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Committee on Nutrient Requirements of Small Ruminants
Board on Agriculture and Natural Resources
Division on Earth and Life Studies
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THE NATIONAL ACADEMIES PRESS · 500 Fifth Street, N.W. · Washington, DC 20001
NOTICE: The project that is the subject of this report was approved by the Governing Board of the National Research
Council, whose members are drawn from the councils of the National Academy of Sciences, the National Academy of
Engineering, and the Institute of Medicine. The members of the committee responsible for the report were chosen for
their special competences and with regard for appropriate balance.
This study was supported by grants from the National Sheep Industry Improvement Center (U.S. Department of
Agriculture), the Joe Skeen Institute for Rangeland Restoration (Montana State University, New Mexico State
University, and Texas A&M University), the American Sheep Industry Association, the Montana Sheep Institute,
internal National Research Council funds, and general support of the Animal Nutrition series provided by the
Department of Health and Human Services (U.S. Food and Drug Administration) under Award No. 223-01-01-2460.
Any opinions, findings, conclusions, or recommendations expressed in this publication are those of the author(s) and
do not necessarily reflect the views of the organizations or agencies that provided support for the project.
Library of Congress Cataloging-in-Publication Data
Nutrient requirements of small ruminants : sheep, goats, cervids, and New World camelids / Committee on Nutrient
Requirements of Small Ruminants, Board on Agriculture and Natural Resources, Division on Earth and Life Studies,
The National Research Council.
p. cm.
Includes bibliographical references and index.
ISBN 0-309-10213-8 (cloth) -- 978-0-309-10213-1 (cloth) 1. Ruminants--Feeding and feeds. 2. Ruminants--
Nutrition--Requirements. I. National Research Council (U.S.). Committee on Nutrient Requirements of Smal
Ruminants.
SF95.N78 2006
636.30852--dc22
2006028306
Additional copies of this report are available from the National Academies Press, 500 Fifth Street, N.W., Lockbox 285,
Washington, DC 20055; (800) 624-6242 or (202) 334-3313 (in the Washington metropolitan area); Internet,
http://www.nap.edu
Copyright 2007 by the National Academy of Sciences. All rights reserved.
Printed in the United States of America
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The National Academy of Sciences is a private, nonprofit, self-perpetuating society of distin-
guished scholars engaged in scientific and engineering research, dedicated to the furtherance of
science and technology and to their use for the general welfare. Upon the authority of the char-
ter granted to it by the Congress in 1863, the Academy has a mandate that requires it to advise
the federal government on scientific and technical matters. Dr. Ralph J. Cicerone is president of
the National Academy of Sciences.
The National Academy of Engineering was established in 1964, under the charter of the
National Academy of Sciences, as a parallel organization of outstanding engineers. It is auton-
omous in its administration and in the selection of its members, sharing with the National Acad-
emy of Sciences the responsibility for advising the federal government. The National Academy
of Engineering also sponsors engineering programs aimed at meeting national needs, encourages
education and research, and recognizes the superior achievements of engineers. Dr. Wm. A. Wulf
is president of the National Academy of Engineering.
The Institute of Medicine was established in 1970 by the National Academy of Sciences to se-
cure the services of eminent members of appropriate professions in the examination of policy
matters pertaining to the health of the public. The Institute acts under the responsibility given to
the National Academy of Sciences by its congressional charter to be an adviser to the federal
government and, upon its own initiative, to identify issues of medical care, research, and educa-
tion. Dr. Harvey V. Fineberg is president of the Institute of Medicine.
The National Research Council was organized by the National Academy of Sciences in 1916
to associate the broad community of science and technology with the Academy's purposes of
furthering knowledge and advising the federal government. Functioning in accordance with gen-
eral policies determined by the Academy, the Council has become the principal operating agency
of both the National Academy of Sciences and the National Academy of Engineering in provid-
ing services to the government, the public, and the scientific and engineering communities. The
Council is administered jointly by both Academies and the Institute of Medicine. Dr. Ralph J.
Cicerone and Dr. Wm. A. Wulf are chair and vice chair, respectively, of the National Research
Council.
www.national-academies.org
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COMMITTEE ON NUTRIENT REQUIREMENTS OF SMALL RUMINANTS
JAMES E. (ED) HUSTON, Chair, Texas A&M University, San Angelo
ROBERT G. WHITE, Vice-Chair, University of Alaska, Fairbanks
BRIAN J. BEQUETTE, University of Maryland, College Park
HUGH DOVE, Commonwealth Scientific and Industrial Research Organisation,
Canberra, Australia
ARTHUR L. GOETSCH, Langston University, Langston, Oklahoma
BRET W. HESS, University of Wyoming, Laramie
MAXIMINO HUERTA BRAVO, Universidad Autónoma Chapingo, Texcoco, Mexico
DAVID G. PUGH, Fort Dodge Animal Health, Waverly, Alabama
SANDRA G. SOLAIMAN, Tuskegee University, Alabama
Staff
AUSTIN J. LEWIS, Study Director
JAMIE S. JONKER, Study Director*
DONNA LEE JAMEISON, Senior Project Assistant**
RUTH S. ARIETI, Project Assistant
PEGGY TSAI, Research Associate
External Support
PAULA T. WHITACRE, Editor
*Through June 2004
**Through January 2006
v
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BOARD ON AGRICULTURE AND NATURAL RESOURCES
W. REG GOMES, Chair, University of California, Oakland
SANDRA J. BARTHOLMEY, University of Illinois at Chicago
ROGER N. BEACHY, Donald Danforth Plant Science Center, St. Louis, Missouri
H. H. CHENG, University of Minnesota, St. Paul
BRUCE L. GARDNER, University of Maryland, College Park, Maryland
JEAN HALLORAN, Consumer Policy Institute/Consumers Union, Yonkers, New York
HANS R. HERREN, Millennium Institute, Arlington, Virginia
KIRK C. KLASING, University of California, Davis
BRIAN W. MCBRIDE, University of Guelph, Guelph, Canada
TERRY L. MEDLEY, E. I. duPont de Nemours & Co., Wilmington, Delaware
ROBERT PAARLBERG, Wellesley College, Watertown, Massachusetts
ALICE N. PELL, Cornell University, Ithaca, New York
BOBBY PHILLS, Florida A&M University, Tallahassee
SONYA B. SALAMON, University of Illinois, Urbana-Champaign
HAL SALWASSER, Oregon State University, Corvallis, Oregon
PEDRO A. SANCHEZ, The Earth Institute at Columbia University, Palisades, New York
B. L. TURNER, II, Clark University, Worcester, Massachusetts
LAURIAN UNNEVEHR, University of Illinois at Urbana-Champaign
TILAHUN D. YILMA, University of California, Davis
JAW-KAI WANG, University of Hawaii, Honolulu
Staff
ROBIN A. SCHOEN, Director
KAREN L. IMHOF, Administrative Assistant
AUSTIN J. LEWIS, Program Officer
PEGGY TSAI, Research Associate
RUTH S. ARIETI, Project Assistant
vi
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Preface
Small ruminants, especially sheep and goats, are an im- elk [Cervus elaphus], and caribou/reindeer [Rangifer taran-
portant part of the economic and social life of many nations dus]), and New World camelids (alpacas [Lama glama] and
throughout the world. Numerous other species of small ru- llamas [Vicugna pacos]). The second important decision was
minants range the Earth in undomesticated herds, some of how to organize the publication--by species or by topic,
which are managed and others which are not. Information such as individual nutrients. Ultimately, the committee de-
about the dietary habits and nutrient requirements of these cided that to organize the report by species would result in
animals is crucial in managing their wellbeing and in con- a large amount of repetition and redundancy. Therefore, the
tributing to the livelihoods of people who depend on them. report is organized by topic. Thus, for example, there are
Proper nutrition and management of small ruminants is also chapters on individual nutrients, and the species are covered
essential for the maintenance of various natural ecosystems within each chapter. The committee recognized that this
from the Arctic Circle to the Sahara Desert. makes the publication somewhat less convenient for readers
The National Research Council has published reports on interested in one species only. To address this, tables of nu-
the Nutrient Requirements of Sheep (6th edition, 1985) and trient requirements are organized by species, and one chap-
the Nutrient Requirements of Goats (1st edition, 1981), but ter discusses practical feeding and nutrition on a species-by-
a large amount of new information has accumulated during species basis.
the last 25 years. Other small ruminants have not been ad- This report represents the concerted efforts of all com-
dressed in the Nutrient Requirements series. This new pub- mittee members. The committee was appointed in April
lication updates the sheep and goat publications and adds in- 2004 and held its first meeting in May 2004. After two more
formation about other small ruminant species. In addition, in-person meetings and several teleconferences, an initial
information about New World camelids (alpacas and llamas) draft was sent to reviewers in November 2005. The commit-
is included to the extent that it is available. tee hopes the publication will serve as an important refer-
The committee appointed to write this report faced two ence for all concerned with the nutrition, feeding, health,
important decisions at the outset. The first decision was and welfare of small ruminants and that it will also serve to
which species to include as "small ruminants." After consid- stimulate more research on this important group of animals.
ering various factors such as body size, amount of scientific
research data, absence of alternative sources of information,
and general interest of potential users, the committee de- ED HUSTON
cided to include the following species: sheep (Ovis aries), Chair, Committee on
goats (Capra hircus), cervids (white-tailed deer [Odocoileus Nutrient Requirements of
virginianus], red deer [Cervus elaphus], wapiti/American Small Ruminants
vii
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Acknowledgments
This report has been reviewed in draft form by persons Although the reviewers listed above have provided con-
chosen for their diverse perspectives and technical expertise structive comments and suggestions, they were not asked to
in accordance with procedures approved by the National endorse the conclusions or recommendations, nor did they
Research Council's Report Review Committee. The purpose see the final draft of the report before its release. The review
of this independent review is to provide candid and critical of this report was overseen by Michael L. Galyean, Texas
comments that will assist the institution in making its pub- Tech University. Appointed by the National Research Coun-
lished report as sound as possible and to ensure that the re- cil, he was responsible for making certain that an independ-
port meets institutional standards of objectivity, evidence, ent examination of this report was carried out in accordance
and responsiveness to the study charge. The review com- with institutional procedures and that all review comments
ments and draft manuscript remain confidential to protect were carefully considered. Responsibility for the final con-
the integrity of the deliberative process. We wish to thank tent of this report rests entirely with the author committee
the following individuals for their review of this report: and the institution.
We are grateful for contributions of Larry Miller, CSREES-
Antonello Cannas, University of Sassari, Sardinia, Italy USDA, who has been a good friend to the industries that will
George F. W. Haenlein, University of Delaware, Newark, be served by this report. Larry is an ever-present entity when-
DE (retired) ever the animal industries, the scientific community, and U.S.
Patrick G. Hatfield, Montana State University, Bozeman, MT governmental agencies interact and always a facilitator, a role
Robert J. Hudson, University of Alberta, Edmonton, that he fulfilled in the production of this document.
Alberta, Canada Members of the author committee express appreciation to
Lyndon N. Irwin, Southwest Missouri State University, the National Research Council and parent agency, The
Springfield, MO National Academies, for entrusting us to prepare this greatly
Woody Lane, Lane Livestock Services, Roseburg, OR anticipated and respected document. Special thanks are given
Jean-Marie Luginbuhl, North Carolina State University, to Charlotte Kirk Baer, former Director of the Board on
Raleigh, NC Agriculture and Natural Resources, for her initiation of this
Daniel G. Morrical, Iowa State University, Ames, IA project and early guidance, and to Robin Schoen, current
John V. Nolan, University of New England, Armidale, Director, for her administrative oversight of this process and
NSW, Australia continued encouragement. A deeply felt thank you goes to
Fredric N. Owens, Pioneer Hi-Bred International, Donna Lee Jameison, Senior Project Assistant, and to Ruthie
West Des Moines, IA Arieti, Project Assistant, for their technical contributions in
Rodney L. Preston, Texas Tech University, Lubbock, TX monitoring, formatting, editing, posting, and other responsi-
(retired) bilities, and for their extremely good nature while perform-
Duane E. Ullrey, Michigan State University, East Lansing, ing these vital services. Finally, the committee wishes to
MI (retired) thank Austin Lewis, Study Director, who led us on a daily
Larry W. Varner, Purina Mills Inc, Seguin, TX basis by giving us information, feedback, critique, and
encouragement and brought us to successful completion.
ix
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Contents
SUMMARY 1
1 INTRODUCTION 3
2 ANATOMY, DIGESTIVE PHYSIOLOGY, AND NUTRIENT UTILIZATION 5
3 INGESTIVE BEHAVIOR AND FEED INTAKE 26
4 ENERGY 39
5 PROTEIN 81
6 ESSENTIAL FATTY ACIDS 106
7 MINERALS 112
8 VITAMINS 150
9 WATER 173
10 PLANT FACTORS AFFECTING NUTRIENT AVAILABILITY 189
11 NUTRIENT SOURCES AND FEEDING PRACTICES 201
12 ENVIRONMENTAL IMPACT 216
13 NUTRITIONAL AND METABOLIC DISEASES, AILMENTS,
AND MODIFIERS 220
14 PROVIDING REQUIRED NUTRIENTS TO SMALL RUMINANTS 232
NUTRIENT REQUIREMENT TABLES 244
Nutrient Requirements of Sheep, 244
Nutrient Requirements of Goats, 271
Nutrient Requirements of Cervids, 300
Nutrient Requirements of New World Camelids, 308
xi
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xii CONTENTS
FEED COMPOSITION TABLES 312
Common Feedstuffs, 312
Novel Feedstuffs, 322
Mineral Supplements, 332
OTHER TABLES 335
Conversion Factors, SI Prefixes, and Termperature Conversions, 335
APPENDIXES
A COMMITTEE STATEMENT OF TASK 337
B ABBREVIATONS AND ACRONYMS 338
C COMMITTEE MEMBER BIOGRAPHIES 342
D BOARD ON AGRICULTURE AND NATURAL RESOURCES
PUBLICATIONS 344
INDEX 347
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Tables and Figures
TABLES
2-1 Digestible Nitrogen Intake and Urea Synthesis and Recycling in DifferentAnimal Species
(g N/day), 19
3-1 Selected Comparisons of Diet Compositions of Small Ruminants, 28
3-2 Potential Intakes (kg DM/day) of Twin-Suckling Ewes (Mature Weight 60 kg; Relative
Size 0.85), when Eating Grass Diets of 8, 10, or 12 MJ ME/kg DM during Lactation, 34
4-1 Predicted Empty Body Concentrations of Fat, Protein, and Energy in Sheep from Body
Condition Score, 44
4-2 Energy and Protein Requirements of Small Ruminants for Pregnancy, 53
4-3 References, Number of Treatment Mean Observations, and Genotypes in the Database
Used to Evaluate Prediction of Average Daily Gain, 55
4-4 Description of the Database Used to Evaluate Prediction of Average Daily Gain, 55
4-5 References, Number of Treatment Mean Observations, and Genotypes in the Database
Used to Evaluate Prediction of Net Energy for Lactation and Average Daily Gain, 61
4-6 Description of the Database Used to Evaluate Prediction of Average Daily Gain and Net
Energy for Lactation, 61
4-7 Adjustment Factors, 67
5-1 EssentialAminoAcid Composition (grams of amino acid per 100 grams protein) of Mixed
Microbial Protein Compared with Carcass Tissue, Milk Casein, Skin, and Wool, 88
5-2 Wool Protein Composition (grams of amino acid per 100 grams protein), 89
5-3 References, Number of Treatment Mean Observations, and Genotypes in the Database
Used to Evaluate Metabolizable Protein Requirements of the CNCPS-S Method, 98
5-4 Description of the Database Used to Evaluate Metabolizable Protein (MP) Requirements
of the CNCPS-S Method, 99
7-1 Copper Absorption Based on Normal Molybdenum and Sulfur Content of the Diet, 128
9-1 Comparison of Estimates of Daily Water Requirements of Goats, 184
10-1 Summary of Some Observed Positive and Negative Effects of Plant Secondary Metabo-
lites after Consumption by Mammalian Herbivores, 191
10-2 Fractionation of Forage Organic Matter According to the Detergent System, 194
10-3 Estimated Digestibility of Neutral Detergent Solubles in a Range of Grass Species, 196
11-1 Long-Chain Fatty Acids in Some Feedstuffs Fed to Small Ruminants, 204
15-1 Nutrient Requirements of Sheep (mature ewes and rams and yearlings maintenance and
lactation), 246
xiii
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xiv TABLES AND FIGURES
15-2 Nutrient Requirements of Sheep (growing, finishing lambs, and yearlings during
growth/gestation), 256
15-3 Mineral Requirements of Sheep, 266
15-4 Nutrient Requirements of Meat and Milk Goats (maintenance-gestation-lactation), 272
15-5 Nutrient Requirements of Goats (growing kids), 281
15-6 Nutrient Requirements of Goats (Angora), 288
15-7 Mineral Requirements of Meat and Milk Goats, 293
15-8 Mineral Requirements of Goats (Angora), 297
15-9 Nutrient Requirements of Cervid Species (white-tailed deer, reindeer/caribou, red deer
and elk/wapiti), 301
15-10 Nutrient Requirements of New World Camelids, 309
15-11 Composition of Common Feedstuffs, 312
15-12 Composition of Pasture or Range Forage, Browse, and Other Novel Feedstuffs, 322
15-13 Compositions of Inorganic Mineral Sources on a 100% Dry Matter Basis, 332
15-14 Table of Conversion Factors, 335
FIGURES
2-1 Drawings of the gastrointestinal tracts of the sheep (a grass/roughage eater morphologi-
cal feeding type) and the llama, 6
2-2 A drawing of the red deer (an intermediate morphological feeding type) showing the
stomach in situ and isolated, 7
2-3 A drawing of the right aspect of the stomach of the goat (an intermediate morphological
feeding type), 7
2-4 Photograph of the four stomach compartments of a yearling white-tailed deer (a concen-
trate selector morphological feeding type), 8
2-5 The inside structure of the four stomach compartments of a white-tailed deer (a concen-
trate selector morphological feeding type), 8
2-6 Drawings of the three stomach compartments of the llama, 9
3-1 Influence of standard reference weight of sheep (mature size when in average condition
score) and relative size (current weight as a proportion of mature weight) on predicted
intakes of an all-grass diet of nonlimiting digestibility (> 0.8), 33
3-2 Influence of relative size (proportion of mature live weight) on predicted DM intakes, ex-
pressed in relation to mature size, 33
3-3 Influence of time since lambing on the predicted DM intake (% of standard reference
weight) of ewes of relative size 0.85 consuming an all-grass diet and suckling either sin-
gle or twin lambs, 34
3-4 Pasture intakes (kg DM/day) predicted for sheep of standard reference weight 50 kg,
grazing improved grass pastures differing in average digestibility, 35
4-1 The relationship between observed ADG (ADGobs) and ADG predicted (ADGpred) by the
NRC method for growing/finishing sheep, 57
4-2 The relationship between observed ADG (ADGobs) and ADG predicted (ADGpred) by the
CNCPS-S method for growing/finishing sheep, 57
4-3 The relationship between observed ADG (ADGobs) and ADG predicted (ADGpred) by the
NRC method for growing finishing/sheep, 58
4-4 The relationship between observed ADG (ADGobs) and ADG predicted (ADGpred) by the
CNCPS-S method for growing/finishing sheep, 59
4-5 The relationship between observed ADG (ADGobs) and ADG predicted (ADGpred) by the
CNCPS-S method for lactating sheep, 62
4-6 The relationship between observed ADG (ADGobs) and ADG predicted (ADGpred) by the
CNCPS-S method for lactating sheep, 63
4-7 The relationship between observed NEl (NEl-obs) and NEl predicted (NEl-pred) by the
CNCPS-S method for lactating sheep, 63
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TABLES AND FIGURES xv
4-8 The relationship between observed NEl (NEl-obs) and NEl predicted (NEl-pred) by the
CNCPS-S method for lactating sheep, 64
5-1 The relationship between metabolizable protein (MP) available (based on the difference
between predicted metabolizable protein intake and metabolizable protein used for main-
tenance; MPg-avail) and used for gain or maintenance (based on predicted metabolizable
protein accreted in tissue or mobilized and used for maintenance; MPg-used), 91
5-2 The relationship between rumen degraded intake protein (DIP) as a percent of total di-
gestible nutrients (TDN) and average daily gain (ADG) relative to body weight (BW)0.75
with a database of treatment mean observations from the literature for growing goats, 95
5-3 The relationship between rumen degraded intake protein (DIP) as a percent of total di-
gestible nutrients (TDN) and average daily gain (ADG) relative to body weight (BW)0.75
with a database of treatment mean observations from the literature for mature goats, 96
5-4 The relationship between rumen degraded intake protein (DIP) as a percent of total di-
gestible nutrients (TDN) and dry matter intake (DMI) relative to body weight (BW)0.75
with a database of treatment mean observations from the literature for growing goats, 96
5-5 The relationship between rumen degraded intake protein (DIP) as a percent of total di-
gestible nutrients (TDN) and dry matter intake (DMI) relative to body weight (BW)0.75
with a database of treatment mean observations from the literature for mature goats, 96
7-1 Selenium requirements of growing lambs based on performance and selenium content of
carcass, 136
10-1 Relationship between plant carbohydrate fractions and analytical components used to de-
scribe them, 194
10-2 Typical changes in dry matter digestibility of a range of pasture grasses (upper panel) and
legumes (lower panel) as spring progresses, 195
10-3 Changes in the digestibility of different plant parts of annual grass pasture during spring-
autumn, 195
11-1 Annual weight change for single-bearing ewe, 208
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