Published 1993

Forage Cell Wall Structure

and Digestibility
 Forage Cell Wall Structure
and Digestibility
Editors
H. G. lung, D. R. Buxton, R. D. Hatfield, and l. Ralph

Based on the International Symposium on Forage Cell Wall

Structure and Digestibility sponsored by the USDA-Agricultural
Research Service and the U.S. Dairy Forage Research Center,
Madison, Wisconsin, 7-10 Oct. 1991.

Organizing Committee
R. D. Hatfield D. R. Buxton
H. G. lung D. R. Mertens
1. Ralph P. 1. Weimer

Editor-in-Chief ASA
G. A. Peterson

Editor-in-Chief CSSA
P. S. Baenziger

Editor-in-Chief SSSA
R. 1. Luxmoore

Managing Editor
S. H. Mickelson

American Society of Agronomy, Inc.

Crop Science Society of America, Inc.
Soil Science Society of America, Inc.
Madison, Wisconsin, USA
1993
Cover Design: Patricia J. Scullion
Cover Artwork: Courtesy of USDA, 1948 Yearbook of Agriculture,
p.659.

Copyright 1993 by the American Society of Agronomy, Inc.

Crop Science Society of America, Inc.
Soil Science Society of America, Inc.

ALL RIGHTS RESERVED UNDER THE U.S. COPYRIGHT

ACT OF 1976 (P.L. 94-553)

Any and all uses beyond the limitations of the "fair use" provision
of the law require written permission from the publisher(s) and/or
the author(s); not applicable to contributions prepared by officers or
employees of the U.S. Government as part of their official duties.

American Society of Agronomy, Inc.

Crop Science Society of America, Inc.
Soil Science Society of America, Inc.
677 South Segoe Road, Madison, WI 53711 USA

Library of Congress Cataloging-in-Publication Data

International Symposium on Forage Cell Wall Structure and Digesti-

bility (1991 : Madison, Wis.)
Forage cell wall structure and digestibility / editors, H.G. Jung
... let al.l.
p. cm.
"Based on the International Symposium on Forage Cell Wall Struc-
ture and Digestibility, sponsored by the USDA-Agricultural Research
Service and the U.S. Dairy Forage Research Center, Madison, Wiscon-
sin, 7-10 Oct. 1991."
Includes bibliographical references and index.
ISBN 0-89118-115-6
1. Lignocellulose in animal nutrition-Congresses. 2. Forage
plants-Cytology-Congresses. 3. Plant cell walls-Congresses.
4. Ruminants-Feeding and feeds-Congresses. 5. Digestion-Con-
gresses. 6. Rumen fermentation-Congresses. 7. Feed utilization
efficiency-Congresses. I. Jung, H.G. II. United States. Agricultur-
al Research Service. III. U.S. Dairy Forage Research Center. IV. Title.
SF98.L54I57 1991
636.2'0855-dc20 93-12202
CIP

Printed in the United States of America

 CONTENTS

Page
FOREWORD ..................................................... xiii
PREFACE........................................................ xv
CONTRIBUTORS ................................................. xvii
CONVERSION FACTORS FOR SI AND NON-SI UNITS............. XXI

SECTION I. PERSPECTIVE
1 Organization of Forage Plant Tissues
J. R. WILSON
I. The Cell Wall 1
II. Tissue Types ............................................ . 3
III. Tissue Proportions in Organs ............................. . 14
IV. Overcoming Plant Structure Limitations .................... . 18
Acknowledgment ............................................. . 27
References ................................................... . 27

2 Utilization of Forage Fiber by Ruminants 33

M. L. GAL YEAN AND A. L. GOETSCH
I. Forages in Ruminant Production ........................... 34
II. Types of Forages and Their Utilization ............... . . . . . . . 39
III. Forages in Mixed Diets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
IV. Other Factors Affecting Forage Utilization. . . . . . . . . . . . . . . . . . 52
V. Other Roles of Forages ................................... 57
VI. Current Limitations to Forage Use ......................... 60
VII. Opportunities for Increased Forage Use ..................... 61
VIII. Opportunities for Greater Animal Productivity from
Forages ................................................. 62
References .................................................... 62

3 Perspectives of Cell Wall Biodegradation-Session Synopsis 73

DANNY E. AKIN
I. Animal Aspects in Plant Cell Wall Utilization .............. . 74
II. Structure and Bioavailability of Plant Cell Walls ............ . 76
III. Rumen Microorganisms and Plant Cell Wall Degradation .... . 77
IV. Conclusion ............................................. . 79
References 80
v
vi CONTENTS

SECTION II. ANALYTICAL TECHNIQUES FOR PLANT CELL

WALLS

4 Quantitative Analysis of Cell Wall Components 83

OLOF THEANDER AND ERIC WESTERLUND
I. Gravimetric Methods for Determination of Total Fiber ....... 84
II. Extraction and Purification of Cell Wall Components. . . . . . . . 85
III. The Uppsala Method for Cell Wall Analysis. . . . . . . . . . . . . . . . . 89
IV. Comparison between the Detergent Gravimetric Methodology
and the Uppsala Method for Cell Wall Analysis. . . . . . . . . . . . . . 93
V. Alternatives for Determining Cell Wall Constituents. . . . . . . . . . 97
VI. Conclusion .............................................. 98
References .................................................... 99

5 Analysis of Forage Cell Wall Polysaccharides 105

DONALD J. NEVINS
I. Preparation of Samples for Analysis ........................ 106
II. Preparation of Soluble Cell Wall Fractions. . . . . . . . . . . . . . . . . . 108
III. Dissociation of Components from the Wall by Specific
Enzymes ................................................ 111
IV. Intermolecular Associations and Spatial Distribution
of Polysaccharides ........................................ 116
References .................................................... 129

6 Application of New Methods for the Investigation of Lignin Structure 133

CATHERINE LAPIERRE
I. Structural Screening and Molecular Profiling of Lignins
and Lignocellulosics by Analytical Pyrolysis ................. 135
II. The Lignin Carbon Skeleton Traced Out by Carbon-13
Nuclear Magnetic Resonance Spectroscopy .................. 140
III. Close Exploration of Lignin Structure by Chemical
Degradation Methods: The Case of Thioacidolysis ............ 150
IV. Conclusion .............................................. 160
Acknowledgment .............................................. 163
References .................................................... 163

7 Analysis of Plant Cell Walls-Session Synopsis 167

JOHN R. OBST
I. Lignin Distribution in Wood ............................... 169
II. Lignin Distribution in Nonwood Plants..................... 177
III. Conclusion .............................................. 178
References .................................................... 179
CONTENTS vii

SECTION III. STRUCTURE OF FORAGE CELL WALLS

8 Composition and Structure of Cell Wall Polysaccharides in Forages 183

PER AM AN
I. Fractionation ............................................ 183
II. Structural Analysis ....................................... 185
III. Structure of Cell Wall Polysaccharides ...................... 187
IV. Composition of Cell Wall Polysaccharides in Grasses. ..... ... 191
V. Composition of Cell Wall Polysaccharides in Dicots .......... 194
VI. Conclusion .............................................. 195
References .................................................... 196

9 Lignin/Hydroxycinnamic Acid/Polysaccharide Complexes: Synthetic

Models for Regiochemical Characterization 201
JOHN RALPH AND RICHARD F. HELM
I. Linkages between Polysaccharides and Hydroxycinnamic
Acids ................................................... 203
II. Structure of Forage Lignin/Hydroxycinnamic Acid
Complexes ............................................... 210
III. Conclusion .............................................. 240
Acknowledgment .............................................. 241
References .................................................... 241

10 Comprehensive Model of the Lignified Plant Cell Wall 247

N. TERASHIMA, K. FUKUSHIMA, L-F. HE, AND K. TAKA BE
I. Deposition of Carbohydrates During the Cell Wall
Differentiation ........................................... 248
II. Deposition of Lignin During the Cell Wall Differentiation ..... 251
III. Comprehensive Structural Model of Lignified Cell Wall
Based on Biogenesis ...................................... 257
IV. Formation and Structure of Cell Wall in Graminaceous
Plants ............. " ..................... " . . .. . . . . . . .. . 263
V. Concluding Remarks and Future Aspects .................... 266
References .................................................... 266

11 Structure of Forage Cell Walls-Session Synopsis 271

DAVID S. HIMMELSBACH
I. Scope ................................................... 271
II. Cell Wall Polysaccharides ................................. 272
III. Lignin/Phenolic-Carbohydrate Complexes ................... 274
IV. Lignified Plant Cell Wall. .. ... ... . . .... ... .. . ... . .. .. . . . .. 275
V. Nondestructive Spectroscopic Methods of Analysis ........... 276
VI. Chemical Microscopy ..................................... 277
VII. Discussion ............................................... 279
VIII. Conclusion .............................................. 280
References .................................................... 280
viii CONTENTS
SECTION IV. CELL WALL MATRIX INTERACTIONS
AND DEGRADATION
12 Cell Wall Polysaccharide Interactions and Degradability 285
RONALD D. HATFIELD
I. Cell Wall Matrix Organization (Component Interactions) ...... 286
II. Cell Wall Degradation .................................... 297
III. Future Direction ......................................... 304
IV. Conclusion .............................................. 307
References .................................................... 307

13 Cell Wall Lignification and Degradability 315

H. G. lUNG AND D. A. DEETZ
I. Lignin Terminology in Forages .......... . . . . . . . . . . . . . . . . . . . 316
II. Empirical Relations ....................................... 317
III. Molecular Mechanisms of Inhibition. . . . . . . . . . . . . . . . . . . . . . . . 327
IV. Model of Cell Wall Lignification and Degradability .......... 332
V. Brown Midrib Mutants: A Case Study. . . . . . . . . . . . . . . . . . . . . . . 337
VI. Conclusion .............................................. 339
References .................................................... 340

14 Mechanistic Models of Forage Cell Wall Degradation 347

ANDREW CHESSON
I. Chemical Composition of Forage Cell Walls. . . . . . . . . . . . . . . . . 348
II. Cell Wall Architecture .................................... 353
III. The Lignin-Carbohydrate Complex as a Unit of Secondary
Cell Wall Structure ....................................... 358
IV. Modelling the Degradation of Individual Cell Walls.......... 362
V. Manipulation of Cell Wall Degradability..................... 367
VI. Integration with Models of Rumen Function ................. 369
References .................................................... 371

15 Cell Wall Matrix Interactions and Degradation-Session Synopsis 377

P. l. VAN SOEST
I. The Concept of Nutritional Uniformity ..................... 379
II. Dietary Fibers ............................................ 380
III. Carbohydrate Fractions-Uronides ......................... 381
IV. Lignin .................................................. 382
V. Other Limiting Factors .................................... 389
VI. Commentary on Models ................................... 390
VII. Discussion ............................................... 391
VIII. Conclusions and Recommendations. . . . . . . . . . . . . . . . . . . . . . . . . . 392
References .................................................... 393
CONTENTS ix

SECTION V. MICROBIAL AND MOLECULAR MECHANISMS

OF CELL WALL DEGRADATION

16 Microbial Adhesion and Degradation of Plant Cell Walls 397

ALICE N. PELL AND PETER SCHOFIELD
I. Adhesion Mechanisms .................................... 398
II. Adhesion of Cellylolytic Bacteria ........................... 403
III. Adhesion of Ruminal Protozoa ............................ 413
IV. Adhesion of Ruminal Fungi.... . ....... ................. .. 414
V. Summary. ... ................ ............... ............. 415
References .................................................... 416

17 Microbial Ecology of Cell Wall Fermentation 425

BURK A. DEHORITY
I. Digestion of Forage Cell Walls by Rumen
Microorganisms .......................................... 427
II. Rumen Microbes That Can Digest the Structural
Polysaccharides in Plant Cell Walls ......................... 429
III. Digestion of Structural Polysaccharides from Intact Plant
Cell Walls ............................................... 433
IV. Synergism between Microbial Species ....................... 438
V. Negative Effects between Microbial Species .................. 443
VI. Relationship of Plant Structure to Digestibility and
Synergism ............................................... 444
VII. Rumen Microorganisms That Use Storage Polysaccharides
or Intermediate Products of Structural
Polysaccharide Digestion .................................. 445
VIII. Conclusion .............................................. 446
References .................................................... 449

18 Enzymatic Hydrolysis of Forage Cell Walls 455

BRYAN A. WHITE, RODERICK I. MACKIE,
AND KINCHEL C. DOERNER

I. Enzymatic Activities of Microorganisms Involved in Plant

Cell Wall Hydrolysis in the Rumen. . . . . . . . . . . . . . . . . . . . . . . . . 456
II. Features of the Enzymatic Degradation of Plant Cell
Walls by Ruminal Microorganisms ......................... 461
III. A Conceptual Model for Hydrolysis of Forage Cell Walls 469
IV. Molecular Biology and the Understanding of Plant Cell
Wall Degradation in the Rumen 473
V. What Can and Needs to be Learned, and Can These Plant
Cell Wall Hydrolyzing Systems be Manipulated? ............. 476
VI. Concluding Remarks ...................................... 478
Acknowledgment .............................................. 478
References .................................................... 479
x CONTENTS

19 Microbial and Molecular Mechanisms of Cell Wall Degradation-

Session Synopsis 485
PAUL J. WEIMER
I. Microbiology and Enzymology of Cell Wall Digestion:
Where Reductionism Reigns ............................... 485
II. The Role of Microbial Adhesion to Plant Fiber in the
Digestion Process ......................................... 486
III. Microbial Ecology of Forage Cell Wall Digestion
in Ruminants ............................................ 489
IV. Enzymology of Forage Cell Wall Digestion. . . . . . . . . . . . . . . . . . 492
V. Genetic Engineering of Ruminal Microbes ................... 495
VI. Concluding Remarks ...................................... 496
Acknowledgment .............................................. 497
References .................................................... 497

SECTION VI. CELL WALL DEGRADATION IN THE RUMINANT

20 Particle-Size Reduction by Ruminants-Effects of Cell Wall
Composition and Structure 499
P. M. KENNEDY AND P. T. DOYLE
I. General Importance of Particle Kinetics ..................... 500
II. Putative Role of Particle Size on Regulation of Voluntary
Intake................................................... 501
III. Microbes Associated with Plant Residues in the
Reticulorumen ........................................... 502
IV. Movement of Particles In, and From, the Reticulorumen ...... 502
V. Methodological Considerations in Particle Kinetics Studies .... 511
VI. Processes and Extent of Particle Comminution. . . . . . . . . . . . . . . 518
VII. Do Constraints Associated with Particle Comminution
Limit Intake? ............................................ 524
VIII. Conclusions and Future Directions ......................... 527
References .................................................... 529

21 Kinetics of Cell Wall Digestion and Passage in Ruminants 535

D. R. MERTENS
I. Kinetics of Digestion ...................................... 538
II. Kinetics of Passage ....................................... 554
III. Integrated Models of Digestion and Passage ................. 561
IV. Relative Importance of Kinetic Variables. . . . . . . . . . . . . . . . . . . . 562
References .................................................... 566

22 Influence of Feeding Management on Ruminant Fiber Digestibility 571

S. TAMMINGA
I. Fiber Digestion Models: Competition between Rates
of Degradation and Rate of Passage ........................ 572
II. Microbiological, Physical, and Physiological Factors
Affecting Fiber Digestion .................................. 573
III. Interaction with Other Dietary Components ................. 579
CONTENTS xi

IV. Feeding Management ..................................... 587

V. Opportunities to Increase Amount of Forage Fed and
Energy Availability from Fiber ............................. 593
VI. Conclusion .............................................. 596
References .................................................... 596

23 Cell Wall Degradation in the Ruminant-Session Synopsis 603

BARBARA P. GLENN AND DALE R. WALDO
I. Challenges of Particle-Size Reduction ....................... 603
II. Challenges of Kinetics of Digestion and Passage ............. 605
III. Challenges of Feeding Management. . . . . . . . . . . . . . . . . . . . . . . .. 607
IV. Importance of Cell Walls in Energetic Efficiency. . . . . . . . . . . . .. 609
V. Future Research and Conclusions. . . . . . . . . . . . . . . . . . . . . . . . . .. 618
References .................................................... 618

SECTION VII. MECHANISMS FOR ALTERING CELL WALL

UTiLiZA TION

24 Cell Wall Biosynthesis and Its Regulation 621

K. IIYAMA, T. B. T. LAM, P. J. MEIKLE, K. NG,
D. I. RHODES, AND B. A. STONE
I.Pathways for Wall Polysaccharide Synthesis ................. 621
II.Pathways for Lignin Biosynthesis .......................... , 635
III.Pathways to Cutin and Suberin ............................ 648
IV. Wall Protein and Glycoprotein Biosynthesis ................. 649
V. Spatial and Temporal Events in Wall Polymer Synthesis
and Their Regulation ..................................... 652
Acknowledgments .............................................. 665
References .................................................... 665

25 Environmental and Genetic Effects on Cell Wall Composition and

Digestibility 685
D. R. BUXTON AND M. D. CASLER
I. Variation Among Plant Parts and Plant Species. . . . . . . . . . . . . . 686
II. Effect of Plant Environment ............................... 688
III. Genetic Regulation and Breeding for Improved Forage
Digestibility .............................................. 694
IV. Potentially Negative Effects of Breeding for Improved
Digestibility .............................................. 705
V. Future Research Needs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 707
References .................................................... 708

26 Postharvest Treatment of Fibrous Feedstuffs to Improve Their

Nutritive Value 715
G. C. FAHEY, JR., L. D. BOURQUIN, E. C. TITGEMEYER,
AND D. G. ATWELL

I. Physical Treatments ...................................... 717

II. Chemical Treatments ..................................... 730
xii CONTENTS

III. Microbial and Enzymatic Treatments ....................... 749

IV. Conclusion .............................................. 757
References .................................................... 758

27 Mechanisms for Altering Cell Wall Utilization-Session Synopsis 767

J.C.BURNS
I. Control Points ........................................... 767
II. Environmental Influence .................................. 770
III. Utilization ............................................... 771
IV. Areas for Future Research. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 774
References .................................................... 776

Subject Index ..................................................... 779

 FOREWORD
It is critical that the efficient use of plant resources be increased to meet
the food and fiber needs of the future. Achievement of this goal will require
that use of forages for livestock production become even more of a feed
resource. However, there is a critical lack of information on the chemistry
of the cell walls of forage plants and especially how this chemistry relates
to digestibility and absorption of the materials and how the digestibility can
be improved through plant breeding.
This book presents the results of a benchmark conference that brought
together scientists from many disciplines ranging from plant cell wall chemis-
try to digestibility. The conference format permitted wide-ranging discus-
sions and they are also captured in the book. The Tri-societies are extremely
grateful to the USDA Dairy Forage Research Center and USDA-Agricultural
Research Service for their support and serving as excellent hosts. The sym-
posium could not have been done without the excellent organizational ef-
forts of Drs. R.D. Hatfield, D.R. Buxton, H.G. Jung, D.R. Mertens, J.
Ralph, and P.J. Weimer.
The Tri-societies are excited about this book and about the possibilities
it brings forth to advance humankind through wise use of our forage
resources.

D. R. KEENEY, president
American Society of Agronomy

C. W. STUBER, president
Crop Science Society of America

D.W. NELSON, president

Soil Science Society of America

xiii
 PREFACE
The chapters for this book were prepared in conjunction with the International
Symposium on Forage Cell Wall Structure and Digestibility, held in Madison, WI
on 7-10 Oct. 1991. About 165 scientists from 15 nations attended the symposium.
In addition to 20 invited speakers and seven session moderators, whose chapters are
presented in this book, the symposium was enriched by the presentation of 46 papers
by participants during two evening poster sessions. The symposium was organized
into seven half-day sessions, with three presentations in each session. An extended
discussion period concluded each session and augmented the regular question-and-
answer period after individual presentations. The format of this book follows the
symposium agenda with the speaker chapters from each session followed by a syn-
opsis chapter prepared by the session moderator. Both the speaker and the session
moderator chapters have been revised or developed to include important points raised
in the discussion periods.
The genesis of the symposium was a question T.K. Kirk posed to the individuals
who later became the organizing committee. He asked if there was a recent confer-
ence book that summarized the state-of-knowledge on the subject of forage cell wall
chemistry and its digestibility by ruminant animals. While there have been confer-
ences in recent years that have addressed aspects of this topic, we did not feel that
a comprehensive reference volume was available that covered forage cell walls and
their digestibility in sufficient breadth or depth. The symposium was subsequently
organized with one of the goals being the production of an up-to-date reference text.
The objective of the symposium was to assemble a multidisciplinary group of
expert scientists to present state-of-the-art information on forage cell wall structure
and digestibility, and to identify limits to our knowledge where more research is re-
quired. It was hoped that this diverse group of individuals could develop a bench-
mark reference publication that describes current knowledge regarding the chemical
structure and digestibility of forage cell walls. The chapter authors were asked to
review the various aspects of this subject and present not only the current state of
the art, but also to provide their personal vision of fruitful future areas of research
that will significantly expand our understanding. Several authors accepted the challenge
to develop conceptual models related to wall structure and development, and wall
degradation in the rumen. The opinions, conclusions, and hypotheses of this collec-
tion of experts from around the world have been compiled in this book. The editori-
al committee wishes to express its gratitude to the authors for their outstanding efforts.
Another goal of the symposium organizing committee was to identify cross-cutting
research topics where critical information is needed to improve forage cell wall diges-
tibility by ruminants. To this end, the symposium speakers and moderators held a
workshop on 10 October upon conclusion of the symposium's general sessions. The
assembled group of scientists developed and refined the following list of research
questions that are addressed in the chapters of this book, but where more informa-
tion is needed.
1. What changes in structural and conformational arrangements of cell wall poly-
saccharides and lignin will lead to increased cell wall digestibility?
2. Must lignin be cross-linked by chemical bonds to cell wall polysaccharides
to limit their digestibility?
3. Which factors within plant cells control lignification and how can these be
modified, genetically or otherwise, to affect the concentration and chemical struc-
ture of lignin?
xv
xvi PREFACE

4. Can microbial or animal genetic modifications increase the amount of net

energy available to animals from cell walls? If so, which modifications will be most
effective?
5. Are numbers, species, and activities of ruminal microorganisms limiting for
forage cell wall digestibility, and how can these be adequately measured in vivo?
6. What are the rate-limiting steps and necessary components for enzymatic
hydrolysis of cell walls by ruminal microorganisms and what methods can be used
to identify these steps?
7. Is availability of adequate forage cell wall surface for colonization by rumi-
nal microorganisms a major deterrent to digestibility of cell walls?
8. Can lignified cell walls of forages be digested from the exterior side of the
cell, or are they digested only from the lumen side outwards by ruminal microor-
ganisms?
9. What plant modifications will lead to increased particle-size reduction in the
rumen and how will these affect the amount of net energy available to animals from
forage cell walls?
10. How will changes in potential rate and extent of forage cell wall digestion
affect the amount of net energy available to animals that consume it?
11. Will improvement in forage cell wall digestibility have effects on yield, vigor,
and ability of forages to withstand stressful environments and pests?
These topics for research represent a group of general questions about cell walls
and their digestibility that have the potential for identifying strategies for the im-
provement of ruminant animal performance when fed high forage diets.
The editorial committee wishes to thank D.R. Mertens and P.I. Weimer for their
assistance in organizing the symposium. Financial and secretarial support for the sym-
posium by the USDA-Agricultural Research Service and the U.S. Dairy Forage
Research Center, and the continued support and encouragement of L.D. Satter, Direc-
tor of the U.S. Dairy Forage Research Center, were greatly appreciated and instrumen-
tal to the symposium's success. The opening remarks at the symposium by R.R. Oltjen,
Associate Deputy Administrator for the USDA-ARS, and F.C. Greene, Assistant
Midwest Area Director for the USDA-ARS, are gratefully acknowledged. The com-
mittee members also express their appreciation to G.C. Marten, Associate Beltsville
Area Director for the USDA-ARS, for his banquet speech entitled, "The Importance
of Unlocking the Secrets Held with the Walls of Forages." The decision by the Ameri-
can Society of Agronomy the Crop Science Society of America, and the Soil Science
Society of America to undertake responsibility for publication of this book has en-
sured production of this high-quality text.
Finally, we wish to thank all the participants at the symposium for their com-
ments, questions, and criticisms. Their input has resulted in significant improvements
in the material presented in this book.

H.G. lung, co-editor R.D. Hatfield, co-editor

USDA-ARS USDA-ARS
Univ. of Minnesota U.S. Dairy Forage Res. Ctr.
St. Paul, Minnesota Madison, Wisconsin

D.R. Buxton, co-editor J. Ralph, co-editor

USDA-ARS USDA-ARS
Iowa State University U.S. Dairy Forage Res. etr.
Ames, Iowa Madison, Wisconsin
 CONTRIBUTORS

Danny E. Akin Research Microbiologist, USDA-ARS, Russell Research Center,

Athens, GA 30613

Per Aman Professor of Plant Products, Department of Food Science, Swed-

ish University of Agricultural Sciences, S-75OO7 Uppsala, Sweden

D. G. Atwell Technical Services and Research, Feed Division, Growmark, Inc.,

Bloomington, IL 61702. Formerly Graduate Research Assistant,
Department of Animal Sciences, University of Illinois, Urbana, IL
61801

L. D. Bourquin Postdoctoral Research Associate, Department of Food Science and

Human Nutrition, Michigan State University, East Lansing, MI
48824-1224. Formerly Graduate Research Assistant, Division of
Nutritional Sciences, University of Illinois, Urbana, IL 61801

J. C. Burns Lead Scientist, Crop Science and Animal Science, Department of

Crop Science, North Carolina State University, Raleigh, NC
27695-7620

D. R. Buxton Research Leader, USDA-ARS, Field Crops Research Unit, Iowa

State University, Ames, IA 50011

M. D. Casler Professor of Plant Breeding and Biometry, Department of Agrono-

my, University of Wisconsin-Madison, Madison, WI 53706-1597

Andrew Chesson Head, Nutritional Sciences Division, Rowett Research Institute,

Greenburn Road, Bucksburn, Aberdeen AB2 9SB, U.K.

D. A. Deetz Graduate Research Assistant, Department of Animal Science,

University of Minnesota, St. Paul, MN 55108

Burk A. Dehority Professor and Associate Chair of Animal Science, Department of

Animal Science, The Ohio State University, Ohio Agricultural
Research and Development Center, Wooster, OH 44691

Kinchel C. Doerner Postdoctoral Research Associate, Department of Microbiology and

Immunology, Medical College of Virginia, Richmond, VA
23298-0678. Formerly Postdoctoral Research Associate, Department
of Animal Science, University of Illinois, Urbana, IL 61801

P. T. Doyle Research Officer, Western Australia Department of Agriculture, Al-

bany 6330, Western Australia

G. C. Fahey, Jr. Professor of Animal Sciences and Nutritional Sciences, Department

of Animal Sciences, University of Illinois, Urbana, IL 61801

K. Fukushima Research Associate of Chemistry of Forest Products, Faculty of

Agriculture, Nagoya University, Nagoya 464-01, Japan

M. L. Galyean Professor, New Mexico State University, Clayton Livestock Research

Center, Clayton, NM 88415
xvii
xviii CONTRIBUTORS

Barbara P. Glenn Research Animal Scientist, USDA-ARS, BARC-East, Beltsville, MD

20705

A. L. Goetsch Associate Professor of Animal Sciences, Department of Animal

Sciences, University of Arkansas, Fayetteville, AR 72701

Ronald D. Hatfield Research Plant Physiologist, USDA-ARS, U.S. Dairy Forage

Research Center, Madison, WI 53706

L-F. He Research Associate, Institute of Biological Chemistry, Washington

State University, Pullman, WA 99163

Richard F. Helm Research Associate, USDA-ARS, U.S. Dairy Forage Research

Center, Madison, WI 53706

David S. Himmelsbach Research Chemist, USDA-ARS, Russell Research Center, Athens,

GA 30613

K.Iiyama Research Fellow, Department of Biochemistry, La Trobe Universi-

ty, Bundoora VIC 3083, Australia

H. G. Jung Research Dairy Scientist, USDA-ARS, University of Minnesota, St.

Paul, MN 55108

P. M. Kennedy Principal Research Scientist, CSIRO, Indooroopilly, Queensland

4068, Australia

T.B. Lam Research Fellow, Department of Biochemistry, La Trobe Universi-

ty, Bundoora VIC 3083, Australia

Catherine Lapierre Laboratoire de Chimie Biologique, Institut National Agronomique,

78850 Thiverval-Grignon, France

Roderick I. Mackie Professor of Microbiology, Department of Animal Sciences, Univer-

sity of Illinois, Urbana, IL 61801

P. J. Meikle Senior Research Officer, Department of Biochemistry, La Trobe

University, Bundoora VIC 3083, Australia

D. R. Mertens Research Dairy Scientist, USDA-ARS, U.S. Dairy Forage Research

Center, Madison, WI 53706

Donald J. Nevins Professor of Plant Physiology, Department of Vegetable Crops,

University of California, Davis, CA 95616

K. Ng Research Fellow, Department of Biochemistry, La Trobe Universi-

ty, Bundoora VIC 3083, Australia

John R. Obst Research Chemist, USDA-FS, Forest Products Laboratory, Madi-

son, WI 53705

Alice N. Pell Associate Professor, Department of Animal Science, Cornell Univer-

sity, Ithaca, NY 14853

John Ralph Rsearch Chemist, USDA-ARS, U.S. Dairy Forage Research Center,
Madison, WI 53706

D. I. Rhodes Postgraduate Student, Department of Biochemistry, La Trobe

University, Bundoora VIC 3083, Australia
CONTRIBUTORS xix

Peter Schofield Research Associate, Department of Animal Sciences, Cornell Univer-

sity, Ithaca, NY 14853

B. A. Stone Professor of Biochemistry, Department of Biochemistry, La Trobe

University, Bundoora VIC 3083, Australia

K. Takabe Instructor, Department of Wood Science and Technology, Faculty

of Agriculture, Kyoto University, Kyoto 606-01, Japan

Seerp Tamminga Department of Animal Nutrition, Agricultural University, Haag-

steeg 4, Wageningen 6708 PM, Netherlands

N. Terashima Professor of Chemistry of Forest Products, Emeritus Professor of

Nagoya University, Faculty of Agriculture, Nagoya University,
Nagoya 464-01, Japan

Olof Theander Professor, Department of Chemistry, Swedish University of Agricul-

tural Sciences, S-750 07 Uppsala, Sweden

E. C. Titgemeyer Assistant Professor of Animal Sciences and Industry, Kansas State

University, Manhattan, KS 66506. Formerly Postdoctoral Research
Associate, Department of Animal Sciences, University of Illinois,
Urbana, IL 61801

P. J. Van Soest Professor of Animal Nutrition, Cornell University, Ithaca, NY 14853

Dale R. Waldo Research Animal Scientist, Ruminant Nutrition Laboratory, BARC-

East, Beltsville, MD 20705

Paul J. Weimer Research Microbiologist, USDA-ARS, U.S. Dairy Forage Research

Center, Madison, WI 53706

Eric Westerlund Professor, Department of Food Science, Swedish University of

Agricultural Sciences, S-75007 Uppsala, Sweden

Bryan A. White Associate Professor of Microbiology, Department of Animal

Sciences, University of Illinois, Urbana, IL 61801

J. R. Wilson Forage Physiologist, CSIRO Division of Tropical Crops and

Pastures, St. Lucia, Brisbane 4067, Australia
Conversion Factors for SI and non-SI Units

xxi
 .s:
Conversion Factors for SI and non-SI Units
To convert Column 1 To convert Column 2
into Column 2, into Column I,
multiply by Column 1 SI Unit Column 2 non-SI Unit multiply by

Length
0.621 kilometer, km (10 3 m) mile, mi 1.609
1.094 meter, m yard, yd 0.914
3.28 meter, m foot, ft 0.304
1.0 micrometer, ~m (10 -6 m) micron, ~ 1.0
3.94 x 10- 2 millimeter, mm (10 -3 m) 25.4 ('l
inch, in
0
10 nanometer, nm (10 -9 m) Angstrom, A 0.1 Z
<:
trl
i:Itl
Area rIJ
0
2.47 hectare, ha acre 0.405 -
Z
247 square kilometer, km 2 (10 3 m)2 acre 4.05 x 10- 3 "'I
0.386 square kilometer, km 2 (10 3 m)2 square mile, mi 2 2.590 >
('l
2.47 x 10- 4 square meter, m 2 acre 4.05 x 10 3 ...,
10.76 square meter, m 2 square foot, ft 2 9.29 x 10- 2 0
i:Itl
1.55 x 10- 3 square millimeter, mm 2 (10 -3 m)2 square inch, in 2 645 rIJ
"'I
0
i:Itl
Volume rIJ

9.73 x 10- 3 cubic meter, m 3 acre-inch 102.8 ->

35.3 cubic meter, m 3 cubic foot, ft 3 2.83 x 10- 2 Z
0
6.10 X 10 4 cubic meter, m 3 cubic inch, in 3 1.64 x 10- 5 Z
2.84 x 10- 2 liter, L (10 -3 m 3 ) bushel, bu 35.24 0
1.057 liter, L (10 -3 m 3 ) quart (liquid), qt 0.946 Z
3.53 x 10- 2 liter, L (10- 3 m 3 ) cubic foot, ft 3 28.3 ~
0.265 liter, L (10 -3 m 3 ) gallon 3.78 ~
liter, L (10 -3 m 3 ) 2.96 x 10- 2 Z
33.78 ounce (fluid), oz ...,
2.11 liter, L (10- 3 m 3 ) pint (fluid), pt 0.473 rIJ
-
 n
o
Mass 2
~
2.20 x 10- 3 gram, g (10 -3 kg) pound,lb 454 t"l
:::tI
3.52 X 10- 2 gram, g (10 -3 kg) ounce (avdp), oz 28.4 rIl
2.205 kilogram, kg pound,lb 0.454 o
0.01 kilogram, kg quintal (metric), q 100 2
"'l
1.10 x 10- 3 kilogram, kg ton (2000 lb), ton 907
1.102 megagram, Mg (tonne) ton (U.S.), ton 0.907 ..,>n
1.102 tonne, t ton (U.S.), ton 0.907 o
:::tI
rIl

Yield and Rate Cl

:::tI
0.893 kilogram per hectare, kg ha - 1 pound per acre, lb acre - 1 1.12 rIl
7.77 x 10- 2 kilogram per cubic meter, kg m- 3 pound per bushel, lb bu - 1 12.87 ..
1.49 x 10- 2 kilogram per hectare, kg ha -1 bushel per acre, 60 lb 67.19 >
2
1.59 x 10- 2 kilogram per hectare, kg ha - 1 bushel per acre, 56 lb 62.71 t:;;
1.86 x 10- 2 kilogram per hectare, kg ha - 1 bushel per acre, 48 lb 53.75 2
0.107 liter per hectare, L ha- 1 gallon per acre 9.35 o2
893 tonnes per hectare, t ha -1 pound per acre, lb acre- 1 1.12 x 10- 3
893 megagram per hectare, Mg ha -1 pound per acre, lb acre- 1 1.12 x 10- 3
r!l
~
0.446 megagram per hectare, Mg ha -1 ton (2000 lb) per acre, ton acre- 1 2.24 2
2.24 meter per second, m s -1 mile per hour 0.447 ::l
rIl

Specific Surface
10 square meter per kilogram, m 2 kg - 1 square centimeter per gram, cm 2 g-1 0.1
1000 square meter per kilogram, m 2 kg- 1 square millimeter per gram, mm 2 g - 1 0.001

Pressure
9.90 megapascal, MPa (10 6 Pal atmosphere 0.101
10 megapascal, MPa (10 6 Pal bar 0.1
1.00 megagram per cubic meter, Mg m- 3 gram per cubic centimeter, g cm- 3 1.00
2.09 x 10- 2 pascal, Pa pound per square foot, lb ft - 2 47.9
1.45 x 10- 4 pascal, Pa pound per square inch, lb in -2 6.90 x 10 3 g,
(continued on next page)
 ~
::;;.
Conversion Factors for SI and non-SI Units
To convert Column 1 To convert Column 2
into Column 2, into Column 1,
multiply by Column 1 SI Unit Column 2 non-SI Unit multiply by

Temperature
1.00 (K - 273) Kelvin, K Celsius, C 1.00 (OC + 273)
(9/5 C) + 32 Celsius, C Fahrenheit, of 5/9 (OF - 32)

Energy, Work, Quantity of Heat (')

9.52 x 10- 4 joule, J British thermal unit, Btu 1.05 X 10 3 o
:2
0.239 joule, J calorie, cal 4.19 <
10 7 joule, J erg 10- 7
0.735 joule, J foot-pound 1.36 ~
2.387 x 10- 5 joule per square meter, J m- 2 calorie per square centimeter (langley) 4.19 x 10 4 o
10- 5 :2
10 5 newton, N dyne
1.43 x 10- 3 watt per square meter, W m- 2 calorie per square centimeter 698 ~
minute (irradiance), cal cm -2 min- 1 Q
Transpiration and Photosynthesis ~
I'!j
3.60 x 10- 2 milligram per square meter second, gram per square decimeter hour, 27.8 o
mg m -2 s-1 gdm- 2 h- 1 rIl
5.56 x 10- 3 milligram (H 2 0) per s~uare meter micromole (H 20) per square centi- 180 ...==
second, mg m -2 s- meter second, ILmol cm - 2 S - 1 >
:2
10- 4 milligram per square meter second, milligram per square centimeter 10 4 o
mgm- 2 s- 1 second, mg cm -2 s-1 :2
35.97 milligram per square meter second, milligram ~r square decimeter hour, 2.78 X 10- 2 o
:2
mg m- 2 s-1 mgdm- h- 1
...ciJ
~
Plane Angle
57.3 radian, rad degrees (angle), 1.75 X 10- 2
..,:2...
rIl
 ("l
Electrical Conductivity, Electricity, and Magnetism o
z
10 siemen per meter, 8 m -1 millimho per centimeter, mmho cm -1 0.1 -<
10- 4 t"l
10 4 tesla, T gauss, G
~
Water Measurement
oZ
'"l
9.73 x 10- 3 cubic meter, m 3 acre-inches, acre-in 102.8 >
9.81 x 10- 3 cubic meter per hour, m 3 h -1 cubic feet per second, ft 3 s-1 101.9 Q
4.40 cubic meter per hour, m 3 h -1 U .8. gallons per minute, gal min -1 0.227 o
8.11 hectare-meters, ha-m acre-feet, acre-ft 0.123 ~
'"l
97.28 hectare-meters, ha-m acre-inches, acre-in 1.03 x 10- 2
8.1 x 10- 2 hectare-centimeters, ha-cm acre-feet. acre-ft 12.33 i
[IJ

Concentrations >
-
Z
o
1 centimole per kilogram, cmol kg- 1 milliequivalents per 100 grams, meq 1 Z
(ion exchange capacity) 100 g-1
0.1 gram per kilogram, g kg- 1 percent, % 10 ~
1 milligram per kilogram. mg kg - 1 parts per million, ppm 1 ~
~
Z
Radioactivity ~
2.7 x 10- 11 becquerel, Bq curie, Ci 3.7 X 1010
2.7x10- 2 becquerel per kilogram. Bq kg -1 picocurie per gram. pCi g-1 37
100 gray. Gy (absorbed dose) rad, rd 0.01
100 sievert, 8v (equivalent dose) rem (roentgen equivalent man) 0.01

Plant Nutrient Conversion

Elemental Oxide
2.29 P P 20 5 0.437
1.20 K K 20 0.830
1.39 Ca CaO 0.715 ~
~
1.66 Mg MgO 0.602 <