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 Second Edition

BARLEY
CHEMISTRY AND TECHNOLOGY

Edited by

Peter R. Shewry
Rothamsted Research
Harpenden, Hertfordshire
United Kingdom

Centre for Food Security

University of Reading,
Reading, United Kingdom

Steven E. Ullrich
Department of Crop and Soil Sciences
Washington State University
Pullman, Washington, U.S.A.
Front cover (left to right in each row): First row, combine harvester in barley
field, courtesy James Hutton Institute; ear of a two-row barley, courtesy
Sabine Gubatz; field of barley during grain-filling stage, courtesy James
Hutton Institute. Second row, barley grain and glass of whisky, courtesy
James Hutton Institute; cattle feeding image, Pavel L Photo and Video/
Shutterstock.com; field of mature barley ready for harvest, courtesy
James Hutton Institute. Third row, barley soup image, Aleksandra Duda/
Shutterstock.com; barley seed image, kzww/Shutterstock.com; beer glass
image, aragami12345s/Shutterstock.com

Reference in this publication to a trademark, proprietary product,

or company name by personnel of the U.S. Department of Agriculture
or anyone else is intended for explicit description only and does not
imply approval or recommendation to the exclusion of others
that may be suitable.

Library of Congress Control Number: 2013956764

International Standard Book Number: 978-1-891127-79-3

©1993, 2014 by AACC International, Inc.

Published 1993. Second Edition 2014

All rights reserved.

No part of this book may be reproduced in any form, including photocopy,
microfilm, information storage and retrieval system, computer database or
software, or by any other means, including electronic or mechanical,
without written permission from the publisher.

Printed in the United States of America on acid-free paper

AACC International, Inc.

3340 Pilot Knob Road
St. Paul, Minnesota 55121, U.S.A.

Byung-Kee Baik, Department of Crop and Soil Sciences, Wash-
ington State University, Pullman, WA. Current address: SWQL,
USDA-ARS, 1680 Madison Ave, Wooster, OH 44691, U.S.A.
Geoffrey B. Fincher, ARC Centre of Excellence in Plant Cell
Walls, School of Agriculture, Food and Wine, University of
Adelaide, Waite Campus, Glen Osmond, SA 5064, Australia
Christine Finnie, Department of Systems Biology, Technical
University of Denmark, Building 224, Søltofts Plads, DK-
2800, Kongens Lyngby, Denmark
B. R. Gibson, VTT Technical Research Centre of Finland, P.O.
Box 1000, FI-02044 VTT, Finland
Mario Gils, Leibniz Institute of Plant Genetics and Crop Plant
Research, Corrensstr. 3, 06466 Stadt Seeland, OT Gater­
sleben, Germany
Andreas Graner, Leibniz Institute of Plant Genetics and Crop
Plant Research, Corrensstr. 3, 06466 Stadt Seeland, OT
Gatersleben, Germany
Sabine Gubatz, Rothamsted Research, Department of Plant Sci-
ence, West Common, Harpenden, Hertfordshire, AL5 2JQ,
U.K.
Kevin B. Hicks, Sustainable Biofuels and CoProducts Research
Unit, Eastern Regional Research Center, ARS, USDA, 600
E. Mermaid Lane, Wyndmoor, PA 19038, U.S.A.
Marta S. Izydorczyk, Canadian Grain Commission, Grain Re-
search Laboratory, 1404-303 Main Street, Winnipeg, Mani-
toba, Canada R3C 3G8
Justin Montanti, Sustainable Biofuels and CoProducts Research
Unit, Eastern Regional Research Center, ARS, USDA, 600
E. Mermaid Lane, Wyndmoor PA 19038, U.S.A.
Matthew Morell, CSIRO Food Futures National Research Flag-
ship, CSIRO Plant Industry, GPO Box 1600, Canberra, ACT
2601, Australia
Nhuan P. Nghiem, Sustainable Biofuels and CoProducts Re-
search Unit, Eastern Regional Research Center, ARS, USDA,
600 E. Mermaid Lane, Wyndmoor, PA 19038, U.S.A.
Contributors
Odd-Arne Olsen, Department of Plant and Environmental Sci-
ences, Norwegian University of Life Sciences, PO Box 5003,
N-1432 Ås, Norway and Faculty of Education and Natural
Sciences, Hedmark University College, N-2306 Hamar,
Norway
A. Ritala, VTT Technical Research Centre of Finland, PO Box
1000, FI-02044 VTT, Finland
Peter R. Shewry, Rothamsted Research, Department of Plant
Science, West Common, Harpenden, Hertfordshire, AL5
2JQ, U.K. and Centre for Food Security, University of Read-
ing, Agriculture Building, Earley Gate, Whiteknights Road,
Reading, RG6 6AR, U.K.
Nese Sreenivasulu, Martin Luther University Halle-Wittenberg,
Interdisciplinary Center for Crop Plant Research, Halle-
Wittenberg, Germany. Current address: International Rice
Research Institute, Los Baños, DAPO Box 7777, Metro
Manila, Philippines
Birte Svensson, Department of Systems Biology, Technical Uni-
versity of Denmark, Søltofts Plads, Building 224, DK-2800,
Kongens Lyngby, Denmark
J. S. Swanston, The James Hutton Institute, Errol Road, Inver-
gowrie, Dundee, DD2 5DA, U.K.
David Topping, CSIRO Food Futures National Research Flag-
ship, CSIRO Animal, Food and Health Sciences, PO Box
10041, Adelaide BC, SA 5000, Australia
Kay Trafford, National Institute of Agricultural Botany, Hunt­
ingdon Road, Cambridge, CB3 0LE, U.K.
Steven E. Ullrich, Department of Crop and Soil Sciences, Wash-
ington State University, Johnson Hall 131, Pullman, WA
99164-6420, U.S.A.
A. Wilhelmson, VTT Technical Research Centre of Finland,
PO Box 1000, FI-02044 VTT, Finland
Winfriede Weschke, Leibniz Institute of Plant Genetics and
Crop Plant Research, Correnstr. 3, 06466, Stadt Seeland, OT
Gatersleben, Germany
a iii
This page intentionally left blank
 Preface to the Second Edition
The AACC International monographs dedicated to the chem-
istry and technology of cereals have become the major reference
works for cereal chemists, food scientists, and crop scientists
alike since the first volume (on wheat, now in its fourth edition)
was launched in 1964.
The volume on barley is one of the youngest, being first pub-
lished in 1993 and edited by Sandy MacGregor and Ron Bhatty.
The massive increase in our knowledge over the last two decades
has therefore necessitated a complete revision, with only three of
the original authors remaining, two of whom provide updates of
their previous chapters.
However, much of the information in the first edition re-
mains valid and relevant, and the emphasis has therefore been
on complementing and updating rather than repeating. Most
readers will continue to refer to their copies of the first edition
for more historical studies. For example, the chapter on seed
proteins (Chapter 6) complements the detailed coverage of hor-
deins in the first edition by focusing on more recent studies of
other grain proteins. Similarly, the inclusion of a new chapter
(Chapter 5) on grain arabinoxylans recognizes that this group of
cell-wall polysaccharides has received relatively little attention
in barley until recent times, compared with the more-abundant
β-glucans. Although knowledge on most topics has advanced
significantly over the past 20 years, there is one notable excep-
tion. This is barley lipids, which have been neglected and are
therefore considered with other “minor components” in Chapter
7. The reader is referred to the comprehensive account of the ear-
lier studies of barley lipids, and their limitations, provided by
Bill Morrison in the first edition.
Despite the inevitability of some overlap, we decided to in-
clude two chapters on grain development, focusing on the control
and pattern of endosperm development (Chapter 2) and on the
broader aspects, including detailed histological studies on control
by nutrition and plant hormones (Chapter 3). These together pro-
vide the most detailed account that has been published to date.
Finally, we would draw the reader’s attention to Chapters 9
and 11, which discuss health benefits and nonfood uses of barley,
respectively. Both of these topics have been widely discussed over
the last decades, and these chapters provide topical and critical
accounts of our current knowledge and further prospects.
We have found it a pleasure to work with a distinguished
group of authors and particularly thank those who contributed
chapters at short notice. We also thank the editorial staff at the
AACC International for their support and their trademark high
quality of production, while Peter Shewry would particularly
like to thank Helen Jenkins at Rothamsted Research for her as-
sistance. The volume has been a labor of love for us, and we hope
that you enjoy reading and using it.
Peter Shewry
Steve Ullrich
av
This page intentionally left blank
 Preface to the First Edition
With the publication of the present volume, the American
Association of Cereal Chemists has completed monographs on
six of the eight major cereals in the family Gramineae, some in
second (rice) and third (wheat) editions. Barley, although the
fourth major cereal in the world (after wheat, rice, and corn), has
been late in making it to the list of AACC published titles. Part
of the reason for this delay was the perception of barley as, first,
a major raw material for the production of alcoholic beverages
(hence of interest largely to the malting and brewing industries)
and, second, a livestock feed. Animal feeds and malt consume
80–90% of the barley production in many Western countries.
At the present time, very little barley is used in human foods
except in the West Asia-North Africa region and until recently
in the Pacific Rim countries, particularly Korea and Japan. Yet,
historically, barley preceded wheat as a food grain in ancient
Egypt and Nubia, and it has been consumed since then in many
cultures. Only at the beginning of this century was barley re-
placed as a food grain by wheat because of the superior bak-
ing quality of wheat flour. Barley, grown in regions of the world
from sub-Artic to temperate, is a versatile cereal. It may be
hulled, thin-hulled, or hull-less, two-rowed or six-rowed, have
normal or high lysine levels, low or high amylose starch levels,
and low or high levels of β-glucan, a major component of solu-
ble dietary fiber implicated in hypocholesterolemia. Moreover,
it does not contain any known antinutritional factors. In view
of these properties, barley remains an underutilized cereal
in human foods and in industrial applications, except for the
malting and brewing industries.
This monograph contains 10 chapters, each written by in-
ternationally acclaimed experts in their respective fields. The
editors suggested only broad guidelines to the authors, who de-
veloped their own chapters. The intention of the editors was to
produce a volume that was broad in scope yet covered each topic
in depth. In our opinion, these objectives have been largely met.
We are indebted to the authors for agreeing in the first place
to write the chapters and for by and large meeting the various
deadlines imposed on them, in spite of their busy schedules; to
reviewers of chapters; and finally to the American Association of
Cereal Chemists for publishing the monograph.
A.W. MacGregor
R. S. Bhatty
a vii
This page intentionally left blank
 Contents

1. The Barley Crop: Origin and Taxonomy, Production, and End Uses. STEVEN E. ULLRICH ♦ 1
Origin and Taxonomy of Barley • 1
Adaptation and Production of Barley • 3
Uses of Barley • 5
Feed Uses | Malting, Brewing, and Distilling Uses | Food Uses | Other Uses of Barley
2. Barley Grain: Development and Structure. SABINE GUBATZ and WINFRIEDE WESCHKE ♦ 11
From Inflorescence Initiation to Flowering • 11
Setting Up Reproductive Structures | Primordia Supply and Survival | Development of Reproductive Structures Until Anthesis
| Anthesis and Fertilization
The Young Developing Grain • 19
Maternal Influences on Early Grain Development | Endosperm: Free Nuclear Phase, Cellularization, and Differentiation
| Early Development of the Embryo | Phytohormones and Early Grain Development
The Grain During Storage-Product Accumulation • 31
Homonal Influences on Endosperm Development | Nutrient Uptake Capacity and Sink Strength
| Endopolyploidization and Storage-Product Accumulation | Aleurone Development | Embryo Growth
The Maturing Grain • 34
Programmed Cell Death of the Endosperm | Desiccation Tolerance of the Embryo and Aleurone | Dormancy
| Seed Coat and Protective Hull of the Grain
3. Genetic and Molecular Aspects of Barley Grain Development. ODD-ARNE OLSEN and
WINFRIEDE WESCHKE ♦ 55
Endosperm • 55
Mutants Affected During Endosperm Development | Endosperm Cellularization | Endosperm Cell Fate Specification
| Maternal Control of Endosperm Development
Embryo • 63
Fertilization and Early Stages of Embryo Development | Embryo Differentiation | Mutants That Affect Embryo Development
| Embryo Development as Influenced by Surrounding Tissues | Coordination of Growth and Development of Embryo and Endosperm
Conclusions • 67
4. Barley Grain Carbohydrates: Starch and Cell Walls. KAY TRAFFORD and GEOFFREY B. FINCHER ♦ 71
Barley Endosperm Starch • 72
Starch Structure and Composition | Starch Granule Morphology | Starch Synthesis During Grain Development
| Subcellular Compartmentation of the Starch Biosynthetic Pathway | Evolution of the Genes Encoding the Cytosolic ADPglucose
Pathway | Genetic Variation in Barley Endosperm Starch
Cell Wall Polysaccharides in Barley Grain • 80
Wall Composition | (1→3,1→4)-β-Glucan Structure and Properties | Synthesis of (1→3,1→4)-β-Glucan in the Developing Barley
Grain | Genetic Variation in (1→3,1→4)-β-Glucan Levels in Barley Grain | Arabinoxylan Structure and Properties | Synthesis and
Remodeling of Arabinoxylans in the Developing Barley Grain | Genetic Variation in Arabinoxylan Levels in Barley Grain
Coordination of Starch and (1→3,1→4)-β-Glucan Synthesis • 88
Concluding Remarks • 89

a ix
x a Contents

5. Barley Arabinoxylans: Molecular, Physicochemical, and Functional Properties.

MARTA S. IZYDORCZYK ♦ 97
Occurrence and Location • 97
Molecular Structures of Arabinoxylans in Barley Grain • 100
General Molecular Features | Structural Diversity of Arabinoxylans
Extraction of Arabinoxylans from Barley and Barley Co-Products • 107
Physicochemical Properties of Arabinoxylans • 109
Molecular Weight and Solution Properties | Gelation
Biosynthesis of Arabinoxylans • 111
Arabinoxylans in Malting and Brewing • 112
Arabinoxylans in Human Nutrition and Health • 114
Arabinoxylans as Dietary Fiber | Physiological Effects in the Gastrointestinal Tract
Animal Nutrition • 116
6. Barley Grain Proteins. CHRISTINE FINNIE and BIRTE SVENSSON ♦ 123
Technical Developments Advancing Grain Protein Research • 123
Protein Mass Spectrometry | Proteome Analysis | Dissection of Seed Tissues and Laser-Capture Microdissection
| MALDI Imaging | Surface Plasmon Resonance and Isothermal Titration Calorimetry | Recombinant Protein Production
and Structure Determination | Molecular Farming: Production of Recombinant Proteins in Barley Grains
Classification of Barley Grain Proteins • 129
Storage Proteins | Amino Acid Composition of Barley Grains and Relation to Protein Quality
Temporal and Spatial Analysis of Proteins in Barley Grains • 132
Changes in Protein Profiles During Grain Development, Germination, and Malting | Tissue-Specific Analysis of Protein Profiles
| Proteins of the Barley Aleurone Layer | Proteins in Subcellular Compartments of the Barley Grain: Amyloplast and Starch Granule
| Proteins in Subcellular Compartments of the Barley Grain: Plasma Membrane Proteins | Membrane Proteins: Sucrose Transporters
| Membrane Proteins: Peptide Transporter HvPTR1 | Genetic Variations in Barley Grain Protein Profiles
| Environmental Effects on Grain Proteins
Functional and Structural Analysis of Barley Proteins • 139
The Barley Grain Thioredoxin System | The Ascorbate-Glutathione Cycle in Barley Grains
Properties of Selected Barley Seed Proteins • 142
Protein Z (Barley Serpin) | Phytase
Structural Characterization of Barley Proteins • 143
Chitinases | β-Amylase | Hordothionin | Protease and Carbohydrase Inhibitors | Lipid Transfer Proteins
| Thaumatin-Like Proteins | Barwin | Peroxidases | Oxalate Oxidase | α-Amylase | Limit Dextrinase | α-Glucosidase
| Exo-β-Glucanases, Endo-β-Glucanases, and β-Glucosidases | Cysteine-Endoprotease | Aspartic Proteinase | Barley Lectin
Perspectives • 157
7. Minor Components of the Barley Grain: Minerals, Lipids, Terpenoids, Phenolics, and Vitamins.
PETER R. SHEWRY ♦ 169
Minerals • 169
Lipids • 171
Lipids in Whole Grain | Lipids in Grain Tissues | Starch Lipids
Terpenoids • 175
Tocols | Sterols | Carotenoids
Phenolic Compounds • 178
Phenolic Acids | Flavonoids | Alkylresorcinols | Other Phenolic Compounds
B Vitamins • 188
 Contents a xi
8. Malting, Brewing, and Distilling. J. S. SWANSTON, A. WILHELMSON, A. RITALA, and B. R. GIBSON ♦ 193
Malting • 194
Commercial Mlat Production | Phases of Malting
Brewing • 198
Outline of Wort Production | Mashing and Wort Separation | Wort Boiling
Brewery Fermentation and Beer Character • 203
Influence of Barley-Derived Components
Distilling • 206
History and Background | Types of Whiskeys
Other Uses of Malt • 209
Food Use
Barley and Malt Testing • 209
Germination Tests | Malt Tests | Breeding and Screening Tests
Future Directions • 212
Technical Developments Within Industries | Barley Breeding Targets and Approaches
9. Barley Foods and Public Health. DAVID TOPPING and MATTHEW MORELL ♦ 223
Diet, Whole Grains, and Disease Risk • 224
Barley Constituents and Human Health Outcomes • 225
The Evolving Relationship Between Dietary Fiber and Human Health • 226
Dietary Fiber | Dietary Fiber, Resistant Starch, and the Australian and African Paradoxes
Barley Components and Health Outcomes • 226
Barley Fiber Components and Health Outcomes • 227
Total and Insoluble Dietary Fiber and Laxation | Barley β-Glucan and Plasma Cholesterol
Barley Foods, Health, and Value Addition • 228
Rihane and Large-Bowel Disease | High-Glucan Barley | High-Amylose Barley
Conclusions and Future Directions • 228
10. Processing of Barley Grain for Food and Feed. BYUNG-KEE BAIK ♦ 233
Processing of Barley Grain for Food Preparation • 234
Whole-Grain Processing | Malting Barley Grain for Food Use | Size-Reduction Processing of Barley Grain
| Changes in Composition of Barley Grain by Processing | Barley Food Preparation
Processing of Barley Grain for Feed • 252
Characteristics of Barley Grain for Feed | Processing of Barley Grain for Feed Preparation
Summary • 259
11. Use of Barley Grain and Straw for Biofuels and Other Industrial Uses. KEVIN B. HICKS, JUSTIN MONTANTI,
and NHUAN P. NGHIEM ♦ 269
Use of Barley Grain (Starch) for Biofuels • 269
Use of Hull-Less Barley Varieties | Very-High-Gravity Fermentations and Use of β-Glucanases to Reduce Mash Viscosity
| Conversion of β-Glucans in Barley Grain to Fuel Ethanol and the EDGE Winter Barley Ethanol Process
| Low-Temperature Granular-Starch-Hydrolyzing Enzyme Technology for Barley Ethanol | Dry-Fractionation of Hulled Barley
Before Fermentation | Mycotoxin Mitigations | Improvements in DDGS Quality | Additional Barley Ethanol Co-Products
| Continued Improvement in Barley Varieties for Fuel Ethanol Production | Life Cycle Assessments of Greenhouse Gas Emissions
from Winter Barley Ethanol, a Potential Advanced Biofuel
Use of Barley Straw and Related Lignocellulosic Barley Biomass for Biofuels • 277
Cellulosic Ethanol | Cellulosic Butanol | Biomethane and Hydrogen
Pyrolysis-Derived Biooils, Biochar, and Syngas • 280
Simultaneous Conversion of Barley Starch and Biomass for Biofuels • 280
Uses of Barley Grain and Straw Components to Produce Biobased Nonfood Products • 280
Starch | β-Glucan | Protein | Antioxidants | Arabinoxylans
Direct Uses of Barley Lignocellulosic Components • 283
Bioremediation | Ground Cover | Fuel Pellets
xii a Contents

12. Molecular Breeding for Malting Quality. MARIO GILS, ANDREAS GRANER, and NESE SREENIVASULU ♦ 293
Overview • 293
Understanding Barley Seed Development: A Prerequisite for Improving Malting Quality • 294
Functional Relevance of Key Genes of Starch Metabolism Studied Through Mutant Analysis • 295
Malting Quality: A Quantitative Genetics Approach • 296
From QTLs to Candidate Genes • 296
Differentially Expressed Genes • 297
Mutation Breeding • 297
Doubled-Haploid Technology • 299
Hybrid Cultivar Breeding • 300
Transformation • 301
GMO Approaches to Improving Malting Quality • 301
Endosperm Protein Expression: Alteration of the Cell Wall or Storage Protein Manipulation
| Expression of Thermostable (1→3,1→4) β-Glucanase
Conclusions and Perspectives on the Future • 303
Index ♦ 311
 Second Edition

BARLEY
CHEMISTRY AND TECHNOLOGY
This page intentionally left blank
 CHAPTER 1

The Barley Crop: Origin and Taxonomy,

Production, and End Uses
Steven E. Ullrich
Deptment of Crop and Soil Sciences
Washington State University
Pullman, Washington, U.S.A.

Origin and Taxonomy of Barley Other domestication traits found in barley samples from parts
The origins of almost all plant species, including the major of the world away from the regions of origin include 4) a reduced
crop species, and the origins of domestication of crop species are vernalization requirement and 5) photoperiod insensitivity that
ancient, far predating the human understanding of such things. facilitates adaptation to different environments and climates
As such, the determination of the origins of crop species relies (Salamini et al 2002).
on evidence from the archeological record and on scientific in- Barley belongs to the globally and economically very im-
vestigation using the most up-­to-­date technologies available at portant plant group, the Triticeae, which is a tribe in the grass
the time, for example, from plant anatomy and morphology, bio- family Poaceae. This tribe is characterized as having spike in-
chemistry, genetics, and agriculture. Therefore, our knowledge florescences, a base chromosome number x = 7, and large ge-
and even theories of crop origin and taxonomy are frequently nomes. In addition to barley, the Triticeae includes the major
incomplete and subject to periodic change. This is definitely temperate small-­grain cereals: wheat (Triticum spp.); rye (Secale
the case for barley (Hordeum vulgare L.), one of the most an- cereale); and triticale (Triticosecale), a human-­developed wheat-­
cient of crop species, which is currently considered to be among rye hybrid crop. In addition, the Triticeae includes forage grasses
the world’s first crop domesticates. Whereas barley domestica- such as the wheatgrasses (Agropyron spp.) and Russian wildrye
tion was once thought to have occurred as a single event in the
Fertile Crescent of the Middle East at least 10,000 years ago, it
is now thought that multiple domestications occurred within
the Fertile Crescent (Zeder 2008), with further domestications
about 1,500–3,000 km further east in central Asia (Azhaguvel
and Komatsuda 2007, Morrell and Clegg 2007).
One approach to determining whether a plant species has
been domesticated is the presence, in archeological samples, of
morphological traits resulting from genetic changes (natural
or caused by humans) that favor agricultural production (von
Bothmer et al 2003b). In the case of barley, such domestication
traits include 1) the presence of a nonbrittle rachis that prevents
the spike from shattering and the grain from scattering upon
maturity (Fig. 1.1); 2) the change from a two-­rowed spike in the
wild progenitor barley (H. vulgare subsp. spontaneum) (Fig. 1.2)
Fig. 1.1. Illustrations of a nonbrittle, nonshattering spike (A) and
to the six-­rowed spike in H. vulgare subsp. vulgare (Fig. 1.3); and brittle, shattering spikes (B) of wild barley (Hordeum vulgare subsp.
3) naked or hull-­less kernels (Fig. 1.4), which also do not occur spontaneum), which probably represent one of the first selections by
in wild barley. Hull-­less kernels facilitate the use of barley for humans that began the domestication process of barley. (Courtesy of
human food because of the reduced insoluble fiber in the hull. Brian Steffenson, University of Minnesota; used by permission)

a1
2 a Barley: Chemistry and Technology, 2nd ed.
(Psathyrostachys juncacea), as well as noxious weeds, including
a close relative of wheat, jointed goatgrass (Aegilops sp.), as well
as quackgrass (Elymus repens), and foxtail barley (H. jubatum).
Relatively recent advances in technological tools have in-
creased our knowledge about the evolutionary age, migra-
tion patterns, and differentiation of new species in the genus
Hordeum, and today the genus is much better understood than
ever before. It is estimated that Hordeum is quite ancient, having
separated from wheat species about 13 million years ago. The re-
lationships among species within a genus can be relatively close
or quite distant. In the case of cultivated barley, most species
are rather distant but still related. Biological understanding of
the members of the genus Hordeum is very useful in identifying
gene sources for improvement of cultivated barley (H. vulgare
Fig. 1.2. Two-­rowed barley spikes of Hordeum vulgare subsp. vul-
gare, which were inherited directly from wild barley (H. vulgare
subsp. spontaneum). (Courtesy of Patrick M. Hayes, Oregon State
University; used by permission)
Fig. 1.3. Six-­rowed barley spikes of Hordeum vulgare subsp. vulgare,
selected from the two-­rowed type sometime during the domestica-
tion process. (Courtesy of Patrick M. Hayes, Oregon State University;
used by permission)
subsp. vulgare). The primary gene pool for cultivated barley is the
wild subspecies progenitor H. vulgare subsp. spontaneum (Figs.
1.5 and 1.6), which is completely cross-­compatible and is increas-
ingly being explored as a source of genes for traits improving
cultivated barley, such as pest resistance, nutritional quality, and
even malting quality (Fetch et al 2003, Matus et al 2003).
Probably the second-­best Hordeum source of genes for cul-
tivated barley is the secondary gene pool species H. bulbosum.
Initial interest in this species was for doubled haploid produc-
tion in cultivated barley and wheat, as the H. bulbosum chro-
mosomes are typically eliminated from hybrids (Kasha and
Kao 1970, Forster et al 2007). Research has been more recently
directed at making viable, stable hybrids between H. vulgare
and H. bulbosum in order to transfer desirable genes, especially
disease-­resistance genes (Pickering and Johnston 2005). Al­
though hybridization is extremely difficult, there is interest in
tertiary gene pool species, such as H. marinum (sea barley) for
salt tolerance (Malik et al 2009).
Fig. 1.4. Hulled (A) and hull-­less (B) barley kernels
of Hordeum vulgare subsp. vulgare. Hull-­less kernels
were selected sometime during the domestication
process. (Courtesy of Byung-­Kee Baik, USDA-­ARS,
Wooster, OH; used by permission)

Hordeum species have migrated from the centers of origin
and evolved to other areas around the world, including the
Americas. In addition to the species mentioned above, other pe-
rennials and annuals are considered to be valuable forage spe-
cies in natural pastures in central Asia and South America (e.g.,
H. murinum). H. chilense, from South America, has been used to
create a new hybrid species, tritordeum, by crossing with wheat.
The contributions from H. chilense include improved baking
Fig. 1.5. Spikes of wild barley (Hor-
deum vulgare subsp. spontaneum).
(Courtesy of Brian Steffenson,
University of Minnesota; used by
permission)
Fig. 1.6. Field of wild barley (Hordeum vulgare subsp. spontaneum).
(Courtesy of Prof. Brian Steffenson, University of Minnesota; used by
permission)
Barley Crop a3
quality and resistance to biotic and abiotic stresses (Martin et al
1999). Other species (e.g., H. jubatum) are weedy or have other
negative characteristics, such as acting as hosts for pests.
This chapter provides only a brief introduction to the origin
and taxonomy of barley. Excellent and comprehensive coverage
of these subjects can be found in von Bothmer et al (2003a) and
von Bothmer and Komatsuda (2011).
Adaptation and Production of Barley
Cultivated barley is adapted to and grown over a wider range
of environmental conditions than any other cereal. It is one of
the most widely adapted crops in the world and is grown further
toward the poles, into deserts, and at higher elevations than any
other crop. The author has observed barley growing for grain at
higher than 65° N. latitude in Alaska and the Nordic countries
of Finland and Norway; toward the margins of the Sahara desert
in Algeria, beyond the range of drought-­tolerant durum wheat
(Triticum durum); and on the Altiplano of Bolivia at 420 m, where
the only other cereal planted is oats (Avena sativa) for fodder.
Barley is a cool-­season crop cultivated in spring and summer at
temperate latitudes and in the tropics at high altitudes. It is culti-
vated in the fall and winter at low and medium altitudes in trop-
ical and subtropical latitudes. Barley is relatively cold-­tolerant
and is considered the most tolerant to drought, alkali, and salt
among small-­grain cereal species, but it is not well adapted to
wet and acidic soil conditions. Relatively rapid emergence, early
maturity, and low water use are the major factors that confer
adaptation to drought and high-­temperature conditions. Spring
types are quite cold-­tolerant, but winter types are less winter-­
hardy than wheat, rye, and triticale. Whereas barley can be and
is grown in some of the world’s agricultural margins, it performs
best under well-­drained fertile loam soils with moderate precipi-
tation (400–800 mm) and temperatures (15–30°C).
The role of barley in cropping systems varies throughout the
world depending on production environments (e.g., climate and
geography) and economic environments (e.g., market supply and
demand and the uses for specific types of barley). The adaptation
of barley allows it a prominent place in crop rotations in the more
marginal production environments of extreme latitude or alti-
tude and low precipitation, but its prominence tends to decrease
as production environments become more favorable in terms of
temperature regimes and water availability. A good example can
be seen in the Pacific Northwest of the United States. Barley is
important in the high semidesert of southern Idaho (which has a
cool and short growing season and crops are mostly irrigated) in
rotation with wheat and potato (Solanum tuberosum). The eco-
nomic environment is also favorable for malting and brewing
cultivars in that there are three malting plants in the area. On
the other hand, almost no barley is grown in the low-­elevation,
semidesert, irrigated Columbia Basin of eastern Washington
because the long, warm growing season favors a plethora (more
than 50) of higher-­value crops such as maize (Zea mays), alfalfa
(Medicago sativa), hops (Humulus lupulus), vegetable crops,
grapes (Vitis spp.), tree fruits, etc. The same trends can be seen
in dryland production environments. Generally, the more crops
that are adapted to and can be grown in an area, the lower the
prominence of barley in crop rotations. Economics tends to be a
4 a Barley: Chemistry and Technology, 2nd ed.

major factor. Barley usually commands a lower price than other 1.2). Barley production peaked in 1990 at 178 million tonnes,
cereals, legumes, and oilseeds because it is primarily used for then steadily declined to 134 million tonnes in 2011, the final
feed. On the other hand, demand for malting barley can equal- year of FAO data available at this writing. Barley production
ize the profitability since malting barley cultivars usually com- ranked fourth in the world from 1961 until 1998, when soy-
mand a significant premium over those of feed. A good example bean production eclipsed that of barley (160 vs. 138 million
is in central and northern Europe, where the temperate and tonnes). While barley production rose and fell over this 50-­
relatively wet growing conditions tend to be quite favorable for year period, the other four crops steadily rose in production,
several cool-­season crops (such as wheat, oilseeds, and Brassica most notably maize and soybean. The production of wheat was
vegetable and forage crops). However, the demand for malting highest among these five crops until 2000, when rice produc-
barley in this area is high, which keeps barley production high. tion eclipsed wheat, and maize eclipsed them both. The rank
France and Germany tend to be the biggest producers of barley of the top five crops has not changed so far in the twenty-­first
in Europe, and they also rank high globally, as described below. century (Table 1.1). The percentage of 2011 production over
Barley also ranks high in production in cool Canada and semi- that of 1961 is quite remarkable for the top five crops: soybean,
arid Australia but less so in the United States, which has more 967%; maize, 431%; rice, 335%; wheat, 317%; and barley, 186%.
favorable growing environments. Maize and soybeans (Glycine Soybean production largely accelerated as a result of increased
max) dominate in the United States, and the highest-­ranking oil and protein demand and maize production because of the
barley-­producing states all border Canada, from the Pacific phenomenal performance of hybrids. Both of these crops have
Ocean through the Great Plains to the Great Lakes. received large increases in breeding and genetic research fund-
Barley is grown under a wide range of growing conditions in ing from the private and public sectors. The improvement and
terms of climate, tillage, fertility, moisture, and crop rotation. proliferation of rice hybrids have contributed to rice produc-
Compared with wheat, which is relatively closely related, bar- tion increases. The rise in population in South and East Asia,
ley generally requires less water and plant nutrients, primarily resulting in a rise in demand, is also a factor in rice production
because it has a shorter growing season. Earlier maturity also increases. Increases in soybean acreage in South America, es-
means that it can escape drought, disease, and insect stresses pecially in Brazil and Argentina, and in the U.S. Midwest have
better than wheat under many conditions. A recent summary of been drivers. Increased demand for biofuels has affected maize
barley agronomic practices can be found in Ullrich (2011), which and soybean acreages. Note that these are somewhat simplified
includes a chapter covering the major barley-­producing areas of explanations of considerably complex situations.
the world. Barley production is widely distributed around the world.
Barley is a major world crop, ranking fourth among cereals The FAO barley database includes production estimates from 98
and fifth overall in millions of tonnes produced per year. The countries. Table 1.3 shows five-­year (2007–2011) average barley
global production of barley, averaged over the first 11 years of production according to continents and regions within conti-
the twenty-­first century, was estimated to average 141 million nents. The greatest production by far is in Europe, including the
tonnes per year, behind maize (715), rice (Oryza sativa) (628), Russian Federation, at 88 million tonnes or 64% of the world
wheat (618), and soybean (207), as shown in Table 1.1. Barley is production, of which approximately two thirds is from the
also ranked fifth based on estimated land area harvested per European Union (EU) of 27 countries. Asia ranks second at 20
year in millions of hectares (ha), after wheat (217), rice (153), million tonnes or 14% of the world production, with about half
maize (148), soybean (89), and barley (55). Production estimates from western Asia; followed by North America (mainly Canada
are based on data from the Food and Agriculture Organization and the United States) at 14 million tonnes (11%); Australia and
(FAO) of the United Nations (FAO 2012b). The trends of the top New Zealand at 8 million tonnes (5%); Africa at 6 million tonnes
five produced crops over the past 50 years from 1961 through (4%), with two thirds of that from North Africa; and finally
2010 indicate some major dynamics and shifts in rank (Table South America at 3 million tonnes (2%). The Russian Federation
heads the list of the top 20 barley-­producing countries in the
world, based on eight-­year averages (2003–2010) of total produc-
tion amount (16.6 million tonnes, or approximately 12% of the
TABLE 1.1
Global Production Average Estimatesa of the Top Eight Crops
142 million tonnes world total) and area of production (8.4%, or
for 11 Years (2000–2010)b
Production Area Harvested Grain/Crop Yield TABLE 1.2
Crop (million t) (million ha) (t/ha) Global Trends for Production (million t) of the Top Five Crops
Maize 715 148 4.8 in the World over 50 Years (1961–2010) a
Rice (paddy) 628 153 4.2
Year Wheat Rice Maize Barley Soybean
Wheat 618 217 2.8
Soybean 207 89 2.3 1961 2,222 216 205 72 27
Barley 141 55 2.6 1971 348 318 314 132 46
Sugarcane (dw) 99 22 4.7 1981 450 410 447 150 89
Potato (dw) 61 19 3.2 1991 547 519 494 170 103
Sorghum 59 43 1.4 2001 590 600 616 144 178
a Expressed 2011 704 723 883 134 261
“as is,” except dry weight (dw) for sugarcane and potato.
b Source: FAO (2012b). a Source: FAO (2013).
 Barley Crop a5
approximately 15% of the 55 million hectares world total) (Table Uses of Barley
1.4). The other top five countries include Germany (11.7 million
tonnes), Canada (11.1), France (10.7), and Ukraine and Spain Barley has had many and diverse uses, which have evolved,
(9.3 each). Sweden, at number 20, produced 1.6 million tonnes. waxed, and waned over the millennia from hunting and gath-
Nine of the top 20 countries are EU members, with the highest ering times, through the development of agriculture, and up to
national average yields being from the northern EU countries today. Carl Nilsson Linnæus must have been thinking of barley
France, Germany, United Kingdom, and Denmark at 7.1, 6.7, 6.7, use when he gave the scientific or Latin name to barley, Hordeum
and 5.8 t/ha, respectively. vulgare. Hordeum derives from hordearii, the Latin name for
gladiators, and translates to “barley men.” This is because bar-
ley was a primary component of the training diet of gladia-
TABLE 1.3 tors, which gave them strength, stamina, and subcutaneous fat
World Distribution of Barley Production Based on Estimates, (Percival 1921, Curry 2008). A layer of fat protected the gladiators
Five-Year Averages (2007–2011)a from relatively superficial wounds, but the bleeding from these
Number Production wounds excited the crowds in the arena (Curry 2008). When
Region of Countries (million t) people think about barley use, animal feed, malt, and/or beer or
World 102 140 whiskey usually come to mind. However, as the term hordearii
Europe 37 88 implies, barley has been and still is used for food. Archaeological
European Union 27 58 evidence also suggests that barley was first used primarily for
Asia 31 20 food (Harlan 1979, Grando and Macpherson 2005). The useful-
West Asia 16 9 ness of fermentation was also discovered early (Hornsey 2003).
South Asia 7 5
As free-­threshing “naked” grains, such as wheat, evolved with
East Asia 5 3
Central Asia 5 3
agriculture and became increasingly popular, the predominantly
North America 2 14 hulled barley for human food was relegated to the poor, and its
Australia/New Zealand 2 8 use for animal feed became increasingly prominent (Grando and
Africa 16 6 Macpherson 2005, Newman and Newman 2008). Nevertheless,
North Africa 6 4 barley remains an important component of the diets of several
East Africa 6 2 cultures across North Africa, the Horn of Africa, the Middle
South America 8 3
East, and Himalayan nations to the present day (Grando and
a Source: FAO (2013). Macpherson 2005).
Globally, barley is currently primarily used for feed, at ap-
proximately 60–70% of the total production. Malt for brewing,
TABLE 1.4 distilling, and food makes up most of the rest at 30–40%, while
Barley Production Estimates by Country, Eight-Year Averages seed use is about 5%, and direct food use is only 2–4% (Newman
(2003–2010)a and Newman 2008). Barley for feed dominates in areas where
Production Harvested small grains dominate cereal production, notably areas that are
Rank Country (million t) (million ha) Yield (t/ha) unable to produce maize, such as northern North America and
World 142 55 2.6 Europe, high and/or dry areas of South America (in the Andes),
1 Russian Federation 16.6 8.4 2.2 Asia (the Middle East, central Asia, and the Himalayan regions),
2 Germany b 11.7 2.0 6.7 and Africa (North Africa and the Horn).
3 Canada 11.1 3.5 3.6 The composition of barley makes it particularly suitable
4 Franceb 10.7 1.7 7.1 as a high-­energy food and feed. In common with other cereal
5 Ukraine 9.3 4.4 2.5 grains, the relatively high energy content of barley is derived
5 Spainb 9.3 3.2 3.1 from starch. The composition data presented here are based on
7 Turkey 7.9 3.4 2.8 several studies and compilations (Åman et al 1985, Åman and
8 Australia 7.4 4.5 2.0 Newman 1986, Ullrich 2002, Newman and Newman 2008). Note
9 United Kingdomb 5.7 1.0 6.7
that these data represent average estimates for barley in order
10 U.S.A. 5.0 1.4 4.0
to give a broad overview, but the contents of components vary
11 Polandb 3.7 1.1 3.6
12 Denmark b 3.4 0.7 5.8
considerably among samples due to genetic and environmental
13 China 3.1 0.8 4.3
impacts. The ranges quoted do not necessarily include data for
14 Iran 2.9 1.5 2.1 particular extreme genotypes or mutants. For example, most
15 Morocco 2.1 0.5 4.2 commercial barley cultivars are hulled, but hull-­less types exist
16 Czech Republicb 2.0 … … and are important in some feed and food applications. Hull-­less
17 Kazakhstan 1.9 … … types (Fig. 1.4) tend to have elevated levels of most inner-­kernel
17 Finlandb 1.9 … … components because the fibrous hull is missing. The waxy, or
17 Belarus 1.9 … … high-­amylopectin and low-­amylose, trait in barley tends to be
20 Swedenb 1.6 … … associated with higher levels of soluble fiber. Extreme differences
a Source: FAO (2012b). can also exist in protein levels due to specific genetics and/or
b European Union member country. extreme levels of moisture or nitrogen in the environments. The
6 a Barley: Chemistry and Technology, 2nd ed.

concentration of starch in whole-­grain barley typically ranges Behall and Hallfrisch 2006, Pins and Kaur 2006, Lazaridou and
from 50 to 65% of the dry weight. Barley also has a moderate Biliaderis 2007). Similar effects on serum cholesterol and lip-
protein level of 10–16%, which is higher than that in many other ids were reported earlier in chickens and eggs (Bengtsson et al
cereals (e.g., maize and rice have <10%). Barley has a fairly bal- 1990), rats (Kalra and Jood 2000, Li et al 2003), and pigs (Bird
anced composition of essential amino acids, except for lysine, et al 2004). Barley has relatively low levels of lipids (2–3%) but is
which tends to be limiting for nonruminants. It also has rela- known to have relatively high levels of vitamin E and other to-
tively high contents of total fiber due to the hull (13–22%) and cols (Moreau et al 2007), which have antioxidant activity. It has
of soluble fiber (5–7%) due primarily to high levels of β-­glucan the highest levels of tocols in the grain compared with the other
(3–6%). β-­Glucan can cause digestive tract disruptions in poul- major cereals, according to Kerckhoffs et al (2002). Detailed re-
try but can have significant advantages for humans, as it has views of barley grain composition can be found in Gubatz and
been implicated in lowering blood lipids and cholesterol in hu- Shewry (2011) and in Chapters 4 (carbohydrates), 6 (protein), and
mans and reducing the glycemic index of foods (summarized by 7 (lipids) of this book.
The trade of barley essentially reflects barley use. The FAO
collects data on import and export activity for various forms of
TABLE 1.5 barley, malt, and beer (FAO 2012a). A general picture of the ex-
World Trade Estimates of Barley and Barley Products port and import of barley and barley products is given in Table
for 2003 and 2009a 1.5, which includes data for the quantity and value of products
2003 2009 for 2003 and 2009. In general, the quantity and value of exports
Commodity Exports Imports Exports Imports and imports increased in 2009 as compared to 2003. However,
Barley grain
some drastic differences can be seen between the two years for
Quantity (million t) 22 22 26 25 some products, i.e., the value of barley and barley products ex-
Value (billion $US) 3 3 5 5 ported and imported generally doubled between 2003 and 2009,
Malt whereas the differences were not as drastic for quantities traded.
Quantity (million t) 6 5 6 7 Table 1.6 presents a similar snapshot of trade, indicating the lead-
Value (billion $US) 2 2 4 4 ing exporting and importing countries. It is not surprising that
Beer the leading exporters (Table 1.6) were also generally the lead-
Quantity (million t) 8 8 11 11
Value (billion $US) 7 7 10 11
ing producers (Table 1.4). With the exception of Mexico and the
Malt extract Netherlands, which are not among the top 20 barley producers in
Quantity (thousand t) 96 148 107 159 the world (Table 1.4), the leading barley producers were the lead-
Value (million $US) 109 157 243 255 ing beer exporters in the world in 2000 and 2009. On the other
Pearled barley hand, the leading importers did not necessarily follow the same
Quantity (thousand t) 18 12 19 30 trends, i.e., low producers were not always the leading importers.
Value (million $US) 5 4 5 9 The top 10 barley beer producers, globally, for the three years
Barley flour/grits
Quantity (thousand t) 20 21 30 31
2006, 2008, and 2010 are listed in Table 1.7. The global produc-
Value (million $US) 4 4 12 10 tion and top annual producers of sorghum, maize, and millet
a Source:
beers for 2006 are listed for comparison. Whereas the produc-
FAO (2012a).

TABLE 1.6
Two-Year Snapshot of Leading Barley-, Malt-, and Beer-Exporting and Importing Countries, Showing Quantity Estimates (million t)a
2000 2009
Barley Malt Beer Barley Malt Beer
Exporters
World 24 World 6 World 6 World 26 World 6 World 11
Germany 6.2 France 1.1 Mexico 1.05 Ukraine 5.5 Belgium 0.97 Netherlands 1.7
France 4.8 Belgium 0.60 Netherlands 0.80 France 4.7 France 0.95 Mexico 1.6
Australia 3.0 Germany 0.55 Germany 0.79 Russian Fed. 3.5 Canada 0.63 Germany 1.4
Canada 1.8 Canada 0.50 Belgium 0.43 Australia 3.2 Australia 0.55 Belgium 1.0
U.K. 1.6 Australia 0.47 Canada 0.39 Canada 1.6 Argentina 0.48 U.K. 0.6
Importers
World 22 World 5 World 6 World 25 World 7 World 11
Saudi Arabia 5.4 Japan 0.74 U.S.A. 2.35 Saudi Arabia 6.0 Brazil 0.83 U.S.A. 3.00
China 2.1 Brazil 0.64 U.K. 0.44 China 1.8 Belgium 0.56 France 0.59
Japan 1.7 Russian Fed. 0.56 Italy 0.42 Belgium 1.6 Japan 0.50 Italy 0.58
Belgium 1.2 Germany 0.31 France 0.37 Netherlands 1.6 Mexico 0.35 Germany 0.57
Iran 1.0 Venezuela 0.26 Germany 0.32 Syria 1.5 U.S.A. 0.33 Canada 0.35
a Source: FAO (2012b).
 Barley Crop a7
TABLE 1.7
Leading Beer-Producing Countries Based
on Estimates of Quantity Produced
(million t), 2006–2010a
2006 2008 2010
Barley beer
World 165 176 169
China 35.9 41.4 36.9
U.S.A. 23.2 23.2 22.8
Russian Fed. 10.0 11.4 10.3
Germany 9.9 9.4 8.9
Brazil 9.4 10.6 11.4
Mexico 7.8 8.2 8.0
United Kingdom 5.4 5.0 4.5
Japan 3.8 3.4 3.5
Spain 3.4 3.3 3.3
Poland 3.3 3.6 3.4
Sorghum beer
World 6.9 … … Fig. 1.7. Spikes of hooded, six-­rowed barley, typical of forage culti-
Tanzania 1.92 … … vars. (Courtesy of Patrick M. Hayes, Oregon State University; used
Uganda 0.82 … … by permission)
Nigeria 0.79 … …
Burkina Faso 0.64 … … mal species. Barley grain may be fed whole and raw, dry-­rolled,
Congo 0.59 … …
steam-­rolled, or ground. It is fed primarily to swine (pigs) and
South Africa 0.56 … …
Cameroon 0.42 … …
ruminants (cattle, sheep, and goats) but is also used for poultry,
Ghana 0.34 … … dogs, and farmed fish (Bregitzer et al 2007, Blake et al 2011). As
Maize beer noted above, poultry are particularly sensitive to soluble fiber
World 2.5 … … components in the grain, so β-­glucanase is often included in the
South Africa 0.90 … … ration (Kellems and Church 2002). Barley plants are also used
Uganda 0.62 … … as forage, including hay, haylage (between hay and silage in
Canada 0.52 … …
moisture content), silage, and greenchop (high-­moisture feed,
Congo 0.14 … …
Zambia 0.14 … …
used directly from the field), as well as for pasturage and straw
Millet beer (Kellems and Church 2002). Barley for forage often uses culti-
World 1.5 … … vars that are awnless or hooded (Fig. 1.7) to avoid the barbed
Uganda 0.34 … … awns that are present on most barley cultivars. This is because
Tanzania 0.33 … … the awns, depending on the stage of maturity, can cause injury
Ethiopia 0.21 … … to the mouths of livestock. Forage barley is typically fed to ru-
a Source: FAO (2012c). minants and, to a lesser extent, swine. Comprehensive reviews of
feed uses and processing of barley can be found in Kellems and
Church (2002), Ullrich (2002), Blake et al (2011), and in Chapters
tion of these beers is relatively small compared to that of barley 9 and 10 of this book.
beer, it is important to the producing countries. All countries
listed for these other beers are from Africa, with the exception of
Canada for maize beer. The top 10 barley beer producers can be Malting, Brewing, and Distilling Uses
considered in three categories. China and the United States have The use of fermentation for making various types of bever-
been consistently the top two producers at roughly 20+ to 35+ ages is ancient, going back to at least the dawn of agriculture and
million tonnes per annum, followed by the Russian Federation, probably associated with permanent settlements (Harlan 1992,
Germany, Brazil, and Mexico (third to sixth ranking, respec- Hornsey 2003, McGovern 2003). Of course, the technology of
tively) at about 8–10 million tonnes, with the United Kingdom, malting, brewing, and distilling of barley has advanced through
Japan, Spain, and Poland at 3+ to 5+ million tonnes. Many fac- the millennia to the high level that exists today. At the same
tors, such as production levels, which are affected by climatic time, tradition plays a key role in the execution of these pro-
and economic conditions as well as political and population cesses, in no small measure because malting and fermentation
situations, may affect the amount and variation of barley and are natural biological processes. Whereas raw barley grain can
barley products that are exported and imported. contribute to the making of some distilled fermented beverages
(e.g., Japanese shochu and Korean soju), malting is a critical first
step in the making of beer and distilled liquors such as Scotch
Feed Uses whiskey and Irish whiskey. Malt also has uses beyond being a
Barley is prepared in various ways for feed, depending upon raw material for beer and whiskey. Malt extract and flour are
the objective (growth, maintenance, or finishing) and the ani- used in confections and in many wheat-­based baked products
8 a Barley: Chemistry and Technology, 2nd ed.
to add flavor and stability, while brewers’ spent grain is used
for feed and has potential use for food (Newman and Newman
2008). Comprehensive current reviews of malting, brewing, and
distilling can be found in Chapter 8 of this book and in Schwarz
and Li (2011).
Food Uses
As noted above, a relatively small proportion of the barley
produced globally (2–4%) is used directly for food. However,
some cultures depend heavily on barley for food. Furthermore,
the dynamics of barley food-­product development and use is
changing in many parts of the world where barley has never been
used extensively or has not been used extensively for many years.
This renewed interest in barley-­based foods is partly the result
of the increased emphasis on including a variety of whole grains
in the diet as a functional-­food approach to preventing or alle-
viating various diseases, including heart disease, hypertension,
diabetes, and obesity. Another significant aspect is the scientific
evidence that barley’s soluble fiber (mainly β-­glucan) can reduce
blood cholesterol and the glycemic index of foods, which can
impact both heart disease and type II diabetes. These findings
were the impetus for the U.S. Food and Drug Administration to
issue a health benefit endorsement for barley foods in 2006 (U.S.
FDA 2006). The increasing use of barley in the United States in
grain products such as multigrain breads, breakfast cereals, and
energy bars has been clearly noticeable in the last decade.
The methods for processing barley for food products include
pearling, grinding or roller milling into flour (whole-­grain or
sifted), steel cutting, rolling or flaking, extruding, and malt-
ing (Newman and Newman 2008). Pearled barley can be used
in several food preparations (such as soups, pilafs, and casse-
roles), while flour from the whole grain or pearled grain can be
included with wheat or rye flour in yeast breads or used alone
in flat breads, pastas and Asian noodles, pastries, etc. Steel-­cut,
flaked, or extruded barley can be used in breakfast cereals and
snack foods. Malt flour is used in an array of wheat-­baked prod-
ucts and can be used for confectionary products. In principle,
barley can be used for almost any product currently made from
wheat, rice, or maize. In fact, the potential is far from being real-
ized commercially in most industrialized nations.
In certain areas of the world today, barley is an important
food ingredient, regularly used in the human diet. These areas
tend to be where other cereals do not grow well, such as arid and
high-­elevation regions of countries in North Africa (Morocco,
Algeria, Tunisia, Libya, and Egypt), in the highlands of the Horn
of Africa (Ethiopia and Eritrea), in the higher dry lands of the
Near East (Iran and Yemen), in the highlands of Central Asia
(from Tibet and Nepal to the republics to the west), and in the
Andean countries of South America (particularly Ecuador, Peru,
and Bolivia). Communities in several East European countries
have also retained traditional barley foods, notably in Estonia,
Latvia, Lithuania, and Moldova (Grando and Macpherson 2005).
The traditional use of barley foods in Nordic and Far Eastern
countries has decreased relatively recently, but these foods are
still popular in certain situations or areas.
Relatively recent comprehensive reviews of barley food pro-
cessing and food use include Grando and Macpherson (2005),
Baik and Ullrich (2008), Newman and Newman (2008), Baik
et al (2011), as well as Chapters 9 and 10 of this book.
Other Uses of Barley
Barley also has other uses, either current or potential. Some
of these are industrial uses of barley starch for paper and fab-
ric manufacturing, paper pulp, construction-­grade fiber board,
biofuels, and industrial chemicals such as ethanol and methanol
(Bjørn Petersen and Munck 1993, Munck 2004). The biofuels in-
dustry is booming in some areas of the world, with ethanol pro-
duction from barley and other grains, and the potential exists for
the production of ethanol from cellulose in barley straw (Kim
et al 2008, Mullen et al 2010, Nghiem et al 2011). See Chapter 11
of this book for a detailed summary of barley use for biofuels.
A myriad of products for various food, feed, and industrial
uses could be derived from barley grain and straw, compara-
ble to those that have been produced for decades by dry and
wet milling of maize grain (Griffey et al 2010). The germina-
tion of barley grain and the fermentation of barley grain and
straw could also generate many products for various industries
(Munck 2004).
References
Åman, P., and Newman, C. W. 1986. Chemical composition of some
different types of barley grains in Montana, U.S.A. J. Cereal Sci.
4:133-­141.
Åman, P., Hesselman, K., and Tilly, A.-­C. 1985. The variation in the
chemical composition of Swedish barleys. J. Cereal Sci. 3:73-­77.
Azhaguvel, P., and Komatsuda, T. 2007. A phylogenetic analysis based
on nucleotide sequence of a marker linked to the brittle rachis
locus indicates a diphyletic origin of barley. Ann. Bot. (Lond.)
100:1009-­1015.
Baik, B.-­K., and Ullrich, S. E. 2008. Barley for food: Characteristics,
improvement, and renewed interest. J. Cereal Sci. 48:233-­242.
Baik, B.-­K., Newman, C. W., and Newman, R. K. 2011. Food uses of
barley. Pages 532-­562 in: Barley: Production, Improvement, and
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Barley Grain: Development and Structure
The outer appearance of the ripe barley grain is deceptively
simple, but its inner organization is complex, reflecting a pro-
gression of developmental steps. Grain development requires a
supply of assimilates and nutrients, at first for the initiation and
growth of the maternal and filial tissues and then to establish
reserves for early seedling development after germination. The
mobilization, influx, distribution, and storage of assimilates and
nutrients are therefore major processes that clearly impact tran-
sient and final grain structures. Maternal-­filial communications
and sink-­source interactions among the grain tissues regulate
normal grain development, which occurs in defined develop-
mental phases.
We describe the developmental biology and structure of bar-
ley grains with regard to these aspects.
From Inflorescence Initiation
to Flowering
Setting Up Reproductive Structures
The barley inflorescence develops from rows of floral meri-
stems that are generated by the shoot apex. After the transition
to sexual reproduction has occurred, double ridges grow on both
sides (Fig. 2.1a–d,h,k) of the elongating shoot apex (Bonnett
1966, Delécolle et al 1989). The upper ridge, the floral meristem,
is initiated together with the meristem of its subtending leaf, the
development of which is later suppressed (Whipple et al 2010).
The transcription factor ramosa2 (ra2) is expressed in the cells
that will constitute the floral meristem in the axil of the bract
(Bortiri et al 2006).
The developmental transition from a vegetative to a floral
meristem is under the control of the MADS-­box transcription
factor VRN1 encoded by the Vernalization 1 gene (Danyluk et al
CHAPTER 2
Sabine Gubatz
Rothamsted Research
Harpenden, Hertfordshire, United Kingdom
Winfriede Weschke
Leibniz Institute of Plant Genetics and Crop Plant Research
Gatersleben, Germany
2003, Hemming and Trevaskis 2011). Vernalization 1 belongs to
a network of genes (Hemming et al 2008, Distelfeld et al 2009)
that together adjust flowering time in barley to environmental
conditions. This group includes genes for photoperiod percep-
tion and regulators of vernalization (Casao et al 2011). The tim-
ing of apex transition has been correlated with thermal time or
with the number of leaves generated (Appleyard et al 1982, Hay
and Ellis 1998, McMaster et al 2005, Arduini et al 2010).
Primordia
The growth and size of primordia change along the ear. Close
to the collar and toward the tip, their size decreases (Fig. 2.2a),
and about a quarter of the initiated primordia do not develop
fertile florets (Kirby 1977). Primordia are generated at regular
intervals over a longer time, but, at flowering, the spikelets of
the ear ultimately flower together in two or three days. This syn-
chronicity is achieved around the end of the primorida initiation
phase by earlier variation in growth rate along the ear (Kirby
and Appleyard 1984). However, a developmental gradient per-
sists along the inflorescence, its lower median part being most
advanced. The heaviest grain later develop here, and this can be
related to the width of primordia as early as the double-­ridge
stage (Fig. 2.1h,i) (Cottrell and Dale 1984) and to a bigger carpel
size at anthesis (Scott et al 1983).
Spikelets
Each floral meristem ridge differentiates a central and two
lateral mounds (triple-­mound stage) (la in Figs. 2.1 and 2.2) and
ultimately develops three spikelets at each node on alternat-
ing sides of the rachis. In the six-­row barleys, all three spikelets
produce grain; in the two-­row barleys, only the median, sitting
a 11
12 a Barley: Chemistry and Technology, 2nd ed.

Fig. 2.1. Establishment of reproductive structures during early inflorescence development. Scanning electron micrography (SEM) of the devel-
oping shoot apex (a–g) and detailed views (h–m). a and b, early development; c, shoot apex emerging from the collar leaf; d, leaf initiation on the
shoot; e–g, ongoing differentiation of the central and lateral (la) spikelet and the glumes (G); h, developing double ridges; i, upper (u) and lower
(lo) part of the developing double ridge; k, further development of the upper ridge; l and m, differentiation of the lateral spikelet, the glumes,
and the lemma (L). Bars are 20 µm in h, 50 µm in i and l, and 100 µm in every other case. (Jean Devonshire and Sabine Gubatz, unpublished
data, 2012). SEM data shown in Figures 2.1, 2.2, 2.5, and 2.7 were acquired on a JSM Jeol (UK) 6360 LV-­SEM using a GATAN (UK) ALTO 2100
cryo system.

spikelet is fertile (Bonnett 1966, Kirby 1977). The rudimentary
character of the lateral spikelets in the two-­row forms (Fig. 2.3)
is linked to the activity of the transcription factor VRS1 in the
primordia of the lateral spikelets at the triple-­mound and glume
primordia stages (Komatsuda et al 2007, Gottwald et al 2009,
Ramsay et al 2011).
A spikelet in barley comprises a single floret associated with
a pair of sterile bracts called glumes. Each fertile floret is com-
posed of lemma and palea, a carpel with a single ovule, three an-
thers, and two lodicules (Figs. 2.3 and 2.4). The floral axis contin-
ues as a vestigal rachilla (Forster et al 2007), which is either short
or reaches half way up the palea (von Bothmer et al 1991), posi-
tioned (above the palea) against the ventral crease of the grain.
The rachilla characteristics are of taxonomical significance in
Hordeum (Bell and Carson 1941, Marchylo and Laberge 1981,
Javaid et al 2009). The inflorescence itself is indeterminate, as
the ear does not build a terminal spikelet. The lemma and palea
are capable of photosynthesis, (Abebe et al 2004, 2009), persist
during grain development, and finally adhere to the surface of
the ripe grain (Figs. 2.3, 2.5, and 2.6) due to a lipid layer on the
pericarp epidermis that is not produced in free-­threshing hul-
less (or naked) barleys (Taketa et al 2008).
Flower ORGAN IDENTITY
Floral homeotic genes are essential for specifying flower
organ identity in angiosperm flowers, and class B, C, D, and E
MADS-­box genes have been identified in the grasses (Thompson
and Hake 2009). However, grass florets include lodicules, lemma,
and palea but not petals and sepals, and the homologies of some
organs are difficult to establish (Kellogg 2001, Rudall et al 2005,
Yoshida 2012). Lodicule identity is promoted by class B MADS-­
box genes (Whipple and Schmidt 2006, Whipple et al 2007),
which, in angiosperm flowers, specify petal identity together
with class A and class E genes (Erbar 2007). The homology of
the lemma and palea is disputed (Prasad et al 2001, Bommert
et al 2005, Preston et al 2009); they may relate to sepals in grasses
(Ciaffi et al 2011) or represent leaflike structures. If so, the lemma
may be interpreted as the bract subtending the floral branch,
and the palea as the first bract on the floral branch, a prophyll
(Clifford 1987, Chapman 1996).
Spikelet differentiation
Spikelet differentiation after the triple-­mound stage begins
with the empty glume initials and then the prominent lemma
primordium (Figs. 2.1g,m and 2.2a). The differentiation of the
anthers and carpel follow, the differentiation of the lemma awn
meristem in the developing ear being the final visible step of
differentiation before primordia generation stops (Fig. 2.7). The
lemma and palea grow only above the developing anthers before
heading. Both the empty glume initials and the lemma primor-
dium are visible from the abaxial side of the apex (Bonnett 1966,
Kirby and Appleyard 1984) (Fig. 2.2b). The developing lodicules

Fig. 2.2. Development of the inflorescence. a, scanning electron mi-
crography showing the more-­advanced development in the middle
of the ear; b, detailed view of developing anthers and lemma; c,
schematic view of lateral spikelet (la), glumes (G), anthers (ST), and
lemma (L). Bars are 200 µm in a, 100 mm in b. (Jean Devonshire and
Sabine Gubatz, unpublished data, 2012)
are covered from view between the primordia of the lemma and
the anterior stamen. On the adaxial side, the differentiating
palea is visible (Fig. 2.7c). The rachilla grows between the lemma
margins. The meristem between the three stamens (Fig. 2.7c)
eventually develops into the female flower organs (Sattler 1973).
Carpel
Development of the carpel begins with growth on the ante-
rior (abaxial) side of the meristem between the three developing
stamens. The carpel grows upward and gradually folds around
and over the early ovule (Barnard 1955, Klaus 1966, Philipson
1985). The style then develops from the two sides of the carpel
wall, which grow, elongate, approach each other, and finally join.
A description of 20 stages of floral development from the transi-
tion apex to anthesis notes 13 different observations of the carpel
after initiation of the gynoecial primordium (Waddington et al
1983). The single carpel in the Poaceae is possibly the result of
a reduction of a tricarpellary structure (Philipson 1985, Rudall
et al 2005).
Grain Development and Structure a 13
Fig. 2.3. Fertile central and lateral sterile spikelets on the ear of a
two-­rowed barley during development (upper panel) and at ripeness
(lower panel). Note the pair of glumes (G) sitting on the rachis (R)
that is associated with the central spikelet, the lemma (L), and lemma
awn (La) of the central spikelet, and the glumes (g), palea (p, see
upper middle image), and lemma (l) of the lateral spikelet. The figure
at the lower right of the panel additionally shows the palea (P) with
its rounded tip (TP).
Carpel and ovule development involve orthologs of the re-
spective Arabidopsis genes, but duplications and subfunctional-
izations occurred in the Poaceae (Ferrándiz et al 2010).
Ovule
The ovule, consisting of the nucellus and two integuments,
develops on the ovary wall without a funicle (Greenham and
Chapman 1990). The chalaza (the region where integuments and
nucellus merge) is in contact with the placenta of the ovary wall,
and tissues are not clearly delineated. This contact zone between
the ovule and carpel wall elongates with the growing grain and
finally becomes oriented along the base of the crease (Frazier and
Appalanaidu 1965, Shamrov 2002). The integuments originate
from the base of the ovule primordium (Reiser and Fischer 1993,
Rudall 1997). In barley, both comprise two cell layers and cover
the surface of the ovule at anthesis, but only the inner integu-
ment builds the micropyle. At the time of megagametogenesis,
the ovule and ovary are not oriented in the same direction but at
14 a Barley: Chemistry and Technology, 2nd ed.

Fig. 2.4. Barley floret around the time of anthesis. a, closed floret at anthesis; b, floret after removal of the lemma, with
arrows indicating the detailed views in e and f; c, anther at anthesis; d, anthers on the day after flowering; e, detailed
view of stigma branch with pollen; f, detailed view of turgescent lodicules at flowering; g, style before pollination. Bars
are 5 mm in a, 2.5 mm in b and d, 1 mm in e and f, and 0.5 mm in g.

a right angle (90°). Because of unequal growth at the base of the
ovule, its orientation changes gradually to the almost anatropous
final orientation (Krauß 1933).
In the tenuinucellate nucellus (Rudall 1997) a subepidermal
cell differentiates directly as the archespore (Norstog 1974). The
specification of cells as gametes under the control of Argonaute
9 has been shown to involve small RNAs (Olmedo-­Monfil et al
2010). In maize, the loss of a DNA methyltransferase function
leads to the formation of more than one embryo sac, indicat-
ing the epigenetic character of the process (Garcia-­Aguilar et al
2010, Gutierrez-­Marcos and Dickinson 2012). During meiosis,
the megaspore mother cell is surrounded by a callose special
wall (Cass et al 1985). Only the chalazal megaspore of the re-
sulting linear tetrad develops further, and the megagametophyte
development follows the Polygonum type. The vacuoles become
prominent at the four-­nuclear stage, and one persists from the
eight-­nuclear stage to become the characteristic vacuole of the
central cell. A possible early role of the vacuole could be to es-
tablish the polar spatial organization of the megagametophyte
(Cass et al 1985). Comparable development of vacuoles can be
observed in other members of the Triticeae (Hause 1991).
Meiosis
Female and male meiosis are coordinated in a single flo-
ret; at 20°C, they were timed to last 43 h (Bennett et al 1973).
Environmental conditions influence the synchrony and duration
of male and female meiosis (Lindgren et al 1969, Bennett et al
1973). High temperature stress at this stage of development can
lead to severe reduction of fertility (see below) (Sakata et al 2000).
Primordia Supply and Survival
Light, nitrogen supply, and genotype influence the number of
generated and surviving fertile florets in barley (Arisnabarreta
and Miralles 2010). When the maximum number of primordia
Another random document with
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 Obad. Gosh all fog horns, no! I found the money box! Not a cent
gone! Everything upset from one end the place to the other, and
nothing took! That young Gordon chap must be a lunatic. Cranberry
has gone his bail, but I reckon he ought ter be in an asylum instead
o’ traveling around loose.
Lem. Wal, it’s the durndest piece o’ business I ever heard tell on,
and everybody don’t believe he took all those pipes even if there is
so much evidence against him. Some says it was a big fellow and
some says it was a little one.
Obad. I stick to it the man that stole my pipe wuz a little bit of a
feller. He up behind me and put his hand in my pocket (illustrating as
he talks) an’ scat before you could so much as wink.
Lem. Yes, an’ Jim Hincks says he wuz a big feller.
Abner. And Sammy said so once, and then he shied off and
wouldn’t say much of anything.
Obad. No wonder. The way Lem talks at the kid is enough to
frighten a whale.
Lem. An’ Cranberry didn’ know when his pipe was took and
wouldn’t say nothin’ if he did. He’s took an awful shine to Miss
Cynthy’s boarder.
Obad. Say, suppose we find the feller and tell him we found out he
didn’t take nothin’ from the post-office an’ see what he says.
All. That’s right. Good idee!
Lem. Maybe he’ll own up ’bout the pipes if he finds out the charge
agin him ain’t so serious.
Obad. Maybe so, but it’s my opinion the feller is plumb crazy.
(They all start to exeunt. Ariel comes to door followed by Hep.)
Ariel. Father, are you here? (Stops abruptly.) Oh, a meeting of the
vigilance committee? Is Sammy up here?
Abner. No.
 Ariel. Hepsy can’t find him, and sometimes he comes up here
when I am home, you know, so I thought perhaps he was here.
Hep. My land! I’ve been all over town. I’m beat out. He didn’t go to
school and I ain’t seen him since he was over to Cynth’s this
morning.
Obad. By tunket, I don’t blame the kid. He will run away some o’
these days an’ never come back, an’ it’ll serve Lem durned right.
Lem. (angrily). What in thunderation would I do? Let him grow up
to be a teetotal liar?
Obad. Gosh all fog horns, yes, if he wants ter be! Half the world
gets their living that way!
(All exeunt but Ariel. Abner comes back.)
Abner. You are going to stay up here, Arey?
Ariel. I think so, dad, a little while. I haven’t been up here since
last summer.
Abner (uneasily). You—you were always happy up here, little girl?
Ariel. Yes, dad, and happiest when I knew you were down-stairs.
Abner. And I was happy when I knew you were up here. Arey, I
have tried to be a good father to you and give you a happy home.
Ariel (going to him and putting her arms around him). You have,
dad! You have! No girl could ask for a kinder father or a happier
home.
Abner. And I want to know you are going to have a happy home
when I’m gone. Arey, Nat’s really in love with you.
Ariel (turning away from him). Oh, father!
Abner. If you knew it meant a good deal to me, would you tell him
yes? To please me? To help me? He could give you a good home.
Could make you happy.
Ariel (slowly turning to him). Do you mean—there is a—a reason
why you want me to marry Nat? That I really could help you by
marrying him?
 Abner. Yes, that’s what I mean.
Ariel. I can’t understand how such a thing can be, but if that is the
case I will do it for you, if you will tell him I don’t care for him and
explain the reason I am marrying him. Will you do that?
Abner. Yes.
Ariel (turning away). Very well.
Abner (stepping toward her). Arey!
Ariel (with an effort). Dad, will you go away please and leave me
alone?
Abner (hesitating a second). Very well.
(Exits with bowed head. Ariel gives a long sigh, goes to c. door
and stands looking out for a second, then goes to doll house,
opens door and takes out a large doll. She goes to bureau,
unlocks a drawer, takes out a small box. Suddenly leans forward
on the bureau, her head on the doll.)
Ariel. Oh, Alicia, has it come to this? (Sam. crawls cautiously out
from under sofa. He steals toward the hammock and throws a pipe
into it, then starts for door. Ariel suddenly turns.) Who is it? Who is
there?
(She starts forward. Sam. hastily crawls under sofa again. Ariel
walks forward with the little box in her hand.)
Ber. (outside). Ship ahoy! Anybody on deck?
Ariel (going to door). Father isn’t here, Captain Berry. I’m up here
alone.
Ber. That so? Can I come up?
Ariel. Why, certainly, if you like. (Ber. enters.) This is my
playhouse up here. I have spent the happiest days of my life up here
in this old room. I think I have spent the only happy ones I will ever
know.
Ber. Nonsense! That’s a pretty way for a young girl like you to be
talking, jest when you got all your life before you.
 Ariel. I wish I didn’t have. I wish this was the last day of my life.
Ber. (aghast). Miss Freeman, you don’t know what you are
saying. Something must have happened to pretty much upset you.
You don’t look like a coward. I can’t somehow believe you are one.
Ariel (surprised). A coward?
Ber. It’s only a coward, a pretty contemptible one at that, who
would rather die than face what’s coming to him.
Ariel (suddenly standing straight and throwing her head up). You
are right. I didn’t think of that. Thank you, Captain Berry. That was
just what I needed to hear. You—you spend a good deal of time over
here in town, don’t you?
Ber. (smiling). Is that a reflection on my duties as keeper of Bay
Point Light?
Ariel (hastily). Oh, no, no, no! I didn’t mean that. I was just
thinking that you must like our town. I know you have an assistant,
and any one doesn’t have to meet you but once to realize that your
duty would stand before anything else in the world.
Ber. (pleased and touched). Why, Miss Freeman, I thank you for
that from the bottom of my heart. You can’t understand how much I
appreciate that coming from you.
Ariel. Why from me, especially? Oh!
(Drops the little box and a little ring.)
Ber. (picking them up and looking at the ring fascinated). What is
this?
Ariel (slowly). That—that is a link, Captain Berry. The only
connecting link between Ariel Freeman and the girl she really is.
Ber. (breathing hard). What do you mean?
Ariel. I suppose you know I’m not Captain Freeman’s daughter?
Everybody knows.
Ber. Yes, I’ve heard.
 Ariel. That little ring is the only thing which could throw any light
on who I really am. That was on my finger, and that was absolutely
all there was to help.
Ber. How did this help?
Ariel. There is a name engraved inside the ring.
Ber. (slowly). And the name is Alicia.
Ariel. Why, who told you that?
Ber. (realising what he has said). Why—why—it’s right here!
(Hastily examines the ring.) You can see it for yourself!
Ariel. You have wonderful eyes. I have always known what the
name is, and I have hard work to trace it. You see what a curious
little old-fashioned ring it is; it has a German mark in it. Captain
Freeman traced this to a German jeweler who in 1878 engraved the
name Alicia in a baby ring for a Mrs. Emerson. The address was a
hotel in Berlin. Father moved heaven and earth to trace it still further,
but that was all he ever found out. He decided that my mother must
have been Alicia Emerson, but whom she married, who my father is,
will probably always remain unknown. Somewhere in the world I may
have a mother—a father.
Ber. You have been happy here?
Ariel. Oh, yes, indeed. I couldn’t have had a kinder father than
Captain Freeman. Gracious, Captain Berry, I can’t imagine why I
have told you all these things! I never speak of them to any one but
Miss Cynthy. Somehow you seem to be the kind of man one tells
their innermost secrets to. It was so kind of you to help Lee Gordon
this morning.
Ber. Nonsense! Nothing kind about it. Justice, that’s all. He ain’t
any more guilty than I am.
Ariel. You really think so?
Ber. Certain! One look at his face will tell you that.
Ariel. But the evidence against him? Such a lot of it!
 Ber. By tunket, the circumstantial kind or I’ll eat my hat.
Ariel. Why, Captain Berry (suddenly grabbing pipe from
hammock), look! Here’s a pipe in my hammock! What in the world
——
Ber. (taking pipe). Jumping jingoes! Seth Mason’s! Got his name
on it!
Ariel. I thought some one was up here just before you came up. It
seemed to me that some one who was small was moving around. I
was over by the bureau and I can’t see across the room, you know. I
was startled for a second and by the time I got my senses together
to cross the room they were gone.
Ber. Um! So? (Looks about. Crosses room away from Ariel. He
suddenly stoops and looks under sofa.) This is a queer mix up, ain’t
it? (Bell rings.) What’s that?
Ariel. That’s the house bell. Dad had it connected so we could
hear it when we were over here and Hannah was out. I think she is
out now, I will just run over to the house if you will excuse me.
Ber. Certain! (Ariel exits. Ber. goes to sofa.) Come out! Come
out here, I say! It’s no use, Sammy! I have caught you square! You
might as well come out! All right then! I’ll bring you out myself.
(Reaches under sofa and pulls Sam. out by one foot. Sam. yells and
kicks.) Belay there! That ain’t no way to act with your old uncle. I ain’t
going to hurt you. (Sits down on sofa and holds Sam. in his arms.)
Now hush up, and tell Uncle Cran the whole business.
Sam. (struggling to get away). No! I ain’t got nothin’ to tell!
Ber. Oh, yes, you have, Sammy! Sit still! You tell Uncle Cran what
you stole all those pipes for. I was the only one in the room that was
watching you this morning, and I guessed somewheres near the
truth. You were frightened when that pipe fell out of Mr. Gordon’s
pocket, and then when you found that no one was going to suspect
you, you opened up the sugar-bowl and took the other one out, and
now you just brought this one up here and put it in Arey’s hammock.
Sam. (thoroughly frightened). Pa will kill me!
 Ber. No, he won’t. Now listen, Sammy, this whole thing has got to
come out some way. Your father will get it out of you, and you know
how he will get it. Now, you tell me the truth and I’ll promise you that
I will make things right with your father. I won’t let him touch you. You
can trust me, can’t you?
Sam. (holding him around the neck). Yes, Uncle Cran.
Ber. Well, then, when did you get a chance to go up into Mr.
Gordon’s room and leave those three pipes there?
Sam. I didn’t take those.
Ber. (reproachfully). Now, Sammy, I said if you told me the truth.
Sam. That’s the truth, Uncle Cran! I took this pipe, and the one in
the sugar-bowl and the one in Mr. Gordon’s pocket, but I didn’t take
the others.
Ber. By Crismus, you look as if you were telling a straight story.
Why in the name o’ all that’s sensible did you steal any of the pipes?
Sam. Well, a man took dad’s away from me, an’ maw didn’t believe
it and I knew there wouldn’t any one believe it, and pa whales the
stuffins out of me for telling things and—and—I thought if some more
people lost their pipes he’d believe me. So I stole three and—and—
pa had to believe me.
Ber. (striking his knee). Well, by tunket, if that ain’t one on Lem!
(Suddenly looks serious and speaks half to himself.) But belay there,
Cranberry! You ain’t got but half the story yet! There’s those other
pipes! Well, Sammy, I’ll pull you out of this some way, although I
don’t know as it’s the right thing for me to do.
Sam. (stretching). Oh, I’m awful tired! I been layin’ under this sofa
awful long. More’n an hour, more’n three hours, I guess. Nat
Williams was up here an’ Cap’n Abner. Say, Uncle Cran, it was
Cap’n Abner that broke into the post-office last night.
Ber. (at the top of his voice). What?
Sam. He was hunting for a paper. And—and—Cap’n Abner cheated
Miss Cynthy. Nat said the land this house stands on belongs to her,
and—and—Cap’n Abner cheated Nat’s father, too,—and—and—
Cap’n Abner is going to make Miss Arey marry Nat—and—and—she
cried over there (pointing to bureau) on top of her doll—and—and—I
guess I don’t remember any more.
Ber. Well, that’s quite sufficient if it’s truth you are telling.
Sam. Of course it is! I wouldn’t tell you any stories, Uncle Cran.
Ber. (a trifle dazed). All right. Now listen, Sammy, you forget this
stuff you have been telling me jest as soon as ever you can. Don’t
you breathe a word of it to a living soul. If I ever hear that you have,
I’ll forget to make peace with your father, and there won’t be any
more trips with me over to the Point to visit the light.
Sam. Oh, I’ll never tell, Uncle Cran! Honest! Hope to die!
Ber. All right. Now you go and find Cap’n Abner and tell him that I
am up here and want to see him. Then you go home as hard as you
can pelt. Your mother is looking for you.
Sam. (exits). All right, Uncle Cran.
(Slight pause. Ber. sits on the sofa looking down at the floor.)
Ber. Great jumping jingoes!
Enter Lee and Peter.
Lee. Captain Berry, I was just talking with Miss Freeman and she
said you were up here. I can’t tell you how much I appreciate your
kindness to me. I don’t see how you ever happened to stand by a
stranger the way you did.
Ber. I guess it was because you are Miss Cynthy’s boarder.
Lee. Captain Berry, this is Mr. Pomeroy and (laughing) he is the
best friend I have got on this earth.
Ber. (shaking hands with Peter). Well, he certain didn’t act the
part this morning.
Peter. Entirely his own fault, I assure you.
Lee. Say, Captain, may we talk with you for a little while?
 Ber. Certain! Glad to have you. Sit down.
Lee. You have proved such a good friend that we are tempted to
make a clean breast of things and tell you our part in this affair.
Ber. Heave ahead! If you can throw a search-light on the mystery
it will be a good thing.
Lee. It is true that I have been expelled from college and that I’m in
trouble with my father.
Peter (heartily). He isn’t to blame in either case, Captain Berry.
Lee (to Peter). Thanks, old man. (Turns to Ber.) When I was at
Yale I was connected with a college magazine and I have several
times been told that I had a great future ahead of me as a
newspaper man or magazine editor. My friend Pomeroy here had to
leave college last year because an invalid uncle who had brought
him up was dying. The uncle left Pete all his money and when I got
into a mess with dad, Pete suggested that he buy a magazine he
had heard about that was on its last legs and see if we could give it a
brace. I jumped at the chance. It was what I would most like on
earth. Pete bought it and we have started to publish and edit a
startling fiction magazine called The Red Cap. For a starter we have
announced a serial detective story—“The Mystery of the Seven
Pipes.”
Ber. Well, by tunket!
Peter. It’s going to be the greatest thing on record. We write all
but the last chapter and offer a prize for the best solution of the
mystery. Gee! Folks run after the first issue as if it was soothing
syrup and they had a kid with the colic. I had been down on the
Cape, so I suggested that we lay the scene down here somewhere.
Lee. And to give local color I suggested that we come down and
carry out some of the story ourselves. Last night when we arrived
there was a thick fog and it seemed a good chance to make a start.
We drew lots and it fell to Pete to steal the pipes.
Peter. We planned on stealing seven but I only got as far as
three.
 Lee (laughing). For the love of Pete, have you any idea who took
the other four?
Ber. Yes, by tunket, I have! This beats the Dutch and no mistake!
You go find Obed and Lem and tell them what you have told me.
Then you bring them back here and we will clear up the mystery.
Lee (jumping up). Done, by jingo!
Peter. Lead us to it!
[They exeunt.
(Slight pause. Abner enters.)
Abner. Did you send Sammy after me?
Ber. Yes, I did. I’ve got something to say to you and I thought this
would be a good place up here. Cap’n Freeman, I’m Ariel’s father.
Abner (staggered). What? What do you mean? (Suddenly
growing furious.) What kind of a trick is this? Do you expect me to
take your word for it? What’s your game?
Ber. (calmly). I married Alicia Emerson twenty years ago this
month. Two years later she and our year old daughter made a trip
with me. We ran into a storm and were wrecked. My wife’s body was
recovered. The child’s was not. I supposed her dead, of course. She
wore a ring, a ring that had been her mother’s. It had her name,
Alicia, engraved in it, also the mark of a German jeweler and——
Abner (hoarsely). Enough! How long have you known this?
Ber. I guessed it last night when I saw Alicia’s face in your
daughter. I made sure of the truth this afternoon.
Abner. Well, what are you going to do? She is under age. The law
would give her to you, I suppose. Are you going to take her away
from me after all these years?
Ber. You love her?
Abner. How can you ask such a question? She is all I have! Ask
any one in Bay Point. Ask the girl herself!
 Ber. (quietly). And yet loving her you ask her to marry a man she
doesn’t love. Kind of curious, ain’t it?
Abner (furiously). Now see here, Cranberry, that’s my business.
Ber. (decidedly). It’s mine, also. She is my daughter. If you will not
give the girl happiness, I will take her from you and give it to her
myself.
Abner (desperately). You mean that—that——
Ber. Just this! Drop this Nat Williams business and never take it
up again, and give Miss Cynthy Tinker what you owe her.
Abner. What do you know about Cynthy Tinker?
Ber. What does it matter? I know you have been cheating her for
years. Do you want me to go into details?
Abner (hastily). No. Supposing I refuse?
Ber. Well, it will be my gain. I shall be able to find a little
happiness with my daughter. I only had her a year. I only had her
mother two years. Then they were both taken from me at the same
time. I’ve had a lonely life, always at sea or keeping a light on some
dismal point. Often when the gales have come and the storms have
lashed around the old house and there’s been nobody but jest old
Cranberry and his pipe, I have looked around my lonely settin’ room
and wondered how it would seem to have Alicia on the other side of
the fire and a little girl on the floor near by.
Abner. My heaven, Berry, I hadn’t thought of your part of this!
Ber. No, we usually get our own point of view. Mighty seldom we
get the other fellow’s.
Abner. And if—if I agree you are willing to swear you will not take
the girl?
Ber. On my word of honor. You do love the girl after all.
Abner. But great heaven man, I can’t square things with Cynthy
Tinker without having people know the facts. Nat will tell.
 Ber. Why don’t you do the telling yourself? If you have been
cheating perfectly innocent people I think you will find that
confession is a good thing for a guilty conscience. Anyway you have
two paths to choose from, and you know what is at the end of each.
(Lee and Peter rush in, Ariel follows. Obad. and Nat. and Lem.,
Cyn. last with a letter in her hand.)
Lee. Well, Captain, we have told them! And it didn’t kill them
either!
Peter. Yes, they rejoice in all the horrible details.
Obad. Gosh all fog horns I should say we did! Did any one ever
hear the like? A story! “Mystery of the Seven Pipes!”
Lem. I always said writers wuz half cracked. I never was so sure of
it as I am this minit! But the mystery ain’t all cleared by a long sight.
Mr. Pomeroy only took three pipes. Who took the other four?
Ber. I suppose, Lem, as constable of Bay Point, you would like to
have the mystery cleared up?
Lem. I sartain would, and then I could go back to work at
something sensible.
Ber. Well, I’ll tell you what you want to know if you’ll jest do me a
little favor.
Lem. (puzzled). Why, sartain, Cap’n, always glad to do anythin’ fer
you.
Ber. Well, Sammy took three pipes.
All. Sammy?
Lem. My Sammy?
Obad. Gosh all fog horns!
Ber. Mr. Pomeroy stole yours from Sammy. You wouldn’t believe
the boy, he was afraid of you and he took some pipes on his own
hook to make his story sound true.
Obad. Rev. Mr. Peters said he was smart and by Crismus, he is!
He’s too smart fer you, Lem!
 Lem. Wal, he’ll find out how smart I am when I get home.
Ber. Lem, you promised me a little favor, you know?
Lem. Sure!
Ber. You are not to touch Sammy.
Lem. (astonished). I’m not to—— (Suddenly stops and grins.) Well,
all right. You’ve got me, Cap’n. You always did spoil that kid. Wal,
there’s still another pipe.
Abner. Yes, mine. I dropped it when I broke into the post-office
last night.
All. What? You? Abner!
Obad. Fer the love o’ Admiral Sampson what fer?
Abner (steadily). I wanted to find some old papers in a business
deal between your father and me. I suppose you remember that your
father sold me this whole shore property here?
Obad. Yes, sartain.
Abner. Well, I found out right after I bought it that half the land
didn’t belong to him to sell. There was a mistake in his title and some
of the land belonged to Nat’s father and some of it belonged to Miss
Cynthy’s father.
Cyn. Land o’ goshen!
Abner. They didn’t seem to know. They must have thought old Mr.
Daniels knew what was his property. I didn’t say anything. You know
why I wanted the property and the money I have made here. I knew
if they realized this land was theirs they wouldn’t have sold. They
would have used it themselves. I knew I had a good thing and I kept
it. I made a pile and I kept them from their chance of making money
when shore property was worth its weight in gold. That day has long
gone by, but I’m ready to pay Miss Cynthy whatever seems right. I’ll
leave it to Cap’n Cranberry to figure it out. I have offered money to
Nat and he has refused to take it. He wants my daughter instead.
Well, he had better take my money, for he can’t have my daughter
unless she wants him.
 Ariel (joyfully). Father!
Nat (furiously). Thunderation!
[Exit.
All. Good work! That’s the talk, Cap’n Abner!
Abner (slowly). I have deceived and cheated all these years. Of
course I realize that I’ve got to lose my friends, that I will be the talk
of the town.
Ber. I don’t see why. It’s taken a pile of courage to come out and
say you were wrong and make it right. If your neighbors are good
friends they will stand by you. They won’t go back on you.
Obad. That’s right, Abner. Of course we won’t.
Lem. And there ain’t a mite of reason why any one should know
anything about this. I for one shan’t mention it to Hepsy, and (grimly)
I guess if she don’t know it you’re safe.
Cyn. (earnestly). And I ain’t got the least bit of feeling against you,
Cap’n Abner.
Abner. Thank you, Cynthy. Thank you all. I don’t deserve such
treatment from you.
Lee. Say, Miss Tinker, just because you are suddenly coming into
money I hope you won’t be so set up that you’ll bounce your
boarders.
Cyn. Well, I guess it won’t be such an awful lot o’ money that I
need to be set up too high. I said you could stay as long as you
wanted to and I meant it. You can.
Lee. Good news!
Peter. Rah, rah, rah! Thank heaven we don’t lose those muffins!
Cyn. (suddenly). And I declare to goodness, Mr. Gordon, here’s a
special delivery letter that come for you. (Gives it to him. He tears it
open.) I thought it must be important and I come out to look for you
and so much has happened that it went clean out of my head.
 Lee. Well, say, this is certainly the day for unusual things to
happen. Look at this, Pete! (Hands the letter to Peter.)
Peter (glancing it over). What now? Well, glory be! Listen, you
who heard Mr. Williams denounce Lee this morning. The fellow who
was guilty in the Yale mix up has been found out and they want Lee
to go back to college. His father has also found out his mistake and
he wants Lee to go back to work for him.
Ariel (going to Lee). Oh, Lee, I’m so glad!
Lee. Well, they are just too late. Lee is going to stay right here on
his present job until he makes good.
Lem. Wal, I guess we got everything cleared up; we might as well
be moving along.
Obad. Gosh all fog horns, yes! I been so excited I ain’t had hardly
a bite to eat to-day. I’m faintin’ dead away if any one should ask you.
(Ber. walks to door, c., and stands looking out. Cyn., Lee, Peter,
Obad. and Lem. exeunt. Abner starts to follow, then looks back
at Ber. uncertainly. Ariel goes to Abner and throws her arms
around him. Ber. watches them from the door.)
Ariel. Oh, dad, I can’t tell you how happy I am.
Abner. I’m glad to hear you say it, Arey. I come mighty near
asking you to sacrifice your happiness.
Ariel (joyfully) But you didn’t quite do it after all.
Abner. Not quite.
Ariel. Father dear, don’t think anything you said can make a bit of
difference. I only love you more than ever before.
Abner. Lord, Arey, what can I say to a thing like that?
Ber. (coming forward). Say you are a fortunate man to have such
a daughter.
Abner. Yes, that’s what I ought to say. It’s true. I’m afraid I don’t
realize how true. (Pats Ariel on the shoulder.) Run along, little girl,
and be happy. (Ariel exits.) Captain Cranberry, I——
 Ber. If you please, Abner, I don’t believe I can talk any more just
now. If you’d just leave me up here alone for a while. It’s all right that
you should have her. It’s you that’s been the real father to her. You
have had the privilege that I have missed. You—you see she’s
Alicia’s little girl and I’ve jest got to get over it, that’s all.
Abner (grasping his hand). All right. I won’t talk. Some day—later,
I’ll try to tell you all that I feel.
[Exit.
(Ber. stands in the center of the room with bowed head.)
Ber. Alicia’s little girl! (Slight pause.)
Cyn. (outside). Arey! Arey! (Enters.) Land, Cap’n, I thought Arey
was here. Are you here all alone?
Ber. (slowly). Yes, alone, Cynthy. The way I’ve lived the most of
my life. The way I’ll always have to live it.
Cyn. My land, Cap’n, you must be dreadful down and out to talk
that way.
Ber. I ain’t a mite o’ good to any one on the face of the earth.
Cyn. You? You ain’t? Well, I’d like to see the man, woman or boy
in Bay Point that you ain’t some good to. Cap’n Berry, I didn’t
suppose you ever got to feeling like this. I think you must have lost
sight of the Beacon.
Ber. (suddenly looking up). By tunket, Miss Cynthy, you’re right!
That’s jest what I’ve done! I reckon I’m kind of tuckered out. I was
jest naturally making a fool of myself, thinking there weren’t nobody
on earth that loves me, and by Crismus, why should there be? I ain’t
got relations same’s other men and I ain’t got no right to expect the
same kind of happiness as other men. Well, I’ll set sail for the Point
and go on duty. That’s the thing for me to do. I’ve been spending too
much time over here and I need to go to work.
Cyn. Cap’n Berry, I should think you would be ashamed to talk so.
Nobody loves you! Why, everybody in Bay Point loves you, and you
know it.
Ber. (suddenly and bluntly). Do you, Miss Cynthy?
 Cyn. (very much confused and upset). Why, my gracious, Cap’n!
What a way to put it! How you talk!
Ber. (looking at her curiously). I’m a regular old fool, Cynthy. I’ve
had this on my mind for a long time and now, by tunket, I’m going to
get it off and then I’ll stop mooning around like a sixteen-year-old kid!
The first day I met you I loved you and I have been loving you a little
more every time I have seen you since. I wouldn’t want you to marry
unless it meant the same to you as it does to me, and I can’t believe
that’s possible. I reckon I know what you’re thinking. I reckon I know
what your answer will be, but I might as well have it from you
straight. (Goes to her, and puts his hands on her shoulders.) Cynthy,
do you think it would be possible for you to find happiness with a
frost-bitten old Cranberry?
Cyn. (looking up at him). I think it would, Cran, if you were the
Berry.
CURTAIN
 MUCH ADO ABOUT BETTY
A Comedy in Three Acts
By Walter Ben Hare
Ten male, twelve female characters, or seven males and seven
females by doubling. Costumes, modern; scenery, two easy interiors.
Plays a full evening. Betty, a moving picture star, going south on a
vacation, loses her memory from the shock of a railway accident,
and is identified as a rival, Violet Ostrich, from a hand-bag that she
carries. In this character she encounters the real Violet, who has just
eloped with Ned O’Hare, and mixes things up sadly both for herself
and the young couple. An exceptionally bright, clever and effective
play that can be highly recommended. Good Negro, Irish and
eccentric comedy parts.
Price, 35 cents
CHARACTERS
Lin Leonard, Betty’s one best bet.
Major Jartree, of Wichita, not only bent, but crooked.
Ned O’Hare, a jolly young honeymooner.
Mr. E. Z. Ostrich, who has written a wonderful picture-play.
Dr. McNutt, solid ivory from the neck up.
Jim Wiles, a high-school senior.
Archie, a black bell-boy at the Hotel Poinsettia.
Officer Riley, who always does his duty.
Officer Dugan, from the Emerald Isle.
Mr. Ebenezer O’Hare, a sick man and a submerged tenth.
Mrs. Ebenezer O’Hare, “Birdie,” the other nine-tenths.
Aunt Winnie, Betty’s chaperone.
Lizzie Monahan, Betty’s maid, with a vivid imagination.
Ethel Kohler, a high-school admirer of Betty.
Violet Ostrich, a film favorite, Ned’s bride.
Mrs. K. M. Diggins, a guest at the Hotel Poinsettia.
Daffodil Diggins, her daughter, “Yes, Mamma!”
 Miss Chizzle, one of the North Georgia Chizzles.
Pearlie Brown, Violet’s maid, a widow of ebon hue.
Violet, Violet Ostrich’s little girl aged seven.
Diamond, Pearlie’s little girl aged six
and
Betty, the star of the Movagraph Co.
Jartree may double Dugan; Ned may double Riley; Jim may double Archie; Mrs.
O’Hare may double Ethel; Aunt Winnie may double Pearlie and Lizzie may double
Miss Chizzle, thus reducing the cast to seven males and seven females. The two
children have no lines to speak.
SYNOPSIS
Act I. Betty’s apartments near New York. Married in haste.
Act II. Parlor D of the Hotel Poinsettia, Palm Beach, Fla. Three days later. Betty
loses her memory.
Act III. Same scene as Act II. A full honeymoon.

JUST A LITTLE MISTAKE

A Comedy in One Act
By Elizabeth Gale
One male, five female characters, or can be played by all girls.
Costumes, modern; scenery, an easy interior. Plays forty minutes.
Mrs. Ball receives a cablegram from her sister Lucy stating that Jerry
will arrive that day and begging her to be cordial. Mrs. Ball then goes
out to hire a cook, leaving three young friends to receive the
unknown guest. The cook, sent down from the agency in haste, is
greeted and entertained as Jerry and when the real Jerry (Miss
Geraldine Take) arrives she is sent out to the kitchen. After
considerable confusion and excitement she is discovered to be the
“Little Miss Take.” Strongly recommended.
Price, 25 cents
 THE SUBMARINE SHELL
A War Play in Four Acts
By Mansfield Scott
Seven males, four females. Costumes, modern; scenery, two
interiors. Plays two hours. Royalty for amateur performance $10.00
for one, $15.00 for two performances. Inspector Malcome Steele, of
the U. S. Secret Service, devotes himself in this thrilling play to
unravelling the German plots that surround Prof. Middlebrook’s
submarine shell that is to bring the downfall of the Hun. The battle
between his wits and those of “Tom Cloff,” the German secret agent,
is of absorbing interest. An easy and effective thriller that can be
recommended for school performance.
Price, 35 cents
CHARACTERS
Hans Kraft, alias James McGrady.
Otto Herman, alias William.
Mr. Warren Middlebrook.
Monsieur Charles LeClair.
Professor Henry Westerberg.
Detective Albert Bradbury.
Inspector Malcome Steele.
“Tom Cloff.”
Mrs. Middlebrook.
Eleanor Middlebrook.
Margaret Linden.
Delia.
SYNOPSIS
Act I. The living-room. August 11, after dinner.
Act II. Same as Act I. August 12, 1:30 p. m.
Act III. The private laboratory. That evening, 7:30.
Act IV. Same as Act III. Later, 10 p. m.