effects of maternal nutrition on placental and fetal Ji et al. (2017), NRC et al. (2012), Wu et al. (1996, growth 2006) Fetal composition of AAs and other nutrients in the Kim et al. (2009), Pond et al. (1969), Wu et al. body (1999) Tissue-speciﬁc and whole-body metabolism of Blachier et al. (2013), Reeds et al. (1996, 1997) amino acids Mammary gland synthesis of bioactive products Hurley (2019), O’Quinn et al. (2002) Nutrient requirements of neonates and adults Davis et al. (2002, 2008, 2010), Wang et al. (2014) Development of infant formulas and TPN solutions Brunton et al. (1999), Jamin et al. (2010), Odle et al. (2017) Regulation of vasodilator production by endothelial Wang et al. (2011), Wu et al. (2001) cells Cardiovascular diseases and responses to exercise Tsang et al. (2016); Walters et al. (2017) and diets Novel biochemical pathways and metabolic defects Wu (1997), Wu et al. (1994, 2000, 2004, 2005) a Development of immune systems Furukawa et al. (2020), Johnson et al. (2006), Wu (1996) Prevention and treatment of IUGR in mammals Ashworth (1991), Rehfeldt et al. (2004), Wu et al. (2006) Production of recombinant proteins and Hay et al. (2022), Monzani et al. (2022) antimicrobials Microbial development in the small and large Dai et al. (2022); Mu et al. (2022); Ren et al. intestines (2020) Sheep Composition of AAs and PAs in fetal ﬂuids and Kwon et al. (2003a, b; 2004a, b) placentae Expression of AA and sugar transporters in the Gao (2020), Moses et al. (2022) conceptus Syntheses of NO and PAs in placentae Kwon et al. (2003a, b), Wang et al. (2015a, b, 2016) Prevention of IUGR in underfed dams through Arg Gilbreath et al. (2021), Lassala et al. (2009, 2010, supplementation 2011), McCoard et al. (2013, 2014, 2016), Sales et al. (2016) Growth and lactation performance Reynolds et al. (2019), Wu et al. (2022) Skeletal muscle growth, development, and Gonzalez et al. (2020), Govoni et al. (2019) adaptation Cow Transgenic cattle with desirable production traitsb Hay et al. (2022), Monzani et al. (2016, 2022) As bioreactors to produce recombinant proteinsc Hay et al. (2022), Monzani et al. (2016; 2021) d Develop therapeutic treatment of citrullinemia Harper et al. (1986), Lee et al. (1999) Chicken Elucidate mechanisms responsible for gout, retinal Cebulla et al. (2012), Lim et al. (2012), Larger degeneration and detachment, diabetes, and ovarian et al. (2004), Ulshafer and Allen (1985) cancer, as well as their prevention and treatment Study cardiovascular development and angiogenesis Vilches-Moure (2019) and Ziche et al. (1994) (continued)
8 G. Wu Table 1.3 (continued) Animal Nutrition and biomedical research References Fish Study mechanisms for the utilization of high dietary Ballantyne (2001), Li et al. (2020a, d, e, f) protein Study mechanisms for glycogenic hepatopathy Li et al. (2020a, b; 2022) Study mechanisms for hepatic steatosis Li et al. (2022) Study mechanisms for black skin syndrome Li et al. (2021a, b, c, d) Study mechanisms for the absence of cancer in the Jia et al. (2021) intestine Study mechanisms for Arg deﬁciency in growth and Li et al. (2021a, b, 2022) survival a Including intestinal intraepithelial lymphocytes and Peyer’s patches b Including resistance to diseases and improved growth and lactation performance c Including tissue-type plasminogen activator, recombinant human growth hormone, recombinant human albumin, recombinant anti-CD20 monoclonal antibody, human lactoferrin, a-lactalbumin, myelin basic protein, and human bile salt-stimulating d A rare Holstein and Holstein–Friesian-speciﬁc metabolic genetic disorder of cattle AAs = amino acids; Arg = arginine; IUGR = intrauterine growth restriction; NO = nitric oxide; PAs = polyamines (putrescine, spermidine, and spermine); TPN = total parenteral nutrition 2004, 2014a). Examples of discovery research carcinogenesis (Jia et al. 2021; Li et al. 2021a, b, using ovine models has greatly advanced the 2022). following aspects of AA nutrition research: ((1) Over the past two decades, genetic engineer- the unusual high abundance of the arginine- ing techniques [e.g., recombinant DNA technol- family AAs and serine in fetal ﬂuids (Kwon et al. ogy and genome editing (Fig. 1.1)] have been 2003a, b), (2) the expression of AA and sugar used to generate transgenic cattle that possess transporters in the conceptus (Gao 2020; Huang desirable production traits (including resistance et al. 2018; Moses et al. 2022; Reynolds et al. to diseases and improved growth and lactational 2022; Satterﬁeld et al. 2010); (3) the syntheses of performance) and produce recombinant proteins nitric oxide and polyamines (key regulators of with nutritional and therapeutic values (Hay et al. angiogenesis) in placentae (Kwon et al. 2003a, 2022; Monzani et al. 2016, 2022). Those proteins b); and (4) the prevention of IUGR in underfed include tissue-type plasminogen activator, dams by increasing the availability of AAs to the recombinant human growth hormone (nutrient fetus through either the dietary realimentation or metabolism), recombinant human albumin, modulation of the uterine arginine-nitric oxide recombinant anti-CD20 monoclonal antibody, pathway (Gilbreath et al. 2021; Lassala et al. human lactoferrin (antimicrobial in the small 2009, 2010, 2011; Satterﬁeld et al. 2012, 2013). intestine), a-lactalbumin (milk protein), myelin In additions, chickens are useful models to study basic protein (neurological development), and mechanisms responsible for gout, retinal degen- human bile salt-stimulating lipase (lipid diges- eration and detachment, diabetes, and ovarian tion). In addition, transgenic pigs have been cancer (Cebulla et al. 2012; Lim et al. 2012; generated to produce porcine growth hormone, Larger et al. 2004; Ulshafer and Allen 1985, as carbohydratases, phytase, antimicrobials, and well as cardiovascular development (Vilches- anti-viral antibodies (Wu and Bazer 2019). Moure 2019) and angiogenesis (Ziche et al. Advanced biotechnology holds great promise for 1994). Finally, ﬁsh can be used to investigate conserving the diverse breeds of animals, mechanisms for the utilization of high amounts enhancing their food efﬁciency and productivity, of dietary protein, glycogenic hepatopathy, hep- and developing new alternatives to in-feed atic steatosis, black skin syndrome, and intestinal antibiotics in the future.
1 Nutrition and Metabolism: Foundations for Animal Growth … 9 Fig. 1.1 Gene (genome) editing of animals using the zinc homologous template for repair, which often inserts or ﬁnger nuclease (ZFN), transcription activator-like effector deletes nucleotides (indels) to cause gene disruption nuclease (TALEN), or clustered regularly interspaced (knockout). The HDR pathway requires the provision of short palindromic repeats-associated nuclease-9 an exogenous DNA template along with a site-speciﬁc (CRISPR/Cas9) technique. A designer nuclease (ZFN, genome editing nuclease to repair the DSB DNA, thereby TALEN or CRISPR/Cas9) cleaves a DNA molecule to causing the knock-in of a desired sequence of DNA into generate a double-strand break (DSB) at a desired the genome of an embryo or animal cells. Because of its genomic locus. Thereafter, one of two endogenous repair more precise targeting of genes, CRISPR/Cas9 is gaining mechanisms may repair the DSB DNA: non-homologous momentum in life sciences as the preferred editor of gene end joining (NHEJ) and the homology-directed repair editing of livestock species. Reproduced from Wu and (HDR). In the NHEJ pathway, the two ends of the Bazer (2019), with permission DSB DNA are brought together and ligated without a 2016; NRC 2002, 2012; Wu 2018). All nutrients 1.4 Nutritional Requirements, interact with each other, intestinal microbes, and Deficiencies, and Health the environment to inﬂuence growth, develop- of Animals ment, and health of animals (Fig. 1.2). Grazing ruminants without protein or mineral supple- All animals have dietary requirements for ments exhibit suboptimal growth and lactational energy, AAs, lipids, carbohydrates, minerals, performance (Bergen et al. 2021; Cao et al. 2021; vitamins, and water in their life cycles (Greene Gilbreath et al. 2021; Govoni et al. 2019).
10 G. Wu Fig. 1.2 Metabolic disorders (including metabolic dis- anemia and hemorrhage (iron), ammonia toxicity (man- eases) in ruminant and nonruminant animals. Defective ganese), Keshan’s disease (selenium), goiter (iodine), biochemical pathways due to deﬁciencies in their dental caries (ﬂuorine), Menke’s and Wilson's diseases enzymes, coenzymes, or cofactors can cause abnormal (copper), and infertility (phosphorus). cDisorders (includ- nutrient metabolism (either inherited or acquired), result- ing diseases) caused by an excessive production of amino ing in multi-organ dysfunctions, diseases and even death acid metabolites include hyperhomocysteinemia, hyper- in livestock and poultry. Reproduced from Wu (2020c), ammonemia, gout, melanosis, and porphyria. dDisorders with permission. aDisorders (including diseases) caused (including diseases) caused by an excessive lipid-soluble by the deﬁciency of a vitamin include polioencephalo- vitamin include hypervitaminosis A, D, E, and K, whereas malacia in ruminants and beriberi (thiamin), pellagra diseases caused by an excessive mineral include polioen- (niacin), burning-foot syndrome (pantothenic acid), neural cephalomalacia in ruminants (sulfate), hypertension tube defects (folate), scurvy (vitamin C), photophobia (sodium), hyperkalemic periodic paralysis in horses (riboﬂavin), xerophthalmia and keratomalacia (vitamin (potassium), copper toxicity, and selenosis (selenium). A), rickets and osteomalacia (vitamin D), liver steatosis e This reaction is catalyzed by urease in the rumen ﬂuid of (choline), myopathy and liver necrosis (vitamin E), ruminants and the intestine of all animals. As, ascites hemorrhage (vitamin K), and infertility (vitamins A and syndrome; CLA, conjugated linoleic acid; FLHS, fatty E). bDisorders (including diseases) caused by the deﬁ- liver hemorrhagic syndrome; FLKS, fatty liver and kidney ciency of a mineral include milk fever in cows, rickets, syndrome and osteomalacia (calcium), grass tetany (magnesium), Likewise, nonruminants (e.g., swine and poultry) et al. 2007). The pages of AEMB have high- fed typical corn- and soybean meal-based diets lighted recent advances in the functions of AAs without dietary supplementation with deﬁcient in the different organ systems of animals AAs cannot achieve their maximum genetic (Beaumont and Blachier 2020; Chen et al. 2020; potential for growth and egg production (He Durante 2020; Flynn et al. 2020; Gao 2020; He et al. 2021a; Zhang et al. 2021). Thus, because and Wu 2020; Hou et al. 2020; Li et al. 2020a; protein is the most abundant dry matter compo- Sandoval et al. 2020; Solano 2020; Wu 2021), nent in growing animals (e.g., about 65–67% in the inter-organ metabolism of AAs (He et al. skeletal muscle on the dry matter basis) and a 2021b; Posey et al. 2021; Ryan et al. 2021), and major nutrient in human foods (e.g., meat, eggs, the roles of AAs in gene expression and cell and milk) and feedstuffs for animal diets, protein signaling (Halloran et al. 2021; Paudel et al. nutrition and metabolism has been an active 2021; Sah et al. 2021; Shen et al. 2021; Yang research area in animal nutrition over the past et al. 2021). Additional AEMB papers focus on century (Baker 2008; Bergen 2007, 2008; Firkins the nutrition and metabolism of humans and
1 Nutrition and Metabolism: Foundations for Animal Growth … 11 other animals, including aquatic, companion, zoo intimately interacts with a diverse community of animals (Che et al. 2021; Herring et al. 2021; Jia intestinal antigens and bacteria to inﬂuence gut et al. 2021; Li et al. 2022; Oberbauer and Larsen and whole-body health (Ren et al. 2020; Wang 2021; Sarkar et al. 2021; Wu et al. 2021). et al. 2020). Compared with humans, farm ani- Compelling evidence indicates that animals (in- mals are at greater risks for infections by cluding humans) have dietary requirements for pathogens such as bacteria, fungi, parasites, and proteinogenic AAs that are not synthesized de viruses. Thus, maintaining a healthy gut is novo (e.g., leucine and lysine) and proteinogenic essential to the survival, growth, and reproduc- AAs that are synthesized de novo in animal cells tion of the animals (Ren et al. 2020). Since the (e.g., glutamate, glutamine, glycine, and proline; 1950s, sub-therapeutic levels of feed antibiotics Wu 2021). Some of the AAs, known as func- have been included in conventional diets to tional AAs in nutrition (Wu 2010), participate in improve the growth performance and feed efﬁ- and regulate key metabolic pathways to improve ciency of swine and poultry. However, due to the the health, survival, growth, development, lacta- development and spread of bacteria resistant to tion, and reproduction of animals. Notably, these antibiotics, feed antibiotics have been banned in two new nutritional concepts are now trans- many countries (e.g., the European Union, the U. forming the feeding practices for animals (in- S., and China) and are being phased out in many cluding livestock, poultry, and ﬁsh) worldwide other nations. Some bacteria are resistant to one (Chalvon-Demersay et al. 2021; Li et al. 2021a, class of antibiotics, and others are resistant to b; Rodrigues et al. 2021; Rossi et al. 2021). In multiple antibiotics, thereby posing a serious addition, this volume of AEMB includes articles global health concern (Koch et al. 2017). Several on mechanisms for the transport of water (Zhu comprehensive articles in this volume of AEBM et al. 2021), fructose (a major sugar in the con- highlight the species and metabolism of intestinal ceptuses of ungulates; Moses et al. 2022), cal- microbes, as well as their interactions with the cium, and phosphorus (Stenhouse et al. 2022b) in intestinal immune systems and the host intestinal adult animals, particularly in pregnant dams with epithelium in swine and poultry (Dai et al. 2022; developing conceptuses. Adequate nutrition is Mu et al. 2022; Swaggerty et al. 2022). This new crucial for preventing and treating metabolic knowledge can help to mitigate antimicrobial disorders (including metabolic diseases) and resistance through the development of prebiotic infectious diseases in all species of animals (Li and probiotic alternatives to in-feed antibiotics in et al. 2007; Wu 2020c; Table 1.4). animal production systems. The small intestine is not only the terminal site for nutrient digestion and absorption, but also plays an important role in AA metabolism 1.5 Protein Foodstuff Sources (Fig. 1.3). Notably, the synthesis of citrulline for Animals from glutamine, glutamate, and proline occurs in the enterocytes (the columnar absorptive epithe- Protein is the most expensive nutrient in animal lial cells of the small intestine) of most mammals, diets (Gatlin et al. 2007; Kim et al. 2009; Li and Wu including humans, pigs, rats, mice, cattle, and 2020; Li et al. 2020b, c; Wu et al. 2014a). Plant- sheep (Blachier et al. 2013; Wu et al. 2021, 2022; and animal-sourced foodstuffs are the sources of Zhang et al. 2021), but not in any cells of protein and AAs for omnivores, whereas carni- chickens (He et al. 2021a, b). As for pigs (Zhang vores consume animal carcasses or animal-derived et al. 2021), the proximal intestine of ﬁsh (e.g., products. As reviewed in this volume of AEBM hybrid striped bass and largemouth bass; Li et al. (Jia et al. 2022; Li and Wu 2022; Li et al. 2021e), 2020d, e, f; Jia et al. 2021) and crustaceans (Li the composition of most AAs differ substantially et al. 2021b) extensively oxidizes glutamate, between plant and animal proteins. Animal- glutamine, and aspartate to provide most of the sourced feedstuffs are generally superior to plant- energy needed by the tissue. In addition, the gut sourced ones for the growth and health of
12 G. Wu Table 1.4 Metabolic disorders (including metabolic diseases) in ruminant and nonruminant animals due to nutrient deﬁciencies Nutrient Metabolic disorder (or Cause Major syndromes disease) Glucose Type 1 diabetes Lack of insulin secretion from Hyperglycemia, retinal damage, pancreatic b-cells, impaired use of blindness, ketosis, impaired blood ﬂow glucose (leading to amputation), muscle loss and weakness, and dyslipidemia Type 2 diabetes Insulin resistance (impaired) insulin Hyperglycemia, retinal damage, signaling, or obesity), impaired use of blindness, impaired blood ﬂow (leading glucose to amputation), excessive glycogen in muscle, and dyslipidemia Hypoglycemia Inadequate glucose provision or Brain damage, coma, and death synthesis Ruminal acidosis in High intake of starch and Inhibits the growth of cellulolytic ruminants monosaccharides, a sudden decrease in bacteria and acetate-producing bacteria, ruminal ﬂuid pH to
1 Nutrition and Metabolism: Foundations for Animal Growth … 13 Table 1.4 (continued) Nutrient Metabolic disorder (or Cause Major syndromes disease) behavior (e.g., walking in circles), and reduced blood pH Low-fat milk syndrome Low intake of ﬁber and high intakes of A reduction in the concentrations of milk (milk fat depression) in starch and unsaturated fatty acids; high fats (up to 50% or more) with little or no dairy cows amount of CLA in the rumen change in concentrations of lactose or protein in milk, and reductions in ruminal pH and Ac/Prop ratio Pregnancy toxemiaa Excessive mobilization of adipose Reduced feed intake, reduced blood pH, (prevalent in ewes and does tissue due to low feed intake, reduced fetal growth, skeletal muscle loss with multiple fetuses in late excessive KB production by the liver and weakness, abnormal behavior (e.g., pregnancy) walking in circles), and maternal and fetal death Fatty liver hemorrhagic Excessive energy intake, particularly High amount of abdominal fats, and syndrome in laying hens in heat stress (often in proliﬁc laying possibly pale combs; the enlarged liver is hens housed in cages) prone to damage and bleeding. Hemorrhage often occurs when a hen is straining to lay her egg. A high rate of mortality Fatty liver and kidney in Biotin deﬁciency, and thus impaired The liver and kidneys are pale and chickens carboxylationb swollen, and contain high lipid deposits; lactic acidosis; death Yellow fat diseasec Excessive intake of oxidized Marked inﬂammation of white adipose (steatitis or pansteatitis) unsaturated fatty acids, and reduced tissue, lipid peroxidation, and the intakes of vitamin E deposition of yellow-wish pigment in adipocytes; possibly myopathy Gangliosidoses A lack of or low activity of enzymes to Excessive lysosomal accumulation of hydrolyze GM lipids known as gangliosides in tissues Gaucher’s disease Deﬁciency of b-glucocere-brosidase to Excessive lysosomal accumulation of degrade GCB lipids known as glucocerebroside in tissues Amino Amyloidosis Abnormal synthesis, degradation, or Deposition of amyloid ﬁbrils (ﬁrm and acids folding of extracellular amyloidd solid extracellular substances) in tissues; (AAs) organ failure, and death Kwashiorkor Dietary protein deﬁciency Weakness, poor health, stunting, anemia, muscle wasting, calcium and bone losses, edema, reduced number of red blood cells, and impaired immunity Hyperammonemia Excessive ammonia in blood, an Arg Impaired ﬂow of blood to the brain, or UCE deﬁciency in mammals; tissue damage, oxidative stress, impaired excess AA intake Krebs cycle, pregnancy loss, coma, and death Gout Excessive urate production, an Arg or High amount of uric acid crystallizes in UCE deﬁciency in mammals; excess joints, tendons, and surrounding tissues, AA intake resulting in red, tender, hot, and swollen tissues; pain Hyperhomocysteinemia Excessive homocysteine in blood due Deﬁciency of NO, impaired ﬂow of to low intakes of vitaminse or high blood to tissues, high risk for SAA intake (continued)