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THE WHEAT PLANT

BY THE SAME AUTHOR.

AGRICULTURAL BOTANY: Theoretical
and Practical. By JOHN PERCIVAL, M.A., F.L.S.
With 265 Illustrations. Sixth Edition, Revised
1921, Large Crown 8vo. i8s. net.

AGRICULTURAL BACTERIOLOGY:

Theoretical and Practical. By JOHN PERCIVAL,
M.A., F.L.S. With 59 Illustrations. Large
Crown 8vo. 125. 6d. net.

THE WHEAT PLANT

A MONOGRAPH

BY

JOHN PERCIVAL

J n

M.A., F.L.S.

PROFESSOR OF AGRICULTURAL BOTANY AT UNIVERSITY COLLEGE, READING, ENGLAND
AUTHOR OF "AGRICULTURAL BOTANY," "AGRICULTURAL BACTERIOLOGY," ETC.

WITH 228 ILLUSTRATIONS

LONDON
DUCKWORTH AND CO.

3 HENRIETTA STREET, COVENT GARDEN, W.C.

First published in 1921.
(^4// rights reserved')

MAiN L1BRARY-AGRICC

TO MY

FATHER AND MOTHER

475719

PREFACE

MORE than twenty years ago, when first engaged in the teaching of botany
to agricultural students, I became aware of the botanist's neglect of culti-
vated plants, and in the preparation of my text-book of Agricultural
Botany found it necessary to make a first-hand study of all the common
crops before any living interest could be created in the work I had under-
taken.

I then determined upon a more intensive study of the chief farm plants
with a view of writing a series of botanical monographs upon them as
opportunity permitted.

Administrative duties and teaching have prevented me from carrying
out the project as designed. Nevertheless, I have kept alive the intention,
and the present monograph is an embodiment of researches made upon
the wheat plant.

I hope that it may serve in a modest way as a model of the research
that is needed upon all farm plants, and act as a stimulus to further effort.

The plants with which the farmer has to deal provide limitless fields
of exploration for the plant physiologist, anatomist, student of variation
and heredity, and indeed all ranks of botanists. Their study will satisfy
the most ardent aspirations of the academic mind in its pursuit of so-called
pure science, and the discoveries to be made cannot fail to be of service
in enabling the cultivator to obtain higher economic returns from the soil
on which all animal life ultimately depends.

In Part I. of the monograph are given the results of my investigations
of the morphology, anatomy, growth, and development of the wheat
plant. Except where otherwise indicated, the various measurements
and illustrations refer to the ordinary dense-eared type of Bread wheat
(T. vulgar e) commonly grown in Great Britain.

In Part II. are described and discussed the taxonomy, characters, and
relationships of the wild species of wheat and the races and varieties of

viii THE WHEAT PLANT

cultivated forms. This section of the work is based upon a prolonged
study of the living plants, forming what may be termed the Reading
Collection. The latter, which is grown annually at the University College
Farm, is probably the most representative collection in existence, and
includes all the races of wheats numbering nearly two thousand forms
derived from almost all wheat-growing regions of the world.

For assistance in obtaining these I am especially indebted to the
Ministry of Agriculture and Fisheries and the Foreign Office ; to Dr.
Regel and M. C. Flaksberger of Petrograd and Dr. Vavilov of Moscow
for Russian and Central Asiatic wheats ; Professor Kozai, Tokio, for
Japanese wheats ; H. M. Leake, Esq., Economic Botanist to the Govern-
ment of the United Provinces, and D. Milne, Esq., Economic Botanist,
Punjab, for Indian wheats ; and numerous workers in other countries.

I desire also to express my thanks to the Council of the College for the
provision of facilities for research ; to Miss Mason and Mr. F/O. Mosley
for assistance with experimental work ; and to Professor Pennington for
his enthusiastic interest in the wheat collection and the cultivation and
testing of selected forms.

To Miss Erith, Lecturer in Agricultural Botany, I owe a debt of
gratitude for faithful assistance with records and experimental investiga-
tions during the last three years, and for the onerous task of proof-reading.

All the illustrations are original, those of the ears of the various races
of wheat being natural size.

JOHN PERCIVAL.

UNIVERSITY COLLEGE, READING.

The Wheat Plant

ERRATA

Page i 5, line 1 9, after " in " insert " some forms of," and for hermonis

read dicoccoides.

„ 15, line 24, for "Fig. 16" read "Fig. 15."
,, 50, in the tables for amyleuin read dicocciim.
,, 52, line 4, after "are" insert "sometimes."
„ 85, in the table for T. hermonis read T. dicoccoides.
„ 154, last line, for "usually 4-nerved " read " 2 to 4-nerved."
,, 167, line 34, for hermonis read dicoccoides.
,, 179, line 19, for " They" read " In some forms they."
,, 1 88, fifth line from bottom, for "four" read "two."
,, 192, line 2, for "4- (3-6)" read "2-6."
„ 196, line 24, for "3" read "2."
,, 400, line 30, after "secured" insert "in cross/."

his enthusiastic interest in the wheat collection and the cultivation and
testing of selected forms.

To Miss Erith, Lecturer in Agricultural Botany, I owe a debt of
gratitude for faithful assistance with records and experimental investiga-
tions during the last three years, and for the onerous task of proof-reading.

All the illustrations are original, those of the ears of the various races
of wheat being natural size.

JOHN PERCIVAL.

UNIVERSITY COLLEGE, READING.

CONTENTS

PART I

CHAP. PAGE

I. INTRODUCTORY ....... 3

II. THE GRAIN: STRUCTURE, COLOUR, AND OTHER CHARACTERS . 7

The Pericarp, 7 : The Seed, 10 : Colour of the Grain, 17 :" Hard "
" Flinty " and " Soft " " Mealy " Grains, 18.

III. GERMINATION OF THE GRAIN ..... 24

IV. THE ROOTS . . . . . . -35

General Morphology, 35 : Anatomy of the Roots, 39.

V. THE LEAVES ....... 44

General Morphology, 44 : Anatomy of the Leaves, 5 1

VI. THE STEMS: " TILLERING," " SHOOTING," AND " LODGING " . 62

General Morphology, 62 : " Tillering," 64 : " Lodging," 78 :
" Shooting" of the Stems and Escape of the Ear, 81 : Anatomy of the
Culm and Rhizome, 92.

VII. THE INFLORESCENCE ...... 100

Structure of the Rachis, Glumes, and Awns, 105.

VIII. THE FLOWER : FERTILISATION AND DEVELOPMENT OF THE

GRAIN . . . . . . . 110

Development of the Spikelet and Flower, 114 : Anthesis and Pollina-
tion, 122 : Fertilisation and Development of the Grain, 129.

PART II

IX. CLASSIFICATION . . . . . . .147

X. WILD SMALL SPELT (T. aegilopoides, Bal.) . . .164

XI. SMALL SPELT (T. monccoccum, L.) . . . .170

XII. WILD EMMER (T. dicoccoides, Korn.) . . . .178

XIII. EMMER (T. dicoccum, Schiibler) . . . . .186

x THE WHEAT PLANT

CHAP. PAGE

XIV. KHORASAN WHEAT (T. orientale, mihi) . . . 204

XV. MACARONI WHEAT (T. durum, Desf.) .... 206

XVI. POLISH WHEAT ( T polonicum, L.) . . . . 230

XVII. RIVET OR CONE WHEAT (T. turgidum, L.) . . . 241

XVIII. EGYPTIAN CONE WHEAT ( T. pyramidale, mihi) . . .262

XIX. COMMON BREAD WHEAT (T. vulgare, Host) . . . 265

XX. CLUB, DWARF, CLUSTER, OR HEDGEHOG WHEAT (T. compactum,

Host) ....... 307

XXI. INDIAN DWARF WHEAT (T. sphaerococcum, mihi) . . 321

XXII. SPELT OR DINKEL (T. Spelta, L.) . . . .325

XXIII. THE ORIGIN AND RELATIONSHIPS OF THE RACES OF WHEAT 335

XXIV. VARIATION . . . . . . -347

XXV. HYBRIDISATION AND WHEAT HYBRIDS .... 360

XXVI. THE IMPROVEMENT AND BREEDING OF WHEAT . . 407

XXVII. YIELD . . . . . . . .415

LIST OF WHEATS ...... 433

BIBLIOGRAPHY . . . . . . 441

INDEX . . . . . . .455

PART I

CHAPTER I

INTRODUCTORY

AMONG the world's crops wheat is pre-eminent both in regard to its
antiquity and its importance as a food of mankind.

In prehistoric times it was cultivated throughout Europe, and was
one of the most valuable cereals of ancient Persia, Greece, and Egypt.

Although rice is the principal food of a large proportion of the human
race, a greater amount of wheat is grown, and this, in the form of bread,
constitutes the chief food of the most highly civilised races.

In Russia and parts of Northern and Central Europe rye is the staple
food, as it was until comparatively recent times in Great Britain and
Western Europe generally ; but with improvements in the standard of
living, wheat is substituted for rye in the diet of the people, and its cultiva-
tion and consumption are being continually extended.

On account of the peculiar physical and chemical qualities of the
gluten of its grain, wheat makes more palatable and better bread than
any other cereal.

Its cultivation is simple, and its adaptability to varying soils and
climatic conditions superior to that of any other plant.

Originating from two or three wild species, through hybridisation,
mutation, and the effects of selection and cultivation, the races of wheat
have become as complex in constitution as the human race, and among
the almost endless number of varieties and forms which exist, wheats are
to be found suitable to the needs of agriculturists in all parts of the world.

Wheat is grown in every country in Europe and Asia, with the exception
of Siam, and large areas are devoted to it in Australia and New Zealand.

In Africa it is an important crop in Abyssinia, the Union of South
Africa, and along the Mediterranean coast from Egypt to Morocco.
Within the Tropics it is also grown in British East Africa and Nigeria.

In the Western Hemisphere the vast wheat-fields of Canada and the
United States are well known, and it is grown in Mexico and most of the
Central American Republics. It is also cultivated in nearly all the countries
of South America.

4 THE WHEAT PLANT

: 5 ,TIte> only parts of the world from which it is absent are the hot, low-
lying regions of the Tropics.

The greatest wheat-growing countries are Russia, the United States,
India, France, Canada, Italy, and Argentina. With the exception of
France and Italy, all these produce more than they require, and export
the surplus to regions which do not grow enough for their needs. Larger
or smaller imports enter every country in Western Europe, by far the
largest amount being taken by the British Isles.

The crop is being harvested in one country or another all the year round
as indicated below :

January. — Australia, New Zealand, Argentina, Chili.

February . — India .

March. — India, Upper Egypt.

April. — India, Persia, Asia Minor, Lower Egypt, Mexico, Cuba.

May. — Japan, China, Central Asia, Morocco, Algeria, Tunis, Texas.

June. — South France, Spain, Italy, Greece, Turkey, Japan, United
States south of 40°.

July. — France, Germany, Austria, Hungary, Roumania, Bulgaria,
Southern Russia, Canada, Northern United States.

August. — England, Northern France, Belgium, Holland, Central
Russia, Canada, United States.

September. — Scotland, Sweden, Norway, Canada.

October. — Northern Russia, Finland.

November. — South Africa, Argentina, Peru.

December. — Bdrma, Australia, Argentina.

The great wheat-producing areas of the world are found in the tem-
perate regions between the parallels of latitude 3O°-6o° N. and 27°-4O° S.

Wheat can, however, be grown from beyond the Arctic circle to the
Equator.

In Europe it has yielded ripe grain as far north as 69° 28' N. at Skibotten
on the Lyngenfjord in Western Norway, and in European Russia it is
cultivated around Archangel in latitude 64° 33' N. The northern limit
of growth in Great Britain is the Orkney Isles about 59° N., though its
profitable culture does not extend beyond 51° or 52° N.

In Asiatic Russia Flaksberger reports the growth of spring wheat
at Verkhoyansk in Siberia, within the Arctic circle.

In the Western Hemisphere spring wheats mature in Alaska up to
60° N. In the Peace River valley in Canada a flour-mill is maintained
at Fort Vermilion, and ripe grain has been produced up to 65° N. on the
Mackenzie River.

It is grown at the Equator on the high lands of Ecuador and Colombia,
and its cultivation is carried on in British East Africa and Nigeria.

Other countries within the Tropics in which the cereal is grown are

INTRODUCTORY 5

India, Arabia, the Philippine Islands, Peru, Brazil, Venezuela, Salvador,
and other Central American Republics.

Wheat has also a wide altitudinal range. In this country it is grown
from sea-level up to about 600 feet.

Farther south, in the Swiss Alps and the Pyrenees, spring wheats
can be grown at an elevation of 4000 feet, and in the Tropics their cultiva-
tion is carried on at an elevation of 8000-10,000 feet in Mexico > Colombia,
Ecuador, and Abyssinia.

Humboldt records its growth at an altitude of 14,000-15,000 feet in
Tibet.

The greatest amount of the best wheats are produced in countries
where the winters are cold and the summers comparatively Jiot. So long
as the winter temperature does not fall below — 20° C.'^and the air and
soil are dry, the exposed plants suffer little, but at lower temperatures
the crop is damaged, and in some cases killed outright, unless protected
by a covering of snow.

In regions of severe winters with cold winds and little snow only
spring wheats are grown, the autumn-sown winter wheats being reserved
for districts with a higher mean winter temperature, or where a sheltering
layer of snow is certain.

For the most satisfactory growth and development of the grain a cool,
moist, growing season followed by a bright, dry, and warm ripening period
of 6-8 weeks, with a mean temperature of 18-19° C. (about 66° F.), is
necessary.

In those parts of this country where the mean temperature of June,
July, and August is below 13-14° C. (56° or 57° F.) wheat is a hazardous
crop.

In regard to the water-supply essential for the wheat crop, an annual
rainfall of 20-30 inches is sufficient, so long as the greater proportion of
it falls during the growing season. Some of the Macaroni and Club
wheats are capable of yielding remunerative returns in regions where the
rainfall is not more than 12-15 inches per annum, and there are records
of crops of Macaroni wheats grown in arid districts without any rainfall
between sowing and harvest, the necessary water being derived from the
rain stored in the soil from the previous season.

Wheat is grown chiefly for its grain, which is ground and utilised in
the form of flour for the manufacture of bread and biscuits.

Some varieties are employed in considerable amounts in the prepara-
tion of macaroni and similar pastes, and most of the " cracked,"
" shredded," and " malted " " cereal foods " consist of specially treated
wheat. Occasionally the grains are cooked whole, and consumed with
milk, or used in soups in place of pearl barley.

Although the grain is useful food for farm animals, its production is

6 THE WHEAT PLANT

too costly in comparison with the cereals maize, oats, and barley to allow
of its being used for this purpose on a large scale.

The so-called " milling offals," however, consisting of the bran or
husk and other portions of the grain, obtained as by-products in the
manufacture of flour for bread-making, are among the most valuable
foods for all kinds of stock.

In the great wheat-growing regions the straw of the crop is of little
value, and is burnt or ploughed in, its humus and mineral constituents
serving to improve the physical and chemical condition of the soil. On
more intensively cultivated lands the straw is of greater importance to the
farmer, being employed as thatch for stacks, food for stock, or as litter
for the bedding of farm animals, in which latter case it is ultimately
returned to the soil as manure.

In addition to its agricultural value, wheat straw is utilised in the
manufacture of mattresses, straw hats, and paper.

CHAPTER II

THE GRAIN : STRUCTURE, COLOUR, AND OTHER CHARACTERS

A GRAIN of wheat, often spoken of as a " seed " by farmers, is a nut-like
fruit termed by botanists a caryopsis, a name first used by Richard.

It contains a single seed or kernel enclosed within a thin shell ; the
seed, however, instead of being free as in many nuts, is adherent to the
inner wall of the pericarp or shell, and the two cannot be separated readily.

As indicated later, the grains of different kinds of wheat vary con-
siderably in size, form, and colour : they nevertheless all resemble each
other in fundamental structure, being fruits with thin- walled pericarps,
each containing a single seed, the latter consisting of four_j)arts, viz- •
(i) the seed-coat or testa, sometimes termed the spermoderm ; (2) the
embryo or young wheat plant ; (3) the nucellar layer ; and (4) the endo-
sperm or floury part — a thin-walled parenchymatous tissue stored with
food for the nutrition of the embryo when germination commences.

From careful dissection of several grains and weighing of the separate
portions the embryo was found to weigh from 2-8 to 3-5 per cent of the
grain ; the pericarp, seed-coat, nucellar layer, and aleuron layer from 7-8
to 8-6 per cent ; the rest, or some 87 to 89 per cent, being the endosperm-
parenchyma, containing starch and gluten.

Ordinary white flour consists chiefly of the finely ground endosperm,
the so-called " milling offals " consisting of the broken pieces of pericarp,
seed- coat, aleuron layer, and the embryo, the latter being usually separated
from the other portion of the grain.

The Pericarp. — The dorsal surface of the pericarp of a grain of wheat
is smooth and rounded, the opposite or ventral side having a characteristic
inward fold or furrow called the " crease " by millers.

At the base is a small wrinkled patch — the part of the pericarp which
covers the embryo plant within. The apex is clothed with a number of
short stiff hairs, which curve inwards towards a common point, and form
the " brush " of the grain. The surface covered by the " brush " slopes
slightly to the furrow side, and is somewhat triangular in outline ; among
its hairs the remains of the styles are found.

7

THE WHEAT PLANT

In most parts the pericarp of the ripe grain is 45-50 p, thick and com-
posed of four or five layers of cells, namely, an outer and inner epidermis
with two or three layers of parenchyma between.

The outer epidermis is formed chiefly of elongated cells from 125 to
210 IJL long and 25-30 /z broad ; the cell walls, which are about 5-7 ft thick,
are perforated with pores, and in surface view appear beaded (Fig. i).
The cells are arranged lengthwise from base to apex of the grain ; near
the apex they become shorter, some of them being almost square.

Belonging to the epidermis are the hairs of the " brush " ; these are
single, fine-pointed cells -5 mm. to i mm. long, each with a more or less

FlG. i. — a, Cells of the outer
epidermis of caryopsis( x 260) ;
b, wall of cell showing thicken-
ing ( x 385).

FIG. 2. — Hairsof the"brush"
at the apex of the caryopsis
(xios).

bulbous base ; their cell cavities are of smaller diameter than the cell
walls, which have a thickness of about 6 ju, (Fig. 2).

Beneath the epidermis are one or two layers of thick- walled parenchyma
and an irregular layer of similar tissue, the cells of which have thin walls,
and are crushed and often disorganised.

Succeeding these is a well-defined layer consisting of elongated cells,
the largest diameters of which are arranged at right angles to those of the
epidermal cells, and therefore across the grain. The cells of this " cross
layer " in the young grain contain chlorophyll, but in the ripe grain are
empty like the rest of the pericarp cells. Except at the apex and base of
the grain, where they are shorter, they are usually from 100 to 150 /j, long
and 15-20^1, broad, with walls 5-7^1 thick, in which are irregular slits or
pores.

In adapting themselves to the shrinking grain when ripening takes

THE GRAIN

place, these cells become more or less crumpled or bent, often leaving
small intercellular spaces where the ends meet (a, Figs. 3, 4, and 5).

Immediately within the " cross layer " is the inner epidermis of the
pericarp. It consists of
thinner - walled vermi-
form or hypha-like cells,
termed " tube-cells " by
Vogl, from 1 20 to 250 /z
long and 12 to 15 p wide;
they lie parallel to the
outer epidermal cells
running from base to
apex, and in a ripe grain

are either separated from FIG. 3. — Transverse section of the pericarp and aleuron

layer of a ripe grain ( x 210). x, Epidermis ; a, " cross

g

layer"; b, "tube-cells" (inner epidermis); d, testa;
e, crushed nucellar layer ; g, aleuron layer.

each other altogether by
wide spaces or are joined
only at irregular intervals.

In transverse or longitudinal sections of the ripe grain these " tube-
cells " are difficult to observe, since they are isolated and do not form a
continuous layer, and the individual cells
are crushed into close contact with the
testa beneath ; their outlines, however, are
easily seen when surface views of strips
of the soaked pericarp are examined after
the outer epidermis has been removed (b,
Figs. 3 and 5).

FIG. 4. — Surface view of the cells (a) of the " cross FIG. 5. — a, Cells of the "cross layer "
layer " of the pericarp, with thin-walled cells of of the pericarp ( x 210) ; b, tube-
trie testa crossing them ( x 210). cells of the pericarp.

The pericarp in the central region of the furrow is thicker than on
the opposite side, and in it are usually developed one or two vacant spaces,
where the tissue has been torn asunder during the ripening of the grain.

io THE WHEAT PLANT

From the point at the base of the grain where the latter was attached
to the rachilla, a delicate vascular bundle traverses the pericarp upwards
along the furrow to near the base of the style.

The Seed. — Within the pericarp is the seed, consisting of testa, nucellar
layer, endosperm, and embryo.

(a) Testa. — In a transverse section of a ripe grain of red wheat, the
testa or seed-coat, to which the colour of the grain is chiefly due, is seen
as a reddish-brown line immediately next to the " cross layer " (d, Figs.
3, 109), the " tube-cells " being usually invisible. It is derived from the
inner of the two ovular integuments, and consists of two layers of elongated
cells, the lumina of which have been obliterated by the crushing together
of the upper and lower walls ; the cell walls of the outer layer are almost
colourless, those of the inner one being tinted red or brown. On account
of their extreme tenuity the separate layers cannot be distinguished in
transverse sections of ripe grains, but after treatment of the pericarp with

a solution of caustic potash, surface
views of them may be obtained,
when they appear composed of thin-
walled parenchymatous cells, 100-
150 /z long and 15-20/^1 broad ; the
long axes of the cells of the outer
layer cut the " cross layer " almost
at right angles, the inner cells being
arranged across the latter at an acute

FIG. 6.-Surfaceviewof the twoVell-layers angle (Figs- 4 and 6).
of the testa ( x 210). i, The outer ; 2, the The seed-coat covers the embryo

and endosperm, and is folded in
the centre of the furrow, where it joins the funicle of the seed.

(b) The Nucellar Layer. — Next to the reddish-brown seed-coat is a
bright colourless line — the so-called " hyaline layer " — possessing no
apparent cell structure (e, Figs. 3, 109). It is the epidermis of the nucellus,
the inner and outer walls of which have been crushed together, so that
the lumen of the cells is not visible in transverse sections. The cells are
more or less rectangular and oblong ; their outline may be seen on examin-
ing portions of the soaked pericarp.

(c) The Endosperm.— With the exception of the small space occupied
by the embryo the endosperm tissue fills the interior of the grain. It
may be divided into two parts, viz. : (i) the so-called " aleuron layer "
and (2) the starch- and gluten-parenchyma.

The aleuron layer surrounds the rest of the endosperm, closely following
the contour of the seed-coat. It is almost entirely one cell j:hick, measuring
about 65-70 n across (g, Fig. 3) ; only in the furrow, and occasionally at
other parts, are two or more cells superposed. The individual cells are

THE GRAIN

ii

rectangular in transverse and longitudinal sections, and polygonal, with
rounded corners in surface view, varying in diameter from 25 to 75 fj,,
and having walls about 6 /z thick (Fig. 7). In front of the scutellum the
layer consists of smaller cells, while over the plumule and coleorhiza it
becomes so very thin and shrivelled that it is difficult to recognise it as
a separate layer. Formerly the cells were termed " gluten cells," but
Schenk in 1872 showed that they contain no gluten. In each is a round
or oval nucleus about 12 \i in diameter and possessing three or four nucleoli.
The rest of the cell cavity is filled with minute spherical bodies, usually
described as aleuron grains, im-
bedded in a small amount of waxy
or oily cytoplasm. Each grain
measures about 3 or 4 /x in dia-

FIG. 7. — Surface view of the aleuron
layer ( x 210).

FIG. 8. — Cell of the aleuron layer after
treatment with dilute alcohol, showing
nucleus and insoluble membranes of the
" aleuron grains " ( x 770).

meter, and consists of a highly refractive core surrounded by a thin
membrane. When treated with water, salt solution, or weak acids or
alkalies, the core dissolves, leaving the insoluble membrane as a hollow
ball (Fig. 8). O'Brien considers that the homogeneous core consists of a
soluble protein which, according to Groom, contains a small amount of
magnesium and calcium phosphates, but neither globoid nor crystalloid as
found in typical aleuron grains are present in these granules.

The starch- and gluten-parenchyma, forming by far the greater portion
of the wheat grain, consists of thin- walled polyhedral cells, with their
long axes arranged usually at right angles to the surrounding pericarp.
Adjoining the aleuron layer, they are comparatively small, becoming two
or three times as large in the central parts of the endosperm. In contact
with the back of the scutellum of the embryo the cells of the endosperm

12

THE WHEAT PLANT

parenchyma are crushed together, forming a narrow band 35-40 fju broad,
devoid of starch (r, Figs. 10 and u). The majority of the cells, however,
are filled with starch grains imbedded in a protoplasmic matrix, from

which gluten is derived on treatment
with water ; more or less disin-
tegrated nuclei may be recognised,
but no aleuron grains are present.
The starch grains in the cells
immediately within the aleuron
layer are small — 8 to 10 p in
diameter — and more uniform in
size than those found in the larger
inner cells of the endosperm. In the
latter two forms are seen, namely,
large lenticular grains of circular or

FIG. 9. — Front view of the embryo.
s, Scutellum ; p, plumule ; /, epiblast.

elliptical outline measuring from 28 to 40 ^
across, and much smaller grains almost
spherical, rarely more than 6 or 8 \i in dia-
meter. In each of the larger grains is a small
central hilum surrounded by faint rings.

(d) The Embryo. — The embryo of the
wheat plant lies on the dorsal side at the
base of the grain, one side of it in close
contact with the endosperm, the other being
covered only by the seed-coat and pericarp.

It is a highly differentiated body, pos-
sessing a short hypocotyl, at the apex of
which is the plumule or primary bud, com-
posed of rudimentary leaves surrounded
by the coleoptile or plumule-sheath, and at
its base, the root -system, completely en-
closed by the coleorhiza or root-sheath
(Figs. 9 and 10).

Attached to one side of the axis, and
forming the greater portion of the embryo
as it exists in the grain, is a fleshy shield-like
structure named by Gartner the scutellum ;
on the other side, opposite to the scutellum,
is a small, tongue -like organ termed the
epiblast by Richard (e, Fig. 10).

r

u

FIG. 10. — Longitudinal section of
the embryo ( x 25). p, Pericarp of
the grain ; o, aleuron layer ; r,
crushed empty cells of the endo-
sperm ; s, scutellum ; a, epithe-
lium of scutellum; v, "ventral"
scale of scutellum ; o', coleoptile ;
/, ist grain foliage leaf; e, epi-
blast ; h, primary root ; n, coleo-
rhiza ; d, bud in axil of coleoptile.

THE GRAIN

(i.) The scutellum resembles a broad elliptical shield, measuring
3 mm. by 2-5 mm. and about -3 mm. in thickness, with a thinner rim.
It is slightly curved, the convex surface fitting closely to the endosperm.
The upper free portion extends about 1-3 mm. beyond the point of
attachment to the axis, and partly surrounds the coleoptile, the latter
being sunk in a slight depression in the scutellum. The portion which
projects over the apex of the coleoptile is sometimes termed the coty-
ledonary sheath or ventral scale (v, Figs. 10, 107).

The epidermis of the back of the scutellum in contact with the endo-
sperm consists of elongated cylindrical cells 35-40^ long and 8-10 p
broad, the long axes of which are arranged at right angles to the curved

FIG. ii. — Transverse section of a
portion of the back of the scutellum
and adjacent endosperm (x2io).
«, Epithelium of the scutellum ; r,

crushed empty cells of the endo- FIG. 12. — Gland-like cavity from the scutellum of
sperm. T. dicoccum ( x 200).

surfaces of the scutellum (a, Figs. 10 and n). It is termed the columnar
epithelium or epithelial layer ; and in the embryos of T. dicoccum, T.
durum, and some other wheats, portions of it are deeply inverted into
the form of simple tubular glands, especially near the upper and lower
edges of the scutellum (Fig. 12). Its cells secrete diastase, the enzyme
which renders soluble the reserve starch stored in the endosperm tissue,
and it functions also as a haustorium through which all the dissolved
plastic materials of the endosperm are absorbed and transferred to the
embryo when germination occurs.

(ii.) The epiblast or lobule is a short, thin scale (e, Fig. 10) about 170 p
long, 550 p broad at the base, and about 82 /x thick ; it is attached to the
axis slightly above the point of insertion of the scutellum, and consists of
parenchyma only, with no vascular system.

14 THE WHEAT PLANT

(iii.) The coleoptile or plumule-sheath is a protective leaf, which com-
pletely encloses the plumule, except near the apex, where, on the side
opposite to the scutellum, there is a minute opening (Figs. 9 and 10),
through which the first green leaves of the plumule ultimately make their
way. It springs from the same side of the axis as the scutellum and
immediately above the point of insertion of the latter organ. In the
resting embryo it is short — about i to 1-25 mm. long — but as it pushes
its way upwards through the soil when the grain is sown, it lengthens,
and may grow to a length of i or 2 inches.

(iv.) The plumule within the coleoptile consists of two or three rudi-
mentary leaves surrounding the growing point. These have a divergence
of ^, the first being on the side of the axis opposite the scutellum. In
addition to the primary bud there is in the wheat embryo a secondary
lateral bud in the axil of the coleoptile on the side next to the scutellum
(d, Fig. 10).

(v.) The Root-system. — The embryo in a grain of Bread wheat possesses
a well-differentiated root-system, consisting of five rootlets, namely, a
primary radicle directed slightly forwards and two pairs of secondary
lateral rootlets, which arise from the short axis of the embryo in a plane
almost parallel to the face of the scutellum.

The roots of the lower older pair, which arise just above the point of
insertion of the epiblast, are not so large as the primary radicle ; those
forming the upper younger pair are very much smaller. Another single
rootlet often grows out in front above the epiblast between the lateral
pairs, but it is rarely differentiated in the resting embryo of wheat, although
it frequently appears after germination is well advanced (b, Figs. 18, 22).

Later still, a pair of opposite rootlets arise above the other pairs and
parallel to them, the seedling plant then possessing eight rootlets.

Each of the rootlets terminates in a growing point covered by a root-
cap, and the primary and two lowest pairs just mentioned are enclosed
within the coleorhiza or root-sheath, a parenchymatous tissue connected
with the hypocotyl and the lower half of the scutellum.

(vi.) The Vascular System. — The vascular system is clearly defined
in the resting embryo in the form of procambial strands, consisting of elon-
gated thin-walled conducting cells of polygonal section, each from 50 to
80 /A long and 6 to 8 ju. in diameter.

A broad strand, oval or elliptical, in transverse section (Figs. 13 and
14), curves out almost at right angles from the hypocotyl, and extends
upwards in the free half of the scutellum in a median plane, nearly to the
upper edge (Fig. 10). For a short distance it becomes broader in its
progress upwards, but finally divides and spreads out in a fan-like fashion,
giving off a number of fine branch strands, each five to ten cells thick.
Some seven or eight of these on the right and a similar number on the

THE GRAIN

left curve round and extend backwards in sinuous lines more or less
parallel to each other, a few on each side being continued almost to the
base of the scutellum (Fig. 13).

In the young state the cells of the strands are thin- walled, but after
germination and growth of the young
plant, xylem tracheids, with lignified
walls and annular or spiral thickening,
are differentiated, and, in addition, a
certain proportion of the parenchyma
of the scutellum exhibits similar,
though less extensive, thickening of
most of its cell walls, transverse as
well as longitudinal.

In the majority of wheats two
straight, unbranched bundles traverse
the coleoptile from base to apex, one
on each side, arranged in a plane
almost parallel to the face of the
scutellum ; in T. hermonis and Indo-
Abyssinian forms of T. dicoccum two
to four additional bundles are present in the outer half of the coleoptile
(Fig. 16).

FIG. 13. — Diagram of the bundles of the
scutellum.

FIG. 14. — Transverse section of portion of scutellum ( x 105). e, Epithelium; b, section
of large central bundle ; a, sections of small bundles (cf. Fig. 16).

The rudimentary first green foliage leaf of T. vulgare usually possesses
eleven bundles, the second leaf seven.

The arrangement and number of the bundles of the scutellum, coleop-
tile, and first and second foliage leaves are shown in Fig. 15.

Traced downwards, four bundles of the first foliage leaf (two from
the right and two from the left of the midrib, Nos. 2, 4, 8, and 10) and

i6

THE WHEAT PLANT

a

r

FIG. 15. — Transverse sections of the embryo
showing the arrangement and number of
the vascular bundles at different levels.
5, Central bundle of scutellum ; a, smaller
bundles of scutellum ; i-n, bundles of first
foliage leaf ; m, midrib of the same ; B, D, F,
bundles of second foliage leaf ; r , lateral roots .

four belonging to the second
foliage leaf disappear before the
point of insertion of the first
foliage leaf is reached.

Lower down, near the point of
insertion of the coleoptile, the
scutellum bundle divides, the
right and left halves joining the
bundles from the coleoptile just
outside the stele.

Still lower, the bundle 5 from
the first foliage leaf coalesces
with bundle B from the second
leaf, and similarly bundles 7 and

FIG. 1 6. — Transverse section of the
scutellum and plumule of Abyssinian
Emmer (T. dicoccuni). Coleoptile with
six vascular bundles.

F on the other side of the midrib
(6) unite with each other : these
all join the vascular elements of
the pair of lateral rootlets which
arise at this point.

The midrib bundle 6 appears
to pass down through the node
into the primary root without
combining with others, although
the anatomy at the node is
obscure.

In regard to the homology of
the various parts of the embryo
many different views have been
advanced. The scutellum is
usually considered a cotyledon,
although Regel, Hofmeister,

THE GRAIN 17

Sachs, and others look upon it as an outgrowth of the hypocotyl or
radicle.

The epiblast, according to Bruns and Celakowsky, is a reduced second
cotyledon ; others suggest that it is a scale-like trichome ; to Worsdell
it is a part of the cotyledon, and corresponds to the auricles at the base of
the blade of a green foliage leaf.

The coleoptile or plumule-sheath is regarded by Hofmeister and
Sachs as the true cotyledon, while Van Tieghem, Klebs, and Worsdell
describe it as the ligule of the chief cotyledon, the blade of which is the
scutellum ; to Bruns and others it represents a bladeless leaf, the first
of the plumule, the second being the first green foliage leaf of the plant.
The view which I think is most in agreement with the development and
structure of the embryo of wheat (see pp. 133-136) is that which regards
the scutellum, epiblast, coleoptile, and first green leaf as the first four
leaves of the plant. The alternate distichous disposition of these structures
on opposite orthostiches also supports this conclusion.

A comparison of the arrangement and relationship of the adventitious
roots to the buds and leaves at the " tillering " nodes of the older plant
suggests that the three pairs of opposite seminal rootlets of the young
wheat plant are associated with three leaves — scutellum, epiblast, and
coleoptile — the plane through them being at right angles to the median
plane of these reduced leaves, an arrangement similar to that of later
roots which spring from the older parts of the axis bearing the foliage
leaves (see pp. 36-38).

The rootlet which appears at right angles to the plane of the rest,
above the epiblast, suggests the suppression of a bud in the axil of the
latter, a corresponding opposite rootlet frequently developing from the
base of the axillary bud of the coleoptile.

Colour of the Grain. — One or two Abyssinian varieties of T. dicoccum
possess grains which are a rich purple tint when freshly ripened, the
colour being due to the presence of an anthocyan pigment located chiefly
in the chlorophyll or " cross " layer of the pericarp (p. 8).

All other wheats have grains which are usually classified as " white " 1^
or " red." The distinction is based on the character of the testa : in
" white " wheats the latter is colourless, while in " red " wheats an oily
or resinous material develops in the cell lumen and cell walls of its two
cell layers during ripening.

The particular shade exhibited by the grain is, however, dependent
not only upon the amount of colour in the testa, but upon the thickness,
tint, and transparency of the superposed pericarp, and the " mealy "
and " flinty " character of the endosperm.

The pericarp is translucent, and in grains ripened in a hot, dry season
is a pale creamy tint, like that of clean unweathered straw.

c

1 8 THE WHEAT PLANT

" White " wheats show a range of tint from an opaque creamy- white
to a translucent amber or yellowish shade ; the former, of which beautiful
examples are seen in Australian and some Asiatic wheats, possess " mealy "
endosperm, while grains of the amber tint have yellowish " flinty "
endosperm which is visible through the thin pericarp.

Among " red " wheats the colour passes by fine gradation from the
palest tint to a dark brick- red or brown.

The colour is most readily distinguished in grains with " mealy "
endosperm, for the latter provides an opaque chalk-like background,
against which the colour of the testa and pericarp are clearly seen.

In many Squarehead forms of wheat with opaque chalky endosperm
the grains are an orange tint, a pale red testa being modified by the
yellowish- white pericarp : others, especially certain Chinese and Japanese
forms with a highly coloured testa and " mealy " endosperm, have rich
brick-red grains.

Red grains with " flinty " endosperm are always darker than mealy
grains of the same variety.

Where the grain is shrivelled or the pericarp discoloured by exposure
to rain or the attack of fungi, its true tint cannot be determined with
certainty. There is, however, usually little difficulty in assigning clean,
well-grown samples of wheat to their respective " white " or " red "
classes, except in certain forms of T. durum with translucent flinty endo-
sperm : among the latter wheats, pale " red " grains and amber-tinted
examples belonging to the " white " class can only be separated by the
most careful examination in a favourable light on a dark background.

" HARD " " FLINTY " AND " SOFT " " MEALY " GRAINS.— The endo-
sperm of certain grains of wheat, when cut transversely, is found to be
dense and translucent, resembling horn or flint in colour and transparency ;
to such the terms " flinty," " glassy," " horny," and " steely " have been
applied. Flinty grains are comparatively hard, with a somewhat vitreous
fracture, breaking into angular fragments, and market samples largely
composed of such grains are often designated " hard " wheats.

So-called " soft " wheats have " mealy " or " starchy " grains, possess-
ing a very white, chalky endosperm ; the grains are opaque and com-
paratively soft, yielding a loose mass of floury meal when crushed.

Sometimes the endosperm of a single grain is mealy in one part and
flinty in another : in such semi-flinty grains the apical portion is always
flinty, the white opaque part being found especially on the dorsal side
near the embryo, the mealiness being most conspicuous about the upper
edge and back of the scutellum, and extending inwards across the endo-
sperm to the furrow.

In many instances the grains of an ear are either all flinty or all mealy,
but not infrequently some ears contain both forms ; in the latter cases

THE GRAIN 19

I have usually found that the small spikelets near the base and apex bear
a greater proportion of mealy grains than the larger spikelets from the
middle of the ear, and the lower heavy grains of each spikelet are usually
more flinty than the smaller upper ones.

The flinty grains of any pedigree line wheats are, on an average, not
only larger and heavier than those of the same sample with mealy endo-
sperm, but have a higher specific gravity than the mealy grains.

Investigations of two examples gave the following results :

Vol. of 100 Grains. Weight of 100 Grains,
c.c. gr.

Sp. Gr.

Flinty.

Mealy. Flinty. Mealy.

Flinty.

Mealy.

3'4

37

3'1 4'54 4-04

3-46 5-01 4-48

1-335

1*344

1-287
1-292

The flinty grains in these wheats are usually 7-10 per cent larger,
10-12 per cent heavier, and have a specific gravity 3-4 per cent higher
than that of mealy grains.

The white opaque appearance of the endosperm of mealy grains is due
to the presence of minute fissures, which develop between and within the
cells during the dessication which occurs at the time of ripening of the grain.

On examination of carefully prepared transverse sections x from grains
showing different degrees of mealiness, it is seen that interstices have
formed along the line of union of adjacent cells and around the starch
grains within the latter, and the contents of some of the cells have shrunk
more or less away from the surrounding cell wall ; these changes appear
first near the furrow, and spread radially outwards across the endosperm
towards the aleuron layer on the dorsal side, especially in the basal half
of the grain near the embryo.

Minute irregular cavities are seen also in the aleuron cells.

From investigations on mealy grains of barley, Brown and Escombe
concluded that the interstices are vacuous or only partially filled with air.

Such minute spaces are absent from flinty endosperm in which all
the cells of the tissue are completely filled with starch grains imbedded in
a protoplasmic matrix, the whole forming a dense coherent mass.

The production of flinty or mealy grains is a hereditary character of

1 The endosperm of uninjured flinty grains appears to be under a stress which tends
to rupture the cells and cell contents, and thin sections cut from flinty grains immedi-
ately become opaque and " mealy." To obtain sections sufficiently thin to be examined
with a high power, without disturbing the original physical condition of the endosperm,
thick dry sections may be ground down on a " carborundum " stone with the finger-tip.
Examine the sections in oil or Canada balsam.

20 THE WHEAT PLANT

particular races and forms of wheat. The grains of T. aegilopoides , T.
dicoccoides, T. monococcum, and T. durum are almost always flinty, while
those of T. turgidum and many forms of T. vulgare and T. compactum are
especially mealy.

Mealy grains are most commonly developed among those races and
forms whose individual spikelets bear the greatest number of grains, and
there is strong positive correlation between mealiness and high grain-
yielding capacity in the race of Bread wheats (T. vulgare). Varieties
possessing normally opaque grains are generally slow-growing wheats
with a long vegetative period and adapted for cultivation in humid climates
or on irrigated land. On the other hand, those giving a high proportion
j of flinty grains produce less, grow and ripen more rapidly, and are met
f with chiefly in regions having a comparatively dry continental climate.

While some wheats maintain their distinctive type of endosperm under
widely different circumstances, others can be made to yield flinty or
mealy grains by varying the nutrition and the external conditions of
growth of the plant.

The Macaroni wheats (T. durum) have the most typically flinty grains,
and in these the feature is so strongly inherent that it is rarely or never
changed by environment.

In the opposite class are the Rivet wheats (T. turgidum), whose grains
are generally opaque and mealy ; in these the character is also com-
paratively stable, and only with difficulty are some of them induced to
bear ripe, flinty grain.

The Bread wheats (T. vulgare) are very variable in regard to the
physical appearance of the endosperm. While some of them exhibit a
strong hereditary tendency to bear grains with flinty or mealy endosperm,
the character is much less rigidly established in this race, and in many
forms of it the production of hard, translucent, or soft, opaque grains is
determined by climate and soil.

Among wheats commonly cultivated in this country the flinty grains
of a sample when sown give rise to plants yielding either flinty or mealy
grains according to the season, the texture of the soil, the space allotted
to the plants, or their manurial treatment ; mealy grains behave in a
similar manner, the physical nature of the endosperm being in both cases
controlled by environment.

Wheats grown in cool districts with abundant rainfall or under irriga-
, tion have softer and more mealy grains than those cultivated in drier and
/ warmer regions.

Voelcker's investigations at Woburn show that heavy soils have a
tendency to produce flinty grains, while light sandy loams give a higher
proportion of mealy grains, these effects being independent of the flinty
or mealy nature of the grain sown.

THE GRAIN

21

In the following table are given the results obtained from pot cultures
at Woburn in 1900 :

Produce.

Grain sown.

Mealy Grain.

Flinty Grain.

i Mealy Grain.

Mealy grains in light soil —

per cent.

per cent.

per cent.

Pot i

IOO

Pot 2

477

6-9

45'4

Pots

84-9

*5'*

Flinty grains in light soil —

Pot i

95'2

4-8

. .

Pot 2

79'4

20-6

. .

Pot3 . • • . • •

647

16-3

19-0

Mealy grains in heavy soil .

IOO

. .

Flinty grains in heavy soil .

IOO

Plants allowed plenty of space always yield a higher proportion of
flinty grains than those which are closely crowded ; below is given the
percentage of mealy and flinty grains in a vulgar e wheat, grown at various
intervals in rows from 6 inches to 24 inches apart ; generally in samples
of the same wheat produced under ordinary field culture 90 per cent of
the grains are mealy or half -mealy.

"SWAN" WHEAT
PERCENTAGE OF FLINTY, HALF-MEALY, AND MEALY GRAINS

Area allotted to each Plant.

Inches.

Inches.

Inches.

Inches.

Inches.

Inches.

6xi.

6x3.

6x6.

12 X 6.

12 X 12.

24 x 24.

Mealy

per cent.

9'5

per cent.
IO-I

per cent.

6-4

per cent.

37

per cent.
3'2

per cent.
•2

Half-mealy

22-2

127

3'8

2-6

3'3

1-2

Flinty

68-3

77-2

89-8

937

93'5

98-6

Although the translucency or opacity of the endosperm is a physical
phenomenon it is correlated with the amount of nitrogen present, the
flinty grains being always richer in this element than the mealy grains of
the same sample.

Moreover, flintiness can be induced in any form of T. vulgar e by
application of large amounts of nitrate of soda or sulphate of ammonia.

22

THE WHEAT PLANT

At the University College Farm, Reading, these fertilisers were
applied to small plots of wheat for several years at the rate of i, 2, 4, 8,
and 1 6 cwts. per acre respectively ; comparison of the produce with that
from the unmanured plots showed that i cwt. per acre distinctly in-
creased the percentage of flinty grains in T. vulgare, and the higher amounts
led to the whole of the sample being flinty.

In the case of the soft floury Blue Cone wheat (T. turgidum) it was not
until an application of 4 cwt. or more per acre was given that the mealy
nature of the endosperm was modified, and only when 8 to 16 cwt. per
acre was used were all the grains flinty.

Similar graduated quantities of the ordinary phosphatic and potassic
fertilisers were tried, but none of these increased the flintiness of the
endosperm.

Forms which ordinarily produce mealy grains only do so under con-
ditions which allow of complete development and normal ripening, for
the grains of all wheats harvested in an immature state have flinty endo-
sperm. The results of harvesting two mealy-grained forms of T. vulgar e
at weekly intervals are given below ; they illustrate the progressive in-
crease in the percentage of soft opaque grains up to the time of perfect
ripeness.

" SQUAREHEAD'S MASTER " WHEAT

PERCENTAGE OF FLINTY AND MEALY GRAINS IN EARS CUT AT WEEKLY
INTERVALS UNTIL RIPE

19.14-

Mealy.

Flinty.

Average weight of 100
Grains. (Air-dry.)

gr-

July 4 ...

0

IOO

1-42

„ ii .

0

IOO

2-yo

„ 18 . . .

34

66

4-08

,t 25 • • •

54

46

4'95

Aug. i . .

89

ii

5-01

„ 8 . . .

89

ii

4-81

" SWAN " WHEAT

IQl6.

Mealy.

Flinty.

Average weight of 100
Grains. (Air-dry.)

July 7 ...

0

IOO

77

14 .

0

IOO

I"54

21 .

0

IOO

2-48

28 ...

37

63

3-46

Aug. 4 ...

77

23

4-67

ii .

96

4

4-67

18 .

96

4

4-5i

THE GRAIN 23

" STRONG " AND " WEAK " WHEATS. — The miller and baker usually
grade the Bread wheats and the flour obtained from them into " strong "
and " weak " varieties, the term " strong " being applied to those wheats
whose flour makes large uniformly porous loaves, the " weak " flour
producing smaller denser loaves.

Although there are many exceptions, the " strong " wheats generally
have red grains with hard " flinty " endosperm, the weaker types having
paler red or white grains with opaque " chalky " endosperm.

CHAPTER III

GERMINATION OF THE GRAIN

A NORMALLY ripened wheat grain, sown an inch or an inch and a half
/ deep in good soil, early in September, begins to germinate in two or three
days, the coleoptile and first leaf appearing above ground in about ten
days. The grain which at the time of ripening became comparatively
dry now absorbs water from the ground, and the dormant embryo soon
gives signs of growth. Concurrent with the visible morphological changes
in the embryo, changes occur in the reserve materials stored in the endo-
sperm, the solid proteins and carbohydrates of the latter being rendered
soluble and diffusible by the action of various enzymes.

Before germination can proceed a number of conditions must be
fulfilled, the chief of which are : (i) a suitable degree of moisture of the
seed-bed, (2) an adequate temperature, and (3) a sufficiency of oxygen.
A deficiency of water, heat, or fresh air retards or completely prevents
germination.

A grain which is on the point of germinating is swollen, its volume
compared with that of a dry caryopsis being considerably increased by
the water which it has absorbed. The puckered surface of the pericarp
covering the embryo becomes smoother, and the whole wall of the fruit
is tense.

Very soon the growing embryo bursts its covering near the base of
the grain, the first portion to emerge being the dilated coleorhiza, which
expands at first in a transverse direction, and tears a longitudinal slit in
the pericarp, thus exposing the plumule. Just after its escape it appears
as a short truncate structure, falsely suggesting the presence of two rootlets
within it (Fig. 17), and consists of elongated and distended parenchy-
matous cells between which are many intercellular spaces which give the
tissue a white glistening appearance. Later, a number of long unicellular
hairs, resembling root-hairs in form and function, often arise from the cells
of the coleorhiza, especially near its base.

After the coleorhiza has grown about a millimetre, the enclosed
primary root bores through it, usually on one side, leaving a bulging

24

GERMINATION OF THE GRAIN 25

angular portion opposite. An hour or two later the first pair of lateral
rootlets become visible, each with its thin covering root-sheath ; these
spring from the sides of the short hypocotyl almost in line with the base
of the epiblast (Fig. 18).

Under favourable conditions the three rootlets and plumule grow

FIG. 17. — Base of germinating grains, showing rupture of the pericarp by the coleorhiza
and plumule of the growing embryo.

rapidly, the former elongating much more quickly than the latter. The
primary root is for a time somewhat longer than the secondary ones, but
later there is little difference between them either in length or thickness.
Usually in three or four days, when the first rootlets have attained a length
of about 2 cm., two more appear, one on each side of the hypocotyl imme-

FIG. 1 8. — Young plants, p, Plumule ; c, coleorhiza ; i, primary root ; 2, root of first pair
of lateral roots ; 3, root of second pair; 4, root above the epiblast, inserted at right
angles to the others.

diately above the first pair, a sixth rootlet sometimes developing later,
at right angles to the rest, from a point on the axis behind the epiblast
(Fig. 18).

The germination of the naked caryopses of T. monococcum, T. dicoccum,
and T. Spelta is similar to that of other wheats : sprouting is, however,

26 THE WHEAT PLANT

generally more speedy in these grains on account of the thinner pericarp,
which permits of rapid absorption of water.

When whole spikelets of the " Spelt " wheats are sown, the coleorhiza
and rootlets of the embryo pierce the base of the grain and the enclosing
flowering glume, but are unable to break through the thicker empty
glumes, the entrance of the rootlet into the ground being made past the
edges of the latter at the base of the spikelet.

In these wheats, in which strong glumes closely invest the grain,
no longitudinal slit exposing the plumule is torn in the pericarp by the
expanding coleorhiza and rootlet.

In some cases the flattened wedge-like coleoptile breaks through the
pericarp near the upper edge of the scutellum and appears at the tip of
the spikelet, after growing along the space between the outer surface of
the caryopsis and the inner surface of the flowering glume : most fre-
quently, however, the plumule escapes from the apex of the grain after
pushing its way longitudinally within the substance of the pericarp in
which it travels through the thin parenchyma outside the " cross layer."

Placed in the soil or on damp filter paper, wheat caryopses soon
begin to absorb water, which penetrates most rapidly through the thin
portion of the pericarp covering the embryo. Laid furrow side down on
a wet surface, they imbibe more slowly than when the dorsal half is wetted.

In order to test the amount absorbed, 100 grains were completely
immersed in water. At certain intervals they were withdrawn, and after
the superfluous water had been carefully removed from their surfaces
by means of filter paper, were weighed again. They were then immersed
for a further period. The results obtained are shown in the table on
the following page.

Germination commenced after about 40 hours' immersion, up to which
time the amount of water absorbed by the grains steadily increased, the
flinty grains imbibing more slowly than the starchy ones.

From another series of investigations I found that grains do not
sprout until they have absorbed at least 30 per cent of their weight of water,
and, with the water content maintained at this figure, germination is
feeble and protracted.

The coleorhiza emerges in about 24 hours after the grains have taken
up 35 per cent of water, which they usually do after 12 to 14 hours' soaking
at 17-18° C. The most vigorous germination, however, at this tem-
perature takes place when the amount imbibed is 36-40 per cent of
the air-dry weight of the grains.

In addition to the increased weight due to the absorption of water,
there is always a change in the volume of the grain, the swollen caryopsis
ready to germinate being from 35 to 40 per cent larger than when
air-dry.

GERMINATION OF THE GRAIN

27

" SWAN " WHEAT
Temperature 16° C.

Weight.

Increase.

Percentage
Increase.

grs.

grs.

ioo starchy grains air-dry

4-70

. .

Weight after soaking 30 minutes

5'H

•44

9'4

„ „ i hour .

5'33

•63

*3'4

„ 3 hours .

5'44 .

74

157

55 55 O , . .

5^4

•94

2O'O

5, „ 9 , . .

5-84

1-14

24-2

55 55 28 , . .

6-45

37'2

36 5 • •

6-68

1-98

42-1

53 . •

6-77

2-07

44-0

ioo flinty grains air -dry

4-98

Weight after soaking 3 hours .

5-64

•66

13-2

55 „ 6 „

•83

16-6

55 55 9 »

5-98

i-oo

20- 1

55 55 25 ,,

6-49

1-51

30-3

5> 55 3^ »»

6-73

175

55 55 53 55 •

6-93

39-i

The relative volumes and weights of the grain when dry, and the same
after soaking 5 hours and 28 hours respectively, were found in one trial
to be :

After soaking.

Air- dry.

5 Hours.

28 Hours.

Relative volume

I

1-222

1-416

Relative weight

I

1-190

1-366

The increase in volume is somewhat greater than the addition in
weight, an augmentation of the latter of 19 and 36-6 per cent respec-
tively being attended by a rise in volume of 22-2 and 41-6 per cent.

The optimum temperature at which germination is most rapid lies
between 20° and 22° C., the minimum, below which it is completely
checked, is about 4° C. The influence of different temperature is seen in
the following results obtained with a Squarehead form of wheat harvested
in August and tested in November 1914.

28

THE WHEAT PLANT

PERCENTAGE GERMINATION

After

35° C.

22° C.

i6°C.

12° C.

80 /-.
\-s.

24 hours ,

33

63

10

0

0

48 , ...

51

32

67

0

0

72 , ...

9

5

22

99

14

96 , ...

4

37

120 ,

i

43

144 , . . .

2

6

IOO

IOO

99

99

IOO

At temperatures considerably above the optimum the grains
germinate irregularly, the sprouting of some of them being much delayed.
Moreover, the embryos frequently die or become so much weakened that
they are liable to be destroyed by the attacks of fungi and bacteria ; the
endosperm also often undergoes decomposition through the activity of
these organisms when the moist grains are kept at 35° C. or higher.

Although germination begins in two or three days or less in grain
sown in the open field at any period of the year, the progress made by the
growing embryo is very slow in late autumn and winter when the tem-
/perature is low, especially if at the same season the rainfall is excessive.
The time which elapses before the coleoptile appears above ground in
grains deposited at a depth of an inch may vary between one and ten
weeks.

Observations made upon weekly sowings of wheat during 1912, 1913,
and 1914 gave the following results, the grains being sown at a depth
of one inch :

NUMBER OF DAYS BETWEEN THE TIME OF SOWING AND THE APPEARANCE
OF THE YOUNG PLANT ABOVE GROUND IN WHEAT SOWN IN DIFFERENT
MONTHS OF THE YEAR.

Jan.

Feb.

Mar.

April.

May.

June.

July.

Aug.

Sept.

Oct.

Nov.

Dec.

1912
J9J3

IQIA

50
7<

16

-?!

9

23

22

6
IS

7
8
8

7
9

7

7
H

8
12

10
10

17
II

39

21

71

42

1 V^T-

jj

During germination extensive chemical changes occur both in the
embryo and the reserve materials of the endosperm.

Starch is absent from the tissues of the dormant embryo except in the
merest traces, but minute globules of a fatty oil and small spherical aleuron

GERMINATION OF THE GRAIN 29

grains — granules of protein, each possessing one or more excessively
minute globoids — are present in the parenchyma of the scutellum and other
parts of the embryo.

Before germination begins, the columnar epithelium of the scutellum
is devoid of starch, but its cells contain dense cytoplasm and an oval
nucleus, which usually rests in the lower half of the cell with its long axis
arranged lengthwise in the cell lumen.

In 12 to 1 6 hours after germination commences in grains sprouting
indoors at i8°-2o° C. starch granules make their appearance in the cells
of the coleorhiza, root-cap, and tip of the coleoptile ; these are derived
chiefly from fat globules or from a soluble carbohydrate stored in the
tissues of the embryo and not from the endosperm, since they arise in
portions of the embryo which have been cut from dry grains and laid upon
damp filter paper. Nevertheless, dissolved reserves soon begin to be
transferred from the endosperm to the growing embryo, and in 24 to 36
hours fine-grained starch is found in the epithelium of the scutellum,
and in greater abundance in the parenchyma of the latter. Increas-
ing quantities are seen also about this period in the fully developed
cells of the root-cap, in the coleoptile, especially near its apex, and in the
coleorhiza ; a small amount is present in the endodermis of the primary
root.

The grains are usually spherical, about i //, in diameter, and many
of them compound, consisting of two or three partial granules. Their
special accumulation in the root-cap and apical cells of the coleoptile
supports the view of Haberlandt and Nemec that they act as statoliths
in the geotropic curvatures of these parts.

The reserves stored in the endosperm are rendered soluble by various
enzymes. Among the latter, whose presence has been established in
germinating cereal grains, are cytase and the amyloclastic enzyme diastase ;
presumably, proteoclastic ferments capable of breaking down the reserve
proteins also occur in the grain, but these have not been investigated
in wheat.

The dissolved plastic materials are absorbed by the columnar
epithelium, and passed on through the parenchyma and the vascular
system of the scutellum to the growing regions of the young plant.

In the resting state of the embryo the cylindrical epithelial cells are
from 30 to 40 p long with somewhat rectangular ends and dense proto-
plasmic contents. As germination advances, the cells elongate until they
are 80-90 /u, long, when absorption is most active, their tips, at the same
time, becoming swollen, club-shaped, and separate from each other,
projecting into the endosperm (Fig. 19). The vascular strands per-
meating the scutellum become more highly differentiated, and many of
the walls of the scutellar parenchyma thickened and pitted.

THE WHEAT PLANT

In addition to its absorptive function, the epithelium secretes diastase
which is responsible for the conversion of the reserve starch grains into
the soluble and diffusible maltose. The aleuron cells also secrete
diastase, but to a smaller extent, and according to Griiss, Puriewitsch,
and others a certain amount of the enzyme is present in the amyliferous
cells of the endosperm.

The first visible action of diastase is seen in the starch grains of the
endosperm behind the upper part of the scutellum after two or three days
of germination at 16° C. Small central slit-like cavities and radiating
cracks first appear in the starch grains ; these rapidly enlarge, and

FIG. 19. — Columnar epithelium of the
scutellum. A, In a dormant grain ;
B, in a germinating grain ( x 275).

FlG. 20. — Starch grains in various
stages of dissolution, i and 2,
From a grain after 5-7 days'
germination ; 3 , from a grain
almost depleted of its endo-
sperm ( x 575).

dissolution proceeds also along the concentric lines of stratification. In
eight or ten days pits and channels are seen ; these commence on the
outside and penetrate into the interior of the grain, its circular outline
being thus broken at various points ; the grains are subsequently eroded
into irregular fragments, which ultimately dissolve and disappear
altogether (Fig. 20).

Concurrent with the starch hydrolysis are modifications in the reserve
proteins of the endosperm. The protoplasmic matrix in which the starch
grains are imbedded expands after imbibing water, and changes into
a hyaline glutinous mass capable of being drawn out into long thin sticky
threads ; later, it loses its elastic quality and is broken down into simpler

GERMINATION OF THE GRAIN 31

soluble nitrogenous compounds, which are absorbed by the growing
embryo with the rest of the soluble plastic materials.

An examination of germinating wheat from day to day shows that the
dissolution of the reserves begins in the amyliferous parenchyma behind
the upper part of the embryo, and spreads gradually towards the apex of
the grain beneath the aleuron layer on the dorsal side, extending also
round the flanks towards the ventral furrow.

Rapid action of the enzymes is promoted by the early hydrolysis of the
cell membranes of the endosperm by cytase, which, according to the
researches of Brown and Escombe and others, is secreted by the aleuron
cells. After the destruction of the cell walls, simple, instead of osmotic,
diffusion is established, and the starch and other cell contents are exposed
to the uninterrupted action of the various enzymes present.

Cytohydrolysis commences very soon after germination begins. In
twenty-four hours, or less, the middle lamella of the starch cells in contact
with the aleuron layer shows signs of dissolution, and in two or three
days the walls of these cells swell and separate from each other, dissolving
later and setting free their contents.

The endosperm tissue is gradually converted from the outside inwards
into a white pasty mass, which becomes more and more liquid and limpid
as the starch grains dissolve, the last portions to disappear being the
amyliferous parenchyma around the furrow of the grain and the aleuron
cells. Leakage of dissolved reserves is effectively prevented by the
semi-permeable testa, which with the cross layer of the pericarp closely
overlaps the edges of the scutellum. The aleuron layer, according to
Haberlandt, is a glandular digestive organ secreting cytase and diastase,
and, according to Bolland, a gluten-transforming ferment also. The
protein granules stored within its cells are utilised in the nutrition of the
protoplasts of the cells.

The layer undergoes chemical changes more slowly than the rest of
the endosperm, little alteration in it being apparent during the first two or
three days except its separation from the adjacent starch cells.

In about a week, when the plumule has attained a length of 3 or 4 cm.,
the aleuron granules in many of the cells have lost their refractive char-
acter or have completely disappeared ; later, they vanish from all the
cells and the remaining brownish granular cytoplasm shrinks and dies,
retreating first from the free walls next the endosperm cavity.

The cell walls from which the cytoplasm has shrunk become crumpled,
and are soon attacked by the cytase, the cell cavities being then exposed
(Fig. 21). Ultimately, the radial and outer walls of the aleuron layer
are broken down and dissolved together with the hyaline nucellar layer.
In 10-15 days at i8°-2O° C., when the first foliage leaf of the young
plant is expanded, the pericarp with the adherent testa is found as a hollow

32 THE WHEAT PLANT

skin within which remains only a watery solution of the last dissolved
reserves ; the plant has become independent of the parental store and is
capable of manufacturing its own plastic materials.

The same sequence of events as that described above is observed
in the germination of grain sown in autumn in the open field, but enzyme
action and growth are less rapid, owing to the lower temperature at this
season. Grains sown during the last few days of October (1914) had
their first green leaves completely unfolded in three or four weeks, and
at the end of November the endosperm had completely disappeared
from the majority of them.

The vitality of the wheat grain and the possibility of its germination
capacity being retained for long periods has been the subject of much
controversy.

Wheat grains of great antiquity have been unearthed from ancient

FIG. 21
bej

[ X 2

21. — Section of a portion of the pericarp and aleuron layer of a grain 16 days from
beginning of germination, a, Cross layer of pericarp ; b, testa ; c, aleuron layer
( x ZT:).

stores and tombs in different parts of the world. In many cases they
are completely carbonised, and in this state are merely friable masses of
charcoal with the form of the grain : such are those obtained from the
neolithic pile-dwellings of Switzerland and the pre-Roman settlements
of this country. Specimens from vases deposited in the tombs of ancient
Egypt, where they have been kept dry, away from atmospheric changes,
show much less alteration, and, except that they are a dark reddish-brown
colour as if scorched, appear normal in size and shape.

I examined a number found by Professor Flinders Petrie in the
Graeco-Roman cemetery at Hawara (about first century B.C.). These were
plump, and similar to some forms of T. durum grown at the present day.
The pericarp was a dark reddish-brown tint, its structure unchanged.
The aleuron layer had retained its original shape, with normal cell walls
and typical round aleuron grains within.

The parenchyma of the endosperm contained unaltered starch grains
which stained a purple colour with iodine, but the delicate cell walls of

GERMINATION OF THE GRAIN 33

the tissue had disappeared and the masses of starch grains in each cell
clung together, preserving the form of the cell-cavity.

The endosperm was very brittle and strongly acid, the acidity being
apparently due to decomposition of the proteins of the grain, for these
were completely altered and no longer gave rise to gluten when water was
added to the finely broken grain.

The embryo had become dark brown, its plumule greatly shrivelled,
and little of its structure was visible, although the scutellum with its
epithelial cells could be recognised in some of the grains.

In grains from spikelets of Emmer from a tomb of the eighteenth
dynasty (1400 B.C., see p. 187) the structure of the pericarp, aleuron
layer and its contents, and the endosperm tissue with its starch grains
were clearly recognisable, although all the parts were more brittle and the
embryo more completely disorganised than in the grains from Hawara.
It is scarcely necessary to observe that the embryos were dead.

Professor Petrie tested samples of grain of Graeco-Roman age which
he found at Hawara immediately after exhumation. The grains were
sown on the banks of a canal in soil of varying degrees of moisture from
mud to fairly dry earth, but none germinated.

From time to time, however, during the last century, reports have \J
been given of the germination of grains taken from Egyptian tombs, but
the evidence when tested is invariably unsatisfactory.

In some cases it is clear that modern grain has been inadvertently ,
admitted to mummy cases, the wrappings of mummies, and vessels taken
from tombs. Sometimes imperfectly thrashed straw and even chaff with
grain have been used as packing for mummies, and mummy cases and
tombs have been used in modern times as convenient stores for grain.

Modern wheat grains appropriately stained have sometimes been
fraudulently mixed with wheat taken from ancient vases, the whole being V
sold to the credulous tourist as " mummy wheat," or introduced into
mummy cases, or placed within the wrappings of mummies.

Genuine vases apparently unopened but containing such grain are
also occasionally sold to travellers.

In regard to recent samples of wheat grains, the duration of their
vitality is dependent chiefly upon (i) the degree of ripeness and the water- \/
content of the grain at harvest, and (2) the temperature and moisture of the
place of storage.

Poorly developed grain, and grain with a high percentage of water in
it, loses its capacity for germination in a year or two, while well-ripened
dry samples retain their vitality for several years.

Storage in a warm room, especially if the atmosphere is damp, leads
to rapid death of the embryo.

Many tests have been made with grains of wheat whose age is beyond

D

34

THE WHEAT PLANT

dispute. These have shown that when kept in an ordinary dry room
grains lose their vitality altogether or have their germinating capacity
greatly reduced in ten years or less.

The rate of loss of vitality is shown' in the following records by Burger-
stein, Carruthers, and Gross :

PERCENTAGE GERMINATION

Age of Seed — Years.

i.

2.

3-

4-

5-

6.

7-

8.

9-

10.

Burgerstein

IOO

Q8

95

94

8?

87

85

79

70

75

Carruthers

IOO

97

92

94

?

88

75

?

29

o

Gross

IOO

IOO

94

95

95

76

72

46

15

6

The grains used by Gross had been stored in open glass beakers or
small bags in a dry unheated room.

In 1907 I tested samples of grain grown on the farmyard manure plot
at Rothamsted. These had been artificially dried after harvest before
being stored in glass bottles.

The following results were obtained :

PERCENTAGE GERMINATION
Year of Harvest

1906.

1905.

1904.

1903.

1902.

1901.

1900.

1899.

1898.

94

74

3

80

40

99

74

99

38

1897-

1896.

1895.

1894.

1893.

1892.

1891.

1890.

1889.

o

67

57

45

57

29

o

0

0

1888.

1887.

1886.

1885.

1884.

1883.

1882.

1881.

1880.

0

0

18

o

i

o

16

0

o

Some of the grains of the harvest of 1882 had retained their vitality
for twenty-five years.

CHAPTER IV

THE ROOTS
GENERAL MORPHOLOGY

THE roots of the wheat plant may be divided into two groups, namely,
(i) the seminal roots, or those belonging initially to the embryo, or which
develop later from the hypocotyl or near it, and (2) the adventitious roots,
which spring from the nodes of the plant, within the soil or just above it.

As previously mentioned, the embryo possesses five roots, namely, a
primary radicle and two pairs of lateral rootlets, which originate from the
hypocotyl in a plane parallel to the face of the scutellum ; a sixth is often
produced later from a point just behind the epiblast at the base of the
plumule, and grows out at right angles to the plane in which the others are
found (4, Fig. 18 ; n, Fig. 22).

The primary radicle and the first pair of seminal rootlets burst through
the tissue of the coleorhiza soon after germination commences, these being
followed subsequently by a second pair ; the sixth rootlet, if it appears at
all, does so only after the others have attained a considerable length when
growth is more advanced. A third pair appears immediately above the
second pair and in the same plane, making in all eight seminal roots.

The apex of each root is pointed, smooth, and more or less mucilagin-
ous. The growing point is protected by the root-cap, the outside of
which is continually being exfoliated as the tip of the root is forced through
the soil, and as rapidly renewed on the inside by the activity of its meristem.

The surface of the rootlet for some distance behind the apex is clothed
with extremely delicate root-hairs.

As the root increases in length and penetrates into fresh regions of the
soil, new root-hairs arise at a short distance from the tip, while those on
the older portions of the root, after collapsing, lose their absorptive power
and turn brown and die. The number of root-hairs is extraordinarily
large ; from 700 to 1000 or more per millimetre length of root are found in
the region of most active absorption.

A number of hairs similar in form and function to root-hairs often

35

36 THE WHEAT PLANT

arise on the basal portions of the coleorhiza soon after the primary root
has broken through it.

The seminal roots are all thin and of nearly uniform diameter through-
out their length ; when turgid and growing they measure from -5 to -75
mm. across ; later, when the soft cortical tissues are shrivelled, they may
become less than half this thickness.

Numbers of very delicate lateral branches, often not more than a
tenth of a millimetre in diameter, appear on the rootlets, generally as soon
as these have reached a length of 4 to 6 inches.

The seminal roots form but a small proportion of the total root-system
of the wheat plant, their function being concerned with the absorption of
the water necessary for the growth of the young plant, especially in its
early stages of development, although they appear to be functional through-
out the life of the plant, for they remain undecayed up to near the time of
harvest. The extent of their development and the depth to which they
descend is influenced by the texture of the soil and the depth at which
the grain is sown. Where the latter is deposited near the surface, the
seminal roots reach their maximum development and penetrate to a
depth of 8-12 inches or more ; where the grain is sown at greater depths
they grow feebly, the reserves of the endosperm being chiefly utilised in
enabling the shoot to grow and reach the light above ground (Fig. 60).

The adventitious roots arise just near the surface of the ground from
the nodes of the main axis and its branches, their position being correlated
with the arrangement of the buds and leaves of the plant. The first to
appear is a pair of roots which grow out from the tillering node of the
primary axis (Figs. 22 and 23), one of them on the right, the other on the
left of the first lateral bud, their points of origin being not quite on the
same level. One of them appears before the other, and in its rapid growth
tends to push aside the adjacent bud from the initial position ; especially
does this happen when the opposite root fails to develop.

At the second node another pair develops in like manner, and from
each of the succeeding somewhat thicker nodes, between which the inter-
nodes remain short, two roots appear arranged similarly in regard to the
nodal bud together with one or more on the opposite side to that on which
the bud arises. Of each pair, the first root to develop is always found on
the same orthostich of the stem.

At the base of the straws near the ground, where the internodes are
longer and the nodes, therefore, not so crowded together, four or six roots
are produced at each node, i.e. two or three pairs symmetrically arranged
as in Fig. 24 ; the upper series of these usually form very strong oblique
props which assist in keeping the straw erect.

Longitudinal planes through the points of origin of opposite adventi-
tious roots of the primary axis cut the plane through the buds and midribs

THE ROOTS

37

of the leaves of the axis at right angles, and are parallel to the plane in
which the three pairs of seminal rootlets are found.

The second and succeeding axes or shoots develop their own inde-
pendent system of adventitious roots, which follow the same general
arrangement in pairs as that of the primary stem, except that at the base
of each bud or axis of the second and subsequent order, a single root first
appears. Moreover the roots on one axis are always arranged in a longi-
tudinal plane at right angles to that
which includes the roots on the axis
of the previous order. The longi-
tudinal plane through the roots of the
first lateral shoot at the tillering node
cuts the plane of the roots of the

120,

FIG. 22. — Arrangement of the seminal
roots and the adventitious roots at .
the "tillering" node of a young .
plant, sc, Scutellum pair ; ep, epi-
blast pair ; co, coleoptile pair ; a,
one of the pair of roots at first node
of primary stem ; b, one of pair at
second node of primary stem ; 1 1 ,
first branch of primary axis; iza,
one of pair of roots at first node of
second branch of primary axis.

col

SC

FIG. 23. — Diagram of the arrange-
ment of the roots, stem axes, and
leaves in the previous figure.

primary axis at right angles ; the sixth seminal root, which springs from
a point just above the epiblast, is in this plane («, Fig. 22).

The adventitious roots do not begin to appear until long after the
leaves of the first or second buds have come above ground. In crops
sown in October and November, particularly when the autumn is wet,
their production is usually delayed until the end of the following January,
at which date the first pair at the tillering node is often not more than
half an inch long : after this date they develop more rapidly. Generally
about the first or second week in March the first roots at the base of the

THE WHEAT PLANT

lateral axes appear, four or five roots being usually present at this time at
the tillering node, the longest being about 3 inches, the shortest about
1 1 inches long, the primary shoot then showing 4 or 5 leaf-blades, the two
secondary shoots 3 or 4 blades each. For a considerable period these
roots are unbranched, and clothed throughout their entire length except
the tip by root-hairs. Later, numerous branches appear, after which
time the piliferous layer shrivels and the roots become thinner and brown.
New roots continue to break out from the nodes of the straws until May
or June, by which time an extensive system has been established in plants
which are not crowded. All these adventitious
roots are of greater length and thickness than
the seminal roots ; some, which spring from
near the base of the straws, are 1-5-2 mm. in
diameter, and ultimately become very rigid in
the first inch or so of their lengths ; in the

FIG. 24. — Arrangement of the adventitious
roots at the three basal nodes of a culm
(xi'5). i, Front ; 2, side view; 3,
transverse sections at a and b.

FIG. 25. — "Anchoring "roots
at the base of a culm (nat.
size).

parts exposed to the air, chloroplasts are present in their cortical tissues
(c, Fig. 30). About 60 per cent of the roots are found in the upper 8-12
inches of the soil, the rest extend to greater depths, a few occasionally pene-
trating 4 or 5 feet or more, especially in deep loams.

The root-system of thin-strawed varieties with upright young shoots
is less extensive than that of thick-strawed forms with a spreading habit
when young.

Moreover, the roots of wheats with erect young shoots are thin and
spread very little near the surface of the soil ; plants with roots of this
habit are easily laid by wind and rain.

The roots of the varieties with young spreading shoots are stout ;
they also make a wide angle with the vertical and anchor the plant firmly
to the soil, isolated and normally grown plants of these varieties being
never laid by wind or rain.

THE ROOTS

39

ANATOMY OF THE ROOTS

In a transverse section through a young seminal root about a centi-
metre from its apex the following tissues are readily recognised : (i) an

outer piliferous layer, (2)

a broad zone of cortex

limited internally by a

well-defined endodermis,

and (3) a polyarch cen-
tral vascular cylinder with

its surrounding pericycle

(Figs 26, 27).

The piliferous layer

consists of thin - walled

elongated cells usually

from 170 to 350 n long and

25-30 [i in diameter.

From many of these are

developed root-hairs

which are produced by the

bulging out and growth

of a small portion of the

cell wall, the point of

origin in each case being usually between the middle and the transverse

septum nearest the root-
tip. These hairs first ap-
pear as very short papillae
about i -5 to 2 millimetres
behind the apex of the
root : further back, where
fully developed, they are
seen to be long tubular
structures closed at the
apex, each from 8 to 12 n
in diameter, and reaching
a length of 1-1-5 mm- m
a damp atmosphere. The
wall of the root-hair is

FIG. 27.— Vascular cylinder of a heptarch seminal root Composed of Cellulose
(xaio). px, Protoxylem ; other letters as in the pre- which allows the absorp-
vious figure. . - .. . .

tion or soil solutions to

proceed freely by osmosis ; the cell wall becomes more or less mucila-
ginous and particles of soil adhere to it.

6

FIG. 26. — Transverse section of a seminal root ( x 100).
a, Piliferous layer ; c, cortex ; e, endodermis ; p, peri-
cycle ; p' small cells of pericycle opposite protoxylem ;
/, protoxylem ; b, phloem ; x, large central vessel.

THE WHEAT PLANT

Lining the hair is a delicate peripheral layer of cytoplasm which
increases somewhat in thickness near the tip. The nucleus of the cell
from which the root-hair arises frequently migrates into the hair and
travels along the lumen towards the apex, at the same time becoming
elongated up to 40-50 p (Fig. 28) : ultimately it becomes disorganised
when the root-hair shrivels and dies.

The zone of cortex which measures about 200 /z across consists usually
of 6-8 layers of parenchymatous cells with small intercellular spaces, the
individual cells being from 20 to 40 /JL in diameter.

The endodermis is a single continuous layer of closely fitting cells ;
its outer walls are thin, but the inner walls in older roots are strongly
cuticularised and exhibit lines of stratification ; the radial walls are

thickened also to some extent, especi-
ally where they join the inner surface
of the cells (e, Fig. 29).

The pericycle is a single layer.
Its component cells, whose radial
diameters are somewhat larger than
their breadths, are of fairly uniform
dimensions except at the points
opposite the protoxylem, where they
are very small. After a time their
walls become more or less sclerotised
and lignified.

The xylem strands are usually 7
or 8 in number in the seminal roots.
One or two very narrow vessels, 5-
10 n in diameter, with closely arranged annular or spiral thickening, con-
stitute the protoxylem of each bundle ; the centripetal vessels formed
later are pitted and wider, being 15-20 /x in diameter. In most primary
roots and some secondary ones the centre of the vascular cylinder is
occupied by a single large pitted vessel about 50 /u. in diameter ; in other
roots there is no central vessel, but two or more large vessels are found
distributed irregularly in the conjunctive tissue.

The phloem bundles are arranged alternately with the xylem strands ;
each generally consists of three cells (b, Figs. 26, 27).

The conjunctive parenchyma of the cylinder has moderately thick
cell walls.

The piliferous layer soon ceases its absorptive function ; the layer of
cortical cells immediately within it then becomes transformed into the
exodermis by suberisation of its walls, and acts for a time as a protective
covering to the root.

Later, much of the cortical tissue shrivels, turns brown, and dies, but

FIG. 28. — Root-hairs ( x 260). n, Nucleus.

THE ROOTS

41

the vascular cylinder retains its freshness, even in the case of the delicate
seminal rootlets, during almost the whole life of the plant.

In roots from which the cortex has decayed, the endodermis, pericycle,
and conjunctive parenchyma of the stele becomes very strongly sclerotic,
but the cells of the bast and the
xylem vessels preserve their normal
character (Fig. 29).

The anatomy of the stronger
adventitious roots which spring
from the nodes of the stems at or
just above the surface of the soil
differs in certain points from that
of the seminal or thinner adven-
titious roots. The exodermis

f>

is

FIG. 29. — Transverse section of a portion of
the vascular cylinder of an old seminal root
examined in May ( x 260). c, Cortex ; e,
endodermis ; p, pericycle ; b, phloem.

often more persistent, and imme-
diately within it there is frequently
developed a zone of sclerotised
cortical cells two or three cells thick (Fig. 30), which, for a time, protects

c

d

FIG. 30. — Transverse section of a portion of an adventitious root from the node imme-
diately above the soil surface ( x 1 50). ex, Exodermis ; h, thick-walled cortical cells ; c,
thin-walled cortical cells containing chloroplasts ; e, endodermal cell with thick inner
wall ; e', thin-walled endodermal cell ; d, lignified pericycle.

the turgid cortex within from collapsing. In the cells of the thin-walled
cortical parenchyma of the stout adventitious roots arising from nodes just
above the soil surface, numerous chloroplasts are found, especially in the
inner layers, the central cylinder then appearing to be surrounded by a
zone of green tissue.

42

For a time the endodermis of these roots is composed of thin-walled
cells not very clearly differentiated from the adjacent cortex and pericycle
cells ; later, the inner walls become extensively thickened except at points

T

a

FIG. 31. — Median longitudinal section of the root-tip of a seminal root (xzio). a,
Initial cells of cortex and piliferous layer ; r, meristem of root-cap ; pr, periblem ;
o, outer layer of periblem with mucilaginous cell walls (c) ; n, endodermal layer ;
s, pericycle layer ; x, central column of cells which develop into vessels.

opposite the xylem, where the walls remain thin, allowing the movement
of water from the vascular cylinder into the cortex for a longer period

than is the case in the more slender roots.

The pericycle at first consists of thin-
walled cells which become thickened later.

At the growing-point of the roots of wheat
are found the usual three layers of meristem
which give rise to the vascular cylinder, the
cortex, and the root-cap respectively.

The plerome or meristem of the vascular
cylinder at the apex consists of a number of
cells somewhat irregularly arranged ; its boundary, however, can be
distinguished without much difficulty from the external periblem. The
latter, from which the cortical tissues are developed, originates from
one or two initial cells (Fig. 31).

FIG. 32. — Recently exfoliated
cells of root-cap ( x 210).

THE ROOTS

43

The piliferous layer, when traced to the root apex, is seen to be the
outermost layer of the periblem. Near the tip its cells have thick mucil-
aginous walls (Fig. 31).

FIG. 33- — Transverse section showing the origin of a lateral root (/>') from the pericycle/)
( x 385). e, Endodermis ; b, phloem.

Adjacent to the initial cells of the periblem is the meristem of the
root - cap. As growth
proceeds, the outer cells
of the root-cap separate
from each other along
the middle lamella and
are rubbed off by friction
with the soil, through
which the roots extend,
the loss being made good
by division of the meri-
stem within. The ex-
foliated cells, at the time
of their separation, are
more or less turgid, and FIG. 34. — Transverse section of a root showing a lateral
contain well-defined root (w)' devel°Pins °PP°site the Phloem < x 2I°)-
nuclei and vacuolated cytoplasm (Fig. 32).

Lateral rootlets are derived from the cells of the pericycle opposite
the phloem of the vascular cylinder, and not from points opposite the
xylem as in the roots of many plants (Figs. 33, 34).

CHAPTER V

THE LEAVES
GENERAL MORPHOLOGY

THE wheat plant possesses several forms of leaves, viz. : (i) the scutellum,
(2) the coleoptile, (3) the ordinary green foliage leaves, (4) the prophylls
of the lateral axes, and (5) the glumes of the inflorescence. The scutellum
has been considered already and the glumes will be dealt with later.

The coleoptile or plumule sheath which encloses the first shoot is a
hollow cylindrical structure with a bluntish apex slightly curved to one
side. I regard it as the primary prophyll or a leaf-sheath the blade of
which is absent. Near its tip, on the side away from the scutellum, is a
small slit through which the foliage leaves emerge (Figs. 9, 35).

In some forms of wheat the coleoptile is pale green or colourless,
in others it is pink. The extent of its development varies with the depth
at which wheat-grains are deposited in the ground. In crops drilled
about an inch deep in the ordinary way in the field it comes to the surface
or a few millimetres above it. In the case of grains sown i -2 cm. (about
half an inch) below the surface, the coleoptile grows to a length of 1-5
cm. : in those sown 2-5-5 cm- (I *° 2 inches) it usually measures about
3-5 cm., while in grains covered with 10 cm. (about 4 inches) of soil it
reaches a length of about 6 cm.

It has few chloroplasts in its tissues and possesses little photo-
synthetic power : it functions chiefly as a protective cover to the young
foliage leaves during their upward growth through the soil.

The foliage leaves of wheat, like those of grasses generally, are arranged
on the culms alternately, in two opposite vertical rows, each leaf having a
divergence of 180° from the next above and below it. This relationship
is seen in the position of the coleoptile and first foliage leaf of the embryo.

In the case of lateral shoots, the first leaf is a binerved prophyll, some-
what resembling in form the coleoptile already described. It is semi-
cylindrical, with an opening at the apex, through which the green foliage
leaves emerge later. The upper part is green with deflexed hairs along

44

THE LEAVES

45

the margins of the two nerves : the lower part is colourless. The prophyll
grows usually to a length of an inch or more and is inserted on the lateral
shoot in such a position that its flattened side is turned towards the axis
from which the shoot arises. The first foliage leaf, however, which suc-
ceeds this prophyll has a divergence from
the latter of 90° only, to the right or left,

A

FIG. 35. — Tip of the coleoptile with first foliage
leaf emerging, i, Front; 2, back view.

so that it and subsequent leaves which
follow the ordinary rule of 180° diver-
gence from each other are arranged in a
plane at right angles to the foliage leaves
of the preceding axis (Figs. 37, 38).

This is the normal arrangement of the
leaves upon stems and lateral shoots in
practically all grasses as first indicated by
Godron.

In the bud the leaves are convolute,
the blades being rolled up from one
margin to the other. The rolling alter-
nates from leaf to leaf, one in which the
right margin is folded over the left being
followed normally by another in which
the left margin covers the right. The first foliage leaf of the embryo
above the coleoptile is not, however, always rolled from the same side ;
in some plants the right margin is folded over the left, in others the
opposite arrangement occurs (Fig. 36). There are therefore two forms
of plants which may be termed respectively " right- " and " left-handed,"

FIG . 3 6 . — Young plants . i , " Right -
handed " ; 2, " left-handed " ; A
and B, transverse sections of the
shoots i and 2.

THE WHEAT PLANT

" right-handed " plants being those in which the right margin of the first
green leaf-blade as seen from the front is folded over the left, " left-
handed " being applied to those in which the left margin covers the right.
In Figs. 37 and 38 the order and mode of rolling of the leaves of

the primary axis of
the plant and the
first two or three
lateral branches is
indicated. In the
majority of right-
handed plants the
first green leaf of
all the lateral shoots
of the main stem is
inserted on the left
side of the secondary
axis which bears it,
the axis being
viewed from the en-
dosperm side of the
FIG. 37. — Diagrams of three types of leaf arrangement, A, B, scutelllim : in those
and C, in young primary shoots, s, Scutellum ; c, coleoptile ;

|, first foliage leaf; p1, p2, prophylls of first and second which are lett-
lateral shoots; a, first foliage leaf of shoot in axil of leaf j ; handed the first
ft, first foliage leaf of shoot in axil of second leaf f .

green leaf arises on

the right side of its
branch. In a small
percentage of cases
s an alternate arrange-
ment of the first
leaves on the lateral
shoots occurs (Figs.
37 and 38) : other
abnormalities are

met with both in
FIG. 38. — Diagram of the axes and leaf arrangements in young , . ,

plant, s, Scutellum; c, coleoptile; i, primary axis; n, trie SUCCCSSlOn and
first, 12, second lateral axis of i ; |, first, f second foliage j^ the folding of the
leaf of primary axis ; /j, prophyll of axis 12 ; TV, first leaf of .
axis 12 ; T{T, prophyll of first lateral axis of axis 12. leaves.

Out of 500 seed-
lings of a pure line of T. vulgar e (Squarehead type), I found 276 with the
left margin folded over the right, and 224 " right-over-left " : in a pure
line of T. turgidum (Blue Cone) 256 were " left-over-right " and 244
" right-over-left " : in both these cases there is a preponderance of
" left-over-right " plants.

P

a

THE LEAVES

47

The foliage leaves are of the ordinary linear gramineous type, possess-
ing sheath, blade, and ligule, with a pair of auricles at the base of each
blade.

The leaf-sheath, which is split in the upper parts but entire near the
base, encircles the culm or straw, protecting the latter from damage by
frost, drought, and insect attacks ; it possesses considerable strength and
serves as a support for the young growing internode within it, and especially
for the basal intercalary growth-zone, which remains soft and weak after
the rest of the internode has become rigid.

The leaf-blade develops much more rapidly than the sheath, and in
young plants reaches a length of two or three inches, while the sheath is
not more than a few millimetres long. The
sheath grows very little before the blade
has attained its maximum size, but grows
rapidly when the stem internodes begin to
lengthen.

The greater portion of the sheath is
somewhat thicker and more transparent
than the blade, with thin transparent
margins. For a distance of 3-5 mm.
above the point of insertion on the stem
it is considerably thickened ; this short
swollen part is about i mm. thick, and
is often erroneously assumed to belong to
the node of the culm (Fig. 39).

In many varieties the surface of the
sheath is glabrous, while in others it is
clothed more or less uniformly with short
deflexed hairs. Sometimes there is a line
of cilia near the outer margin of the sheath,
and in T. aegilopoides, T. monococcum, and
T. dicoccoides the hairs on the swollen base are longer than those of other
wheats, forming a conspicuous white muff-like band round it.

The leaf-blade is linear- and parallel-veined ; its midrib projects on the
back and is continued some way along the sheath as a raised line. It is
twisted into the form of a loose right-handed screw, the twist being
increased by holding the tip of a young leaf and turning clockwise (Fig.
40). The direction of the screw is normally the same for all leaf-blades,
although exceptions are occasionally found, and it is independent of the
alternately reversed convolute rolling of the young leaves previously
noticed.

The halves right and left of the midrib are of different widths, the
wider and narrower portions alternating regularly in successive leaves.

FIG. 39. — i, Node; 2, longitudinal
section through the node ( x 2).
b, Diaphragm ; a, thickened leaf-
base.

48

THE WHEAT PLANT

The narrow side is also slightly longer than the wider half, an arrangement
which places the attached auricles one below the other alternately.

The blade of the first green leaf of the wheat plant differs from all the
succeeding ones in having a somewhat stiffened bluntish apex, in outline
like an arch, a form which enables the top to push its way through the
soil without damage (Figs. 35, 41). The characteristic and easily recog-
nised shape of this leaf greatly assists in the determination of the number
of separate plants in a crowded row of seedlings in the field.

The blades of all other foliage leaves are drawn out into long acuminate
points which are slightly hooded. About i| to 2 inches from the top of a
full-grown leaf is a peculiar constriction, the margins curving inwards at

that point giving to the blade the
appearance of being formed of two
pieces (3, Fig. 41).

On the upper surface is a series
of longitudinal and slightly raised
ribs, which on the young leaves of
T. dicoccoides, T. dicoccum, and T.
turgidum are covered with soft
velvety hairs of nearly uniform
length ; in T. monococcum the hairs
are short and scabrid, in T.
durum the blades are glabrous,
while in T. vulgar e, T. Spelta, T.
aegilopoides there are long hairs on
123 the summits and shorter ones on the

Fl?- 41-— ', Apex of the first flanks of the ridges (Figs, in, I64).1
foliage leaf; 2, apex of v 6 ' 7'

second foliage leaf; 3, apex The long hairs of 1. aegilopoides
of older leaf -blade (nat. tend to disappear when the plants
size). .

are brought under cultivation.

The lower surface of the leaf is not ribbed and is usually smoother than
the upper surface.

At the base of the upper surface of the leaf-blade is the ligule, a thin
membranous structure which closely invests the stem and prevents the
access of rain, dust, aphides, and other insects between the leaf-sheath
and the stem (/, Fig. 42). It is colourless,1 and its free edge is irregularly
cut and fringed with minute hairs. The ligules of the upper leaves are
from 3 to 4 mm. long, those of the lower leaves much shorter.

The auricles are curved claw-like appendages attached to the base of
each leaf-blade, where they loosely clasp the sheath and stem from opposite

1 The ridges and hairs on the surface of the leaf are readily seen on bending the blade
outwards and examining with a pocket lens ( x 8) the folded part as a silhouette against
the sky or other light background.

FIG. 40. —
Young leaf-
blade show-
ing spiral
twist (nat.
size).

THE LEAVES

49

sides. They are usually pale green or pinkish in colour, and their margins
and tips are often fringed with a few long unicellular hairs (a, Fig. 42).

As indicated previously, they are not exactly opposite to each other ;
the lower one is always connected with the narrower and longer half of
the leaf-blade, and as this half alternates from right to left in successive
blades, the lower auricle follows the same succession, encircling the sheath
and straw from the right and left alternately.

On young leaves and on those of the lower nodes of the straw which
die off soon, the auricles are small, but on the upper leaves they are more
strongly developed. The size also varies with the race and variety of
wheat, those of T. Spelta and T. dicoccum being the largest ; the free claw
in some cases attains a length of 4 or 5 mm.

The number of leaves on a well-de-
veloped straw of the various races of wheat
depends partly on the variety and the
vigour of the individual plant. In most
varieties of T. vulgare as grown in the
field, about 70 per cent of the straws
possess 6 leaves, 8 or 10 per cent have
7, and about 20 per cent only 5 leaves. In
varieties of T. turgidum 20 per cent have 7
or 8 leaves, about 60 per cent 6 leaves, the
rest only 5 or less.

The total length of each leaf, i.e. the
combined length of its sheath and blade,
increases from below upwards to the fifth,
which is the longest on the straw ; the
sixth or uppermost is invariably shorter
than the one immediately below it. The
average lengths of the leaves (sheath and blade) of the primary straw
of the different races of wheats are given below.

a-

FIG. 42.

i — L

Auricle of leaf ; /, ligule
(X2).

AVERAGE LENGTH OF SUCCESSIVE LEAVES IN INCHES

ist.

Total Length

(Lowest.)

2nd.

3rd.

4th.

5th.

6th.

of Five Uppermost
Leaves (Inches).

T. monococcum

12-5

I3-5

14-0

«'S

11-3

62-8

T. vulgare

12 5

IS'4

16-6

17-7

17-1

79'3

T. Polonicum

16-5

17-0

16-8

17-0

18-0

85-3

T. amyleum

14-5

I5-5

I7'5

16-8

18-0

18-0

85-8

T. compactum

iS'3

17-0

18-0

19-2

17-7

87-2

T. Spelta

I3-3

I5-4

18-0

18-5

19-0

18-5

89-4

T. durum

14-4

16-9

19-4

20-1

20-5

19-7

96-6

T. turgidum

16-2

18-4

20-5

21-8

22-4

2I'I

104-2

5°

THE WHEAT PLANT

The width of the blades of different leaves increases more or less
regularly from the lowest to the uppermost leaf of the culm : the average
breadth across the middle of the blades of the successive leaves of fifty
well-developed straws of T. vulgare (Squarehead form) was :

Leaf.
Width

ist
6-3

2nd
9-2

3rd
10-3

4th

5th
H'3

6th

17-1 mm.

The longest leaves are possessed by T. turgidum and T. durum.

In the following table is given the average separate lengths of sheath
and blade respectively, deduced from measurements of successive leaves
of the chief straw of fifteen plants of the different wheats mentioned.

LENGTHS OF SHEATH AND BLADE (inches)

ist.
(Lowest.)

2nd.

3rd.

4th.

5th.

6th.

Sh. Bl.

Sh. Bl.

Sh. Bl.

Sh. Bl.

Sh. Bl.

Sh. Bl

T. monococcum

4-0 8-5

4'5 9'0

5-0 9-0

5'0 6-5

6-3 5-0

T. vulgare

4-9 7-8

5-8 9-6

6-5 io-i

7'4 i0'3

9-3 7-8

T. Polonicum

7'° 9'5

7-0 io-o

7-3 9-5

8-0 9-0

10-5 7'5

T. amyleum

S'o 9'5

5'5 lO'O

6-3 II-2

6-6 10-2

7'7 10-3

9-5 8-5

T. compactum

5'7 9'6

6-0 n-o

6-6 11-4

7-5 11-7

9-0 8-7

T. Spelta .

5-3 8-0

5-8 9-6

7-0 ii-o

7-5 10-8

8-2 ii-o

9-6 8-8

T. durum .

6-0 8-4

6-6 10-3

8-0 11-4

9-0 ii-i

9-0 11-5

10-7 9-0

T. turgidum

6-4 9-8

7'3 "'I

8-3 12-0

9-1 12-7

10-2 I2'2

n-8 9'3

DIFFERENCE BETWEEN SHEATH AND BLADE

Leaves.

2nd.

3rd.

4th.

5th.

7th.

T. monococcum

4-5

4'5

4-0

i'S

-1*3

T. vulgare

2-9

3-8

3-6

2-9

-i-5

T. Polonicum

2-5

3-0

2-2

i-o

-3-o

T. amyleum

4'5

4'9

3-6

2'5

— i-o

T. compactum

3'9

5'0

4-8

' 4-2

-0-3

T. Spelta

3-8

4-0

3'3

2-8

-0-8

T. durum

3'7

3'3

2'I

2-5

-0-3

T. turgidum

3-8

3'9

3-6

2-0

-2-5

On comparing the figures it is seen that the length of the sheath of
successive leaves increases upwards, the uppermost being the longest.
The blades also exhibit a successive increase from leaf to leaf up to the
fifth, but that of the sixth or uppermost is always shorter than the blade
of the leaf below it.

In all leaves except the terminal one the average length of the blade
is from 2| to 4 inches longer than the sheath associated with it ; but the

THE LEAVES 51

blade of the uppermost leaf is usually 1-2^ inches shorter than its
sheath.

The length of any particular sheath is approximately equal to the
arithmetical mean of the sheath lengths of the leaves immediately above
and below it, a relationship similar to that which exists between the
lengths of separate internodes of fully developed straws.

The figures for the sheaths of T. turgidum, T. Spelta, and T. compactum
are :

•

T. turgidum.

T. Spelta.

T. compactum.

Length.

Length.

Length.

Found.

Calculated.

Found.

Calculated.

Found.

Calculated.

inches.

inches.

inches.

inches.

inches.

inches.

Second sheath
Third sheath
Fourth sheath
Fifth sheath .

7'3

8-3
9-1

IO-2

7'35

8-2

9^5

10-45

5-8
7-0

U

6-75
7-6
8-5

6-0
6-6

7'5

6'-i5

6-75
7-8

In the other races of wheat mentioned, the agreement between the
calculated length and that actually observed is not quite so close, a dis-
crepancy which is possibly due to the averages given in the table con-
taining measurements of leaves which had not all reached their maximum
development or whose normal growth had been disturbed.

ANATOMY OF THE LEAVES
i . The Coleoptile and Prophylk

The tissues of the coleoptile are of a simple character. Outside is
an epidermis of long cells of rectangular outline with thin straight walls :
there are no hairs present. One or two single rows of stomata run from
the base to near the tip, on each side of, and parallel to the vascular strands
within the leaf, the number of stomata increasing towards the apex, where
they are crowded together in an irregular manner.

The few chloroplasts present are found in most abundance in the first
layer of cells beneath the epidermis and in the parenchyma surrounding
the vascular bundles ; they remain colourless except when the coleoptile
is exposed to light.

In the majority of wheats two vascular bundles are found on opposite
sides of the coleoptile. These are simple and straight for the greater
part of their length : at the summit they bend towards each other and

THE WHEAT PLANT

meet in a backward pointing V-shaped curve, one or two very fine strands
of tracheids anastomosing with them just below their point of union.

In T. dicoccoides and Indian and Abyssinian forms of T. dicoccum three
to six vascular bundles are present in the coleoptile (Fig. 16), a fact which
supports the view that the latter is homologous with the leaf-sheath or
prophyll rather than with the ligule. The bundles consist of a few small
scattered xylem elements of nearly uniform diameter and a mass of phloem
protected on the outside by a single or double layer of thick-walled
cells (Fig. 44).

The epidermis of the prophylls is composed of elongated rectangular
cells with thin straight walls. Along the two margins of the flat side of
the prophyll are reflexed unicellular hairs, and
near the apex two lines of stomata run on op-
posite sides of the nerves. The mesophyll con-
sists chiefly of delicate parenchyma ; the cells of

FIG. 45. — Outline

FIG. 43. — Stoma of the and transverse

coleoptile (cf. Fig. 49) FIG. 44. — Transverse section of a vascular section of a
( x 380). bundle of the coleoptile ( x 280). prophyll.

the apical portion round the nerves contain chloroplasts, the rest of the
tissue being colourless. The convex part of the prophyll is transversed
by 10-12 very thin, straight, vascular strands, two stronger bundles
running from the base to near the apex in the angles where the convex
and flat side meet (Fig. 45). Here and there, especially near the tip, are
fine anastomosing lines of tracheids. A well-defined strand of stereome
strengthens each of the two angles, and, in addition, two or three cells
of rudimentary sclerenchyma are found beneath the outer epidermis of
the convex side associated with the fine vascular strands.

THE LEAVES

53

2. The Foliage Leaves

The structure of the green foliage leaves of the wheat plant is more
complicated. Covering the surface is (i) a somewhat elaborate epidermis
with characteristic stomata and trichomes of various forms ; within is (2)
the green assimilating parenchyma, (3) the conducting vascular bundles,
and (4) longitudinal strands of fibrous stereome or supporting tissue.

FIG. 46. — Transverse section of the leaf-blade, s, Stereome.

FIG. 47. — Transverse section of the leaf-sheath ( x 25). o, Outer ; i, inner epidermis ;
/, lacuna ; s, stereome ; a, mesophyll.

(i.) THE BLADE AND SHEATH, (a) The Epidermis. — The epidermis
of the blade of the foliage leaves is composed of a number of diverse
elements arranged in parallel rows along the long axis. Some of the
individual rows consist entirely of elongated cells placed end to end, each
cell appearing in longitudinal section as a narrow rectangle 150-300^
long and 15-20 /z wide. In other rows short square cells 15-20 /JL across
are intercalated here and there between the long cells : characteristic
lines of long cells alternating with stomata are also present, and, except
in T. durum, trichomes or hairs of various lengths are found scattered along
the rows at more or less regular intervals, the whole arrangement as seen
in surface view being illustrated in Fig. 48. On the upper surface of the
blade is a series of longitudinal ridges or ribs, the lower surface being
almost flat. The epidermal cells covering the ridges differ somewhat
in form and arrangement from those over the furrows and along the edge
of the leaf.

Running along the summit of each ridge is a single row of elongated
thick- walled and pitted cells alternating with hairs.

On the flanks of the ridge, right and left of this central line, are three
to five rows of long cells often interspersed with short ones and hairs :

54

THE WHEAT PLANT

parallel to these, at the base of the ridge, are single or double lines of
stomata.

In the furrow between two ridges is a band of three to seven rows of
elongated cells, whose walls are thinner and not so distinctly parallel to
each other as those of the long cells in other parts of the blade : these are
termed " bulliform cells " by Duval-Jouve and " motor cells " by Pe'e-
Laby. They are somewhat shorter than the ordinary long cells of the
epidermis, measuring 100-250 /z in length and 17-20 ^ in width : in
transverse section they are seen to be deeper and of different form from
the neighbouring cells, being spread out in the form of a fan, each cell
being narrow at the outer part, and broader or smaller at the base within
the leaf, where it is in contact with the assimilating parenchyma (a, Fig. 56).

FIG. 48. — Upper surface of the leaf-blade, a, Hair ; b, c, rows of stomata ;
e, rows of motor cells ; n, optical section of mesophyll cells ( x 105).

When transpiration is excessive these cells lose water and the edges
of the lamina curve inwards, tending to check any further loss through
the stomata, which are abundant on the upper surface. On reabsorption
of water the blade becomes flat again.

The rest of the epidermis is of fairly uniform thickness, 25-30 fj, deep,
each cell about as long as broad, with a convex cutinised outer wall 4-5 //,
thick, the inner and lateral walls being much thinner.

Several rows of epidermal cells bordering the edges of the leaf-blade
have specially thick sinuous walls.

The trichomes or hairs on the leaf-blade are always unicellular, and
vary much in length and stoutness ; some of them on the edges of older
leaves are little more than blunt prominences, others are short and stout,
20-30 [L long, with fine curved points rendering the surface scabrid ; on
soft velvety leaves of T. dicoccum and T. turgidum, the hairs often attain

THE LEAVES

55

a length of 300-400 /z, while on T. aegilopoides the sparsely distributed
long hairs on the surface of the leaf-blade measure i mm. or more.

When present the trichomes are distributed in somewhat regular
order, at short intervals along the rows of epidermal cells on the margins
and surfaces of the blade. There are usually more on the upper than the
lower epidermis, and in T. vulgare the longest hairs are found in the row
of thick-walled cells which runs along the summit of each ridge ; shorter
ones are frequent on the flanks of the ribs, but few are seen among the
motor cells.

Each stoma of the leaf-blade consists of four
cells, the two guard cells being narrow, with speci-
ally thickened walls round the stomatal pore and
thin-walled widely dilated ends : the pore when
closed appears as a narrow slit 30-40 /x long

(Fig. 49)-

The stomata are only very slightly sunk below

the general level of their neighbouring cells and
run in single or double longitudinal lines, each
stoma alternating more or less regularly with
an elongated epidermal cell. On the upper sur-
face the rows on one side are contiguous with
the motor cells, and on the other side lie next
to several rows of long cells without stomata.

In transverse section the pores of the stomata
are seen to be in communication with large
intercellular cavities in the mesophyll. The epi-
dermis of the lower surface of the leaf resembles
that of the upper surface in form and arrange- FlG 49_I? Stomaj sur.
ment of its elements, but there are no definitely face view; 2, median
marked ridges upon it, and motor cells are
absent : the cell walls are sinuous and much
thickened, especially in the lower half of the blade.
The hairs are also fewer in number, and the
stomata more often in single lines than on the upper surface.

The ratio of the number of stomata on the upper and lower epidermis
respectively is usually about 10:7, the number found on the upper surface
in the few cases examined being about 7000 per square centimetre.

The outer epidermis of the full-grown leaf-sheath is composed of rows
of elongated cells similar in form to those of the blade, but the walls are
more sinuous and stoutly thickened (Figs. 50, 52).

The stomata are arranged in lines, as on the lamina, and a few hairs
are usually present. Over the thick nodal base of the sheath the epidermis
consists of much shorter cells, in the rows of which, at more or less

transverse section ; 3,
transverse section across
one end ; 4, longitudinal
section of a guard cell
(x38o).

THE WHEAT PLANT

regular intervals, are pairs of cells, the upper one oval, the lower one
broader and curved (Fig. 50) ; trichomes when present arise from the

oval cells.

The cells of the inner epidermis of
the sheath are thin - walled. Inter-
calated in the rows lying between the
points opposite the vascular strands,
especially near the upper part of the
sheath, are short hairs, the tips of
which are directed forwards (Fig. 51).
A few stomata of a simpler pattern
than those of the blade are found in
this region of the inner epidermis ;
these communicate with larger lysigen-
ous lacunae in the mesophyll.

(b) The Parenchyma of the Leaf. —
FIG. so.-Outer epidermis of the base of Th parenchyma of the blade of the
a young leaf-sheath ( x 210). J .

foliage leaf consists chiefly of thin-

walled assimilating tissue, containing lenticular chloroplasts 4-5-6 p in
diameter. Just within both the upper and lower epidermis the cells are

FIG. 51. — Inner epider-
mis of the leaf-sheath
f x 5°).

FIG. 52. — A, Outer epidermis of the leaf-
sheath ; B, transverse section of the same
( x 210).

somewhat elongated and arranged in a more or less regular manner,
with their long axes at right angles to the surfaces of the lamina, both
layers suggesting a resemblance to the palisade parenchyma of a leaf of a

THE LEAVES 57

dicotyledon. In optical surface view the cells appear as circles from
20 to 32 p in diameter («, Fig. 48).

The cells of the chlorophyll-containing tissue in the central part of
the leaf are much more irregular in shape and are loosely packed, with
larger intercellular spaces between them.

The assimilating tissue of the leaf-sheath is arranged in parallel
strips between the vascular strands, chloroplasts being present to a depth
of three or four cells from the outer epidermis only ; the rest of the
parenchyma on the inside of the bundles and near the inner epidermis
is colourless.

Extending longitudinally throughout the greater part of a full-grown
sheath are large lysigenous cavities between the vascular bundles
(/, Fig. 47).

The ground tissue of the thick nodal part of the sheath is composed
of parenchymatous cells somewhat flattened when young and more or
less hexagonal in transverse section, with unlignified walls 4 p thick

(Fig. 58).

Chloroplasts are present especially in the subepidermal layers. In
each cell on the outside of the stereome, and between the vascular bundles,
there is a single crystal or cluster of crystals of calcium oxalate (Fig. 58).
The crystals appear to be waste products connected with the processes
involved in the thickening of the neighbouring cell walls. Starch grains,
4 /z in diameter, are also found in the two or three cell layers of the ground
tissue on the inner side of the xylem of the bundle.

(c) The Vascular Bundles. — Running through the ground tissue of the
blade, somewhat nearer to the lower than the upper surface, are the
vascular bundles, three types of which may be recognised, viz. : (i)
stout bundles running longitudinally from the base to the apex of the
blade, (2) more slender bundles parallel to and lying between the strong
ones, and (3) very fine transverse veins which cross more or less obliquely
from one longitudinal strand to another.

The number present in a leaf depends upon the size of the latter and
its position on the plant, the upper leaf often possessing more than three
times as many bundles as the lowest leaf on the straw. The first foliage
leaf of a seedling plant of T. vulgare usually has 11-13 bundles, 3-5 of
which may be traced to near the apex. In a full-grown leaf on a well-
developed straw there are generally 9-13 stout bundles, and between
each pair of these there may be 1-5 slender intermediate strands, the
most frequent number being 2 or 3.

Below are given the number found in the broadest part of the several
leaves on a full-grown straw of T. vulgare.

When the bundles are traced from the base towards the apex, some of
them, instead of remaining as single strands throughout their course,

THE WHEAT PLANT

Stout Bundles.

Fine

T c,n(

T 1 •

Left Half.

Midrib.

Right Half.

Bundles.

5th or uppermost

6

I

6

66

79

4th ....

6

I

6

42

55

3rd ....

6

I

6

31

44

2nd ....

4

I

4

28

37

ist leaf near base of

straw

4

I

4

18

27

are seen to divide where the leaf becomes wider, the two halves gradually
separating into distinct bundles.

On the other hand, in the upper part of the lamina,
some of the finer bundles disappear not by blindly
ending in the mesophyll, but by turning to one side
and uniting with another ; others somewhat stouter
divide in two, the separate parts then curve rapidly
to the right and left respectively and become joined to
neighbouring parallel bundles as indicated in Fig. 53.
On account of these divisions and fusions of
bundles, the number found varies with the part of
the leaf chosen for examination, being greatest in
the middle and least near the tip.

The vascular bundles from the leaf-blade pass
uninterruptedly into the sheath and downwards into
FIG, 53.— Course of the tne n°de, where they enter the diaphragm between
vascular bundles near two adjoining internodes (pp. Q5-Q7).
the apex of a leaf- ... ; J, , i -I 'u 1 J

blade ( x 5). All the bundles are collateral, with the xylem towards

the upper surface of the blade, the phloem below.

In the xylem of the stout strands there are one or two vessels about
20 IJL in diameter, with annular or spiral thickening, and to the right and
left of these are two vessels, usually of wider bore, with narrow elliptical
pits.

In the slender intermediate bundles few or no vessels are visible,
although xylem and phloem are both present.

The very fine anastomosing bundles, which cross from one parallel
bundle to another, consist of short tracheids about 6 p, in diameter, with
parallel thin-walled parenchymatous cells about 4 ^ across accompanying
them (Fig. 54).

Each bundle is surrounded by an inner and an outer sheath (Fig. 55) ;
the former (the " mestome " sheath of Schwendener) completely encloses
the vascular strand, and is composed of elongated thick-walled cells

THE LEAVES

59

usually with square ends and a few simple pits ; the outer or "parenchyma
sheath " is more conspicuous, especially round the smaller bundles, and
consists of thin- walled cells, often almost circular in transverse section,

FlG. 54. — Fine strand of tracheids crossing from one
large bundle to another (optical view of portion
of leaf cleared with chloral hydrate ( x 135).

FIG. 55. — Transverse section of the
leaf through a small vascular
bundle (xaio). s, Stereome ;
i, inner ; o, outer bundle sheath.

measuring 20-30^ across and 100-150 /z in length, their long axes parallel
to the long axes of the leaf. It only partially encircles the bundle, being

O

FIG. 56. — Transverse section of a leaf through a large vascular bundle ( x 210). a, Motor
cells ; 5, stoma ; n, mesophyll cells ; b, phloem of vascular bundle ; o, outer sheath
of bundle.

discontinuous on the phloem side, and in the larger bundles is united to
the girder of stereome mentioned below. Unlike the ordinary parenchyma
of the blade, the outer sheath contains few or no chloroplasts except when

6o

THE WHEAT PLANT

young ; in the nodal part of the leaf-sheath, however, the outer bundle-
sheath retains its chloroplasts for some time, accumulating later a large
number of fine round starch grains, each about 4 /z in diameter.

(d) Stereome of the Leaf. — Above and below the bundles, and arranged
parallel with them along the leaf, are strands of stereome or supporting
tissue consisting of sclerenchymatous fibres, some of the walls of which
are so much thickened that the lamina of the cells are almost obliterated.

In the case of a thin
intermediate bundle, small
isolated stereome strands,
consisting in some in-
stances of three or four cells
only, are present immedi-
ately within the upper and
lower epidermis above and
below the bundle, the space
between the latter and its
corresponding sclerenchy-
matous strands being
occupied by ordinary thin-
walled ground tissue. The
stereome above and below
a stout bundle is, however,
more extensive, usually fill-
ing up the space and form-
ing a strong girder between
the epidermis and the
sclerenchymatous bundle-
sheath with which it be-
comes united.

In the leaf -sheath the
vascular bundles are gir-
dered to the outer epidermis
by strands of stereome
continuous with those of
the under side of the blade.

FIG. 57, i. — Transverse section of the leaf-base just
above a node (xzs). s, Stem; b, leaf-base;
r, stereome of bundle.

FIG. 57, 2. — Transverse section half an inch above i
(*35).

On the inside of the sheath, which corresponds with the upper surface
of the blade, only a few cells of stereome are seen beneath the epidermis
opposite to the larger bundles but not connected with them as a girder ;
opposite to the smaller bundles none is developed. Traced downwards,
the stereome increases in amount, reaching a maximum in the thick nodal
portion of the sheath, where it forms a strong strand of semi-elliptical
section on the outside of each vascular bundle and free from the epidermis

THE LEAVES

61

(Fig. 57) : the fibres in the middle part of the thick leaf-base remain
unlignified for some time after those in the upper and lower sections
of it have changed.

In addition to the stereome associated with the vascular bundles
there is a stout band of mechanical
tissue 80-100 /A wide, along both
margins of the leaf-blade just within
the epidermis, which greatly assists in
keeping flat the edges of the leaf ; it
is absent from the edges of the leaf-
sheath.

(ii.) THE LIGULE. — The ligule,
often 2-5 to 3 mm. long, is an emer-
gence of thin-walled parenchyma
arising at the point of union of blade
and sheath, its inner epidermis being
directly continuous with that of the
leaf-sheath ; it possesses no vascular
tissue.

At the base it is three or four cells
thick. Along the thinner upper free
edge many of its cells are elongated,
forming an irregular fringe of uni-
cellular hairs, each 60-80 JJL long, and
8-10 /A broad. Both surfaces are with-
out stomata or hairs, and the walls of
the epidermis, especially at the margins
and back of the ligule, frequently show
faint transverse scalariform pucker-
ing. The cells are 5-10 times as long as broad, most of them colourless,
but some contain small chloroplasts, and others a pinkish cell-sap.

(iii.) THE AURICLES. — The auricles consist of parenchyma ; some of
their epidermal cells are unicellular hairs from -5 to i mm. long.

FIG. 58. — c, Transverse section of a
portion of the leaf-base ( x 105) ;
s, stereome of bundles (cf. Fig.
57) ; a, longitudinal section of paren-
chyma cells in c ; b, calcium oxalate
crystals from cells in c.

CHAPTER VI

THE STEMS : " TILLERING," " SHOOTING," AND " LODGING "
GENERAL MORPHOLOGY

THE culms or straws are erect, elastic, cylindrical, and in some wheats,
especially T. dicoccoides, T. turgidum, and T. Polonicum, more or less
furrowed, with smooth or scabrid surfaces. The upper parts in a young
state are green, and carry on the process of assimilation extensively, the
straw and investing leaf-sheaths being able to supply sufficient food to
mature and ripen grain even when the leaf-blades are removed at the
time of flowering of the ear.

When ripe the colour in most wheats is a pale yellow ; in some, how-
ever, the upper internode is a characteristic purple tint about the time of
harvest.

In normally grown plants of Bread wheat the majority of the culms
possess six nodes, although straws with seven and also with five are not
infrequent.

At the nodes, which should not be confused with the swollen bases of
the leaf-sheaths, the stem is much contracted in diameter and always solid,
the vascular bundles being there crowded together and interlaced to form
a strong diaphragm between two internodes (Fig. 39).

In most varieties of Bread wheats the internodes are hollow, but
in many kinds of Rivet and Macaroni wheats the central part of the straw
is completely filled with a soft pith. In the hollow straws the cavity
begins to form when the young internode is about half an inch long, the
separation of the cells appearing first in the upper part of the internode
just below the diaphragm.

The whole length of the lower, and much of the upper internodes are
invested by the leaf-sheaths, which function as protective and supporting
structures to these sections of the straw, especially when the latter are
immature.

The length attained by a straw is influenced by a number of inde-
pendent factors, some races, e.g. Rivet wheats ( T. turgidum) and Macaroni

THE STEMS

wheats (T. durum), usually have tall stems, while Small Spelt (T. mono-
coccum) and Bread wheats (T, vulgar e) are generally shorter, but tall and
more or less dwarf varieties occur among almost all the races of wheat.
The particular length attained is not only dependent upon the race or
variety of wheat, but also upon various external conditions.

The application of nitrogenous manures increases the length of the
straw : phosphates and potash fertilisers tend to reduce it.

Plants grown at wide intervals have taller stems than those which are
more crowded together, and the amount of moisture in the soil greatly
modifies the development of the straw.

The lengths of the several internodes increase from the base to apex
of the straw, the uppermost internode being always the longest in plants
which have received no check in their growth.

From measurements upon Rye and other cereals, Nowacki concluded
that the length of each internode upon a normally grown and fully de-
veloped straw is the arithmetic mean of the lengths of the internodes
immediately above and below it. It is possible to find individual stems
whose measurements closely approximate to this formula, especially
among short-strawed varieties of wheat, but Nowacki's law is not generally
true, neither in respect of the straws of an ordinary field crop nor of those
of plants grown a foot or more apart from each other.

Below are given the average lengths of the internodes of long and
short straws of a variety of T. vulgar e and one of T. turgidum.

AVERAGE LENGTHS OF INTERNODES (cm.)

Internodes.

i.

2.

3-

4-

5- 6.

Total Length
of Straw.

RED SQUAREHEAD (T. vulgare)

Long straws —

cm.

cm.

cm.

cm.

cm.

cm.

cm.

Found
Calculated .

3-6

8-9
7'9

12-3
14-0

19-2

21-3

30'3
34'5

49-8

124-1

Short straws —

Found
Calculated .

2-0

4-6

5'3

8-6
9'5

14-4
16-3

24-0

25-2

36-0

89-6

BLUE CONE (T. '.urgidum)

Long straws —

Found
Calculated .

3'4

*'4
6-9

10-4

II-O

15-6
19-9

29-4

37-8

60-0

125-2

Short straws —

Found
Calculated .

2-1

3-6
4-2

6-4
6-7

9-9
12-8

19-2
24-4

39-0

80-2

64

THE WHEAT PLANT

The diameter of the straw is influenced by the same numerous factors
as those which affect the length ; especially stout stems are generally
found on plants which are grown at wider intervals than one foot apart.

The average diameter of the separate internodes increases from below
up to the fifth ; the sixth or upper internode is more slender than the
rest. Each individual internode is thickest in the middle, from which
point it tapers more or less evenly to both ends.

The average thickness of the wall of the straw decreases from the base
to the apex ; similarly, the thickness of the wall of each internode is
greatest at the base and diminishes gradually from below upwards.

The following are measurements of the diameters of the internodes,
the thickness of the wall, and thickness of the zone of hypoderm of hollow
straws of T. vulgar e (Squarehead) taken from an ordinary field crop.

Internode.

Average
Diameter.

Diameter.

Thickness
of Wall.

Average.

Thickness
of Hypoderm.

mm.

mm.

It,

M-

6th (upper)

3-03

top 2-40

412

440

115-150

middle 3-85 380

"5

base 2-85 528

150

Sth . .

4-18

top 3-90

445

5°7

115-150

middle 5-05

500

85-H5

base 3-60

577

85-15°

4th .

4^5

top 3'95

460 570

85-H5

middle 4-65

5J°

85-100

base 3-85

740

85-15°

3rd .

3-88

top 3-62

528

604

85-"5

middle 4-20

545

85-100

base 3-82

740

85-15°

and .

3-56

top 3-50

577

642

85

middle 3-65

610

85-115

base 3-55 i 740

ISt

3'43

3'43

74°

740

170

' TILLERING "

The primary stem of the wheat embryo within the grain is extremely
short, and consists of hypocotyl, or portion below the point of insertion
of the scutellum, and the epicotyl, or axis of the plumule.

When growth commences in grains sown in the ordinary way in the
field, the hypocotyl remains short, but the lower part of the epicotyl

THE STEMS 65

lengthens into a thin erect rhizome which pushes the terminal bud up-
wards through the soil.

This rhizomatous portion of the stem is from -6 mm. to i mm. in
diameter, much thinner than the aerial straws of the plant, and is usually
the second internode of the epicotyl. When grains are sown at a depth
of four inches or more, the second and third and sometimes other inter-
nodes above these often lengthen to form part of the rhizome (Fig. 61) :
at depths of three inches or less, the rhizome usually consists of the second
internode only.

In plants grown from grains sown three inches deep, the first, second,

r—,

FIG. 59. — Young plants with shoots at the " tillering " node, o, Scutellum ; r, seminal
roots ; r' and a (in 2), adventitious roots ; x, rhizome ; a (in i), shoot in axil of cole-
optile ; c, shrivelled coleoptile ; n, prophyll of first shoot; 5, leaf-blade of ist leaf;
T> T> f> and ^, ist, and, 3rd, and 4th leaves of primary axis ; p, prophyll of first shoot.

and third axillary buds of the main stem remain dormant or give rise to
weak shoots, the fourth lateral bud being the first to produce a strong shoot.

The third axillary 'bud is generally the first to grow strongly when
grains are deposited two inches in the soil, the second developing most
rapidly when the grain is sown one and a half inches down.

The bud in the axil of the first foliage leaf grows out immediately
when the grain is placed nearer the surface of the soil.

The length of the rhizome varies with the depth at which the grains
are deposited. When the latter are placed on the surface, or less than an

F

66

THE WHEAT PLANT

inch below, the rhizome is scarcely visible, being rarely more than two or
three millimetres in length ; at greater depths it lengthens proportion-
ately in such a manner that the plumule of the young plant produced
from a grain sown at any depth between one and three inches is lifted up
to within an inch or less of the surface (Fig. 60).

When the primary bud has reached this point, the rhizomatous inter-
node ceases to lengthen, and the succeeding internodes of the plumule
remain short.

During autumn and spring, however, the primary axis continues to

123 456

FIG. 60. — Wheat seedlings from grains sown at depths of 1-6 inches. Note extent of
tillering, length of rhizome, and number and length of adventitious roots from the
first (tillering) node. (About half natural size.)

grow very slowly, and, at the same time, in the axils of its leaves buds are
formed which expand into short secondary stems ; the latter also bear axil-
lary buds which are capable of developing in a similar fashion into branches
of the third order, and so on. Thus, from the primary bud of a single
wheat grain, a large number of stems may be produced, which remain very
short until April, at which date they usually begin to expand, the strongest
of them ultimately growing out into straws each with its terminal ear.

The production of these numerous shoots with unexpanded inter-
nodes, which takes place near the surface of the soil, is known as the
" stooling " or " tillering " of the plant ; it is the normal process of
branch formation in cereals and grasses generally.

The depth at which " tillering " or " stooling " begins, that is the

THE STEMS

point at which the plumule is arrested in its upward growth through the
soil, is regulated chiefly by the light-perception of the plant, and by the
depth at which the grain is sown.

In shady places the tillering node is found nearer the surface than in
sunny open ground, and a continuance of dull weather after late sowing
tends to shallow tillering.

The following measurements obtained from plants derived from
grains sown at carefully controlled depths illustrate the influence of depth
of sowing on the length of the rhizome, and the depth of the tillering node
below the surface.

Depth
of Sowing.

Length
of Rhizome.

Depth
of Tillering Node.

inches.

inches.

inches.

•5

•I

•4

i

•5

•5

i-S

2

3

I-O

1-6

2-2

•5
'4
•8

The arrangement and order of development of the several branches
may be followed by the dissection of young plants
in different stages of growth.

In the resting embryo within the grain, the
terminal primary bud is visible with its axis and
two or three rudimentary leaves ; the first lateral
bud is also seen in longitudinal sections in the
axil of the coleoptile (Fig. 10).

Ten or fifteen days after germination com-
mences, more lateral buds are formed, one in
each axil of the alternate leaves on opposite sides
of the primary axis (Fig. 62). Expansion of the
latter soon commences ; its first internode,
namely, that below the first lateral bud in the
axil of the coleoptile, always remains very short ;
the second, however, grows considerably, forming
a more or less elongated rhizome as already
described, the terminal bud with its lateral buds
being pushed upwards in the soil to the tillering
point.

After the plumule has been carried upwards

to this position, the buds in the axils of its two ,

, /.. . FIG. 61. — Rhizome of three

lowest roliage leaves develop rapidly, the plant internodes.

68

THE WHEAT PLANT

n

then showing three green shoots above ground, one on each side of the
main stem, arranged in alternate order, i.e. with a divergence of \ or
1 80°. Two more shoots soon spring from the primary stem above the
first pair, and about the same time each of the two first branches give rise
to two alternate branches of the second order arranged in a plane at right
angles to that of the first pair, the plant then consisting of nine shoots,

the relative position of which
can be easily recognised in
upright growing spring
forms of wheat : in the winter
types, however, which for a
time are decumbent, the
shoots soon bend out of the
planes in which they arise
and appear arranged more
$ or less in a semi-circle close
to the ground : the arrange-
ment of the first nine axes
'O of the plant is indicated
in Fig. 63. Additional
branches may arise later, not
only on the primary axis,
but on stems of the second,
third, and higher order, the
plant ultimately consisting
of a very large number of
crowded leafy shoots, with
short unexpanded inter-
nodes, the whole forming a
FIG. 62. — Longitudinal section through a young plant , . r, .1 • ...

14 days old ( x 25). *, Scutellum ; a, coleoptile ; dense tutt> the Visible parts
b, bud in the axil of the coleoptile ; c, bud in the being, of course, the blades
axil of the first foliage leaf ; n, bud in the axil of the r if i • i_

second foliage leaf ; *, terminal bud ; r, adventitious « the leaves With portions

root- of their sheaths, the axes or

stems from which they spring being short and hidden from view.

Schoute's method of indicating the various axes or branches of the
plant is convenient. The primary axis is denoted by the figure i, its ist,
2nd, 3rd . . . branches by the figures n, 12, 13 . . . respectively.

Similarly, the figures i, 2, 3 ... may be joined to these to indicate
the ist, 2nd, 3rd . . . branches of these axes : e.g. 132 denotes the 2nd
branch of the 3rd branch of the primary axis (Fig. 64).

The leaves on the several axes may be expressed in the form of a
fraction, the denominator of which denotes the axis, the numerator re-
ferring to the number of the leaf on that axis : e.g. -?- indicates the 2nd

THE STEMS

69

leaf of the primary axis, T2T the 2nd leaf on the 2nd branch of the first
axis, and so on (Fig. 59).

Very distinct differences in the habit of growth are visible in autumn
and early spring among young wheat plants of different varieties. Two
extreme types are readily recognised, namely, (i) the erect type (Fig. 65),
with shoots which spring up almost vertically, and (2) the prostrate type,
whose leafy shoots lie on the surface of the soil ; between these is (3) an
intermediate type (A and B, Fig. 66), with shoots which grow up at a variable
angle with the horizon. The three types may be indicated by the symbols
| , / , — : these characters are constant and are easily distinguished
except when the plants are abnormally crowded and " tillering " is checked.

A

FIG. 63. — A, Diagram of the arrangement of the axes of a young plant; B, relative
position of the axes later ; s, scutellum ; c, coleoptile ; /, first foliage leaf; p,p', prophylls.

In those of erect habit the young shoots form a somewhat compact
tuft, and the culms of the mature plant converge at the base to a narrow
point just below ground, resembling the ribs of a nearly closed umbrella
(A, Fig. 67) : they are very liable to lodge and are easily pulled out of the
soil. The tendency to grow in this manner is sometimes seen in plants
with only two leaf-blades developed, the first blade then making but a
narrow angle with the second.

In plants of the prostrate habit the first leaf-blade becomes horizontal
soon after the second blade appears. Later, the several shoots of the young
plant curve away from each other and soon come to lie close to the surface

THE WHEAT PLANT

of the soil, the strongly curved parts at this stage being the short leaf-
sheaths (A, Fig. 66). The extreme forms of this type are sometimes
called by farmers " creeping wheats." The culms of the fully developed
plant are bent at their bases, the whole being arranged in the form of a
cup (B, Fig. 67). Wheats with this habit
do not easily lodge and are so firmly
rooted in the ground that they are difficult
to pull up.

In wheat plants of an ordinary field
crop the development of the shoots does
not always proceed with the regularity
described above. The first lateral bud,

a

FIG. 64. — Diagrams of the arrangement
of the culms of tillered plants.

FlG. 65. — Young plant with
erect habit.

namely that in the axil of the coleoptile, generally remains dormant
or dies off altogether ; occasionally it develops and remains close to the
hypoctyl : the internode above it, carrying the growing point and lateral
buds upwards to the tillering region, the first lateral shoot in such an
example appearing to arise from the base of the rhizome some distance
below the rest of the shoots which spring later from the tillering nodes
(a (i), Fig. 59). Many later buds fail to develop, and the branching and
distribution of the shoots are therefore not so symmetrical as would be
the case if the buds in the axils of every leaf grew out in the order of its

THE STEMS 71

production. Moreover, crowding occurs, and the tillering process is often
checked after the formation of a comparatively small number of shoots,
and plants with two or three stems only are frequent.

B

FIG. 66. — Young plants. A, Prostrate habit ; B, semi-erect.

A strongly tillered plant, however, grown in an isolated position from
a single grain sown early in good soil, may possess a hundred shoots or
more in spring, which may develop ears yielding several thousands of
grains at harvest.

THE WHEAT PLANT

Although the buds and branches originating during the tillering
period may, under favourable conditions, amount to a very large number,
unless the plants have ample space for development and the soil is in a
highly fertile condition, many of them are very weak and grow out into
slender straws with small ears. In the majority of instances all the
leafy " tillers " of a plant produce ear-bearing straws, but sometimes in
an ordinary crop, where the individual plants " tiller " very little, some
shoots die off even after they have reached a height of 3-6 inches. This is
most frequently observed in dry seasons when the adventitious root-system
associated with each shoot and essential to its nutrition has developed too
late or has dried up before obtaining an adequate hold of the soil.

The extent to which tillering proceeds among the plants of a field
crop of wheat is comparatively small, the majority of the grains rarely
producing more than three or four fully developed straws.

Schoute found that out of 2895 plants examined in a Dutch crop

Relative Frequency.

•208

•no

•027

•0091

•0037

•0017

•ooio

•0014

•0003

•0003
•0003

63-7 per cent, or about two-thirds of the plants, gave one straw only,
while 95-4 per cent yielded not more than three straws each.

The average number of straws per plant produced by 841 varieties,
including representatives of T. dicoccum, T. turgidum, T. durum, and T.
vulgare, grown in 1912-13 at Reading in rows 6 inches apart, the grains
6 inches asunder in the rows, was 5-4.

7 varieties produced 1-2 straws per plant.

1845 plants produced only i straw
60 1 2 straws

378

3
4

27
n

6

5
3

8

4

9

0

10

i

n

0

12

I

13

I

H

34
137
163

183

J33

74

5°

22

38

2-3

3-4
4-5
5-6
6-7
7-8
8-9
9-10
over 10

THE STEMS 73

While it is the rule that, under the conditions which prevail in fields
receiving customary cultivation, each grain sown yields only a very few
straws, there are nevertheless numerous records of very prolific plants
arising from single grains sown in open rich ground and freed from
competition with neighbouring plants. Patrick Shirreff mentions one
with sixty- three ears containing 2473 grains, and another with eighty
ears yielding 4522 grains ; Haberlandt also refers to one bearing 130 ears
which produced 6855 grains weighing 218 grams.

By dividing a tillered plant into separate parts, each with its ad-
ventitious roots, and transplanting the detached shoots into good ground,
a very large number of ears and grains may be obtained in one season from
a single grain.

C. Miller gives an account of the result of an experiment illustrating
this method of increase. He states that he sowed a grain of wheat on
June 2, 1765, and on August 8 lifted the plant and divided it into
eighteen parts ; the several shoots were then planted out singly and the
division and transplanting repeated in September and October of the
same year, sixty-seven plants being obtained. A third division and
planting was carried out between the middle of March and the i2th of /
April in the succeeding year, producing 500 plants which at harvest
yielded 21,109 ears and 47 Ibs. 7 oz. of grain, the estimated number of
grains deduced by calculation after counting and weighing a certain
number being 576,840.

Incidentally, it may be noted here that as a process for the multiplica-
tion of new varieties of wheat, Miller's system of division and transplanting
does not lead to such rapid increase as the practice of thin seeding ; for,
as Shirreff points out, the single grain which Miller used was sown early
in June, and must have been obtained from the previous year's harvest,
two years being required, therefore, to produce one crop. On the other
hand, the plant mentioned by Shirreff, which from one grain produced
2473 in the first year, would yield 6,115,729 grains at the harvest of the
second, or more than ten times as many as Miller's plant, if those obtained
in the first season were each as productive in the second as the initial
grain.

The extent of tillering which wheats exhibit is governed by internal
physiological peculiarities of the plants, and also by various external
causes.

Most varieties of T. durum , and many of T. turgidum tiller very little,
while plants of T. monococcum, T. dtcoccum, T. Spelta, and T. compactum
usually give rise to a number of straws ; even varieties of the same race,
very similar to each other in form of ear and other morphological char-
acters, often differ in branching and in stem production. Late forms
usually tiller well, early forms tiller very little. The rapid-growing spring

74 THE WHEAT PLANT

varieties of T. vulgare with slender straws tiller more than the thicker
strawed winter wheats.

Tillering capacity is not only dependent upon the inherent vital
constitution of the species or variety concerned ; it is also greatly
influenced by internal qualities associated with difference in the size of
the grain from which the plants are raised. In experiments carried out
upon large and small grains which were sown in rows six inches apart,
and three inches asunder in the row, I found that large grains selected
from the ears of sixteen varieties of T. vulgare gave plants which pro-
duced an average of 8-1 stems per plant, while the small grains taken
from the same ears gave plants yielding only 4-4 straws each. Even
when the grains were grown at wider intervals, namely one foot apart
each way, the plants from the large grains produced an average of 14
straws each, those from the smaller grains only 10-6 per plant.

Much of the increased luxuriance of the plants derived from large
grains is doubtless to be attributed to the abundant food supply in the
endosperm of such grains ; some of it, however, may be the result of
specific biological differences in the embryos of large as compared with
those of small grains, although these differences are probably not of much
magnitude in grains selected from the same ear. Nevertheless, the parti-
cular share in the variation of the tillering power which should be assigned
to each of these two causes can only be determined by further experiment.

In addition to inherent qualities of the plants, other factors are con-
cerned in the tillering process. Profuse tillering is an index of natural
exuberant growth, and although the number, size, and vigour of the shoots
produced rests to some extent upon the hereditary qualities of the species,
variety, or form of wheat concerned, the branching and stem production
attained by any particular kind is due in largest measure to the action of
those external conditions which check or assist vegetative growth generally.
The question is one chiefly concerned with nutrition, and the tillering of
individual plants raised from grains taken from the same ear or spikelet
depends to a greater or lesser degree upon those external conditions which
favour or retard their nutritive functions.

The temperature of the soil and air, the amount and distribution of
the rainfall, the intensity of the sunlight, and other climatic factors
connoted by the term " season " have a controlling influence upon the
tillering process. It is also greatly affected by the physical and chemical
condition of the soil, the density of " seeding " or the amount of space
allotted to each plant, and the date at which the grain is sown.

For the most vigorous growth of the wheat plant comparatively dry
conditions are essential ; excessive moisture of the soil or atmosphere
greatly checks it and reduces tillering, especially when associated with low
temperature as in winter.

THE STEMS

75

The effect of the rainfall upon the number of straws produced by
the plant is strikingly exhibited in the results of some experiments
carried out during three years in order to determine the differences
between autumn- and spring-sown crops.

Nineteen varieties were sown in October, and again in February, in
the seasons 1910-11, 1911-12, and 1912-13, the plants being grown in
rows six inches apart, the individual grain being deposited six inches
asunder in the rows.

The average tillering power and the rainfall are given below :

1910-11.

1911-12.

1912-13.

Rainfall—

inches.

inches.

inches.

Oct. to March .

14-2

19-6

127

Feb. to March .

2-9

5'0

3'2

Number of straws per plant —

Autumn-sown

5'0

2-y

5'4

Spring-sown

6-5

2'5

6-0

The heavy rainfall in the winter of 1911-12 reduced the straws per
plant to about half the number obtained in the previous and succeeding
drier seasons, the effect being seen not only in the autumn- but in the
spring-sown crops.

It was found also that grains sown in spring very frequently produced
more straws and ears, though always of less average weight than those of
the same variety sown some three or four months earlier in the previous
autumn, the increase being most marked in the rapid-growing spring
wheats of normally short vegetative period.

The table on the following page gives the number of straws and ears
per plant produced by five typical winter and four spring wheats, sown in
autumn (Oct.) and spring (Feb.) respectively, during three successive seasons.

Tillering only continues so long as the temperature is above the
minimum for growth, and is most rapid at the optimum temperature :
it almost ceases in mid- winter, but goes on to an almost equal extent in
autumn and spring.

The date of sowing also influences tillering, since the degree of warmth
and the amount of light available for the plant as well as the length of time
for assimilation and vegetative growth are all correlated with it. Wheat
sown early in autumn branches more than it does when sown later,
although in most seasons the difference in the number of straws per plant
is comparatively small among crops drilled at any time between late
October and the end of January. It is in plants produced from grains

THE WHEAT PLANT

1910-11.

1911-12.

1912-13.

Sown in

Sown in

Sown in

Autumn.

Spring.

Autumn.

Spring.

Autumn.

Spring.

WINTER VARIETIES

Red Squarehead
White Squarehead .
Trump ....
Aleph ....
Hunter's White

Average

5'4
4.4

6-4

7-0
4-0

5-2

P

6-4
7-9

5'5

2-2
2-0

2-4
2-4

2-6

1-6
1-6
i-i
1-8
2-8

6-0

3'4
4-8
4-8
4-6

6-3

5'3
57
6-4

57

5-4 6-0

2-3

1-8

47

5'9

SPRING VARIETIES

Calcutta .
March ....

4-4
6-6

7-2
8-0

1-7

2-0

2-8
1-8

6-i

10-0

Manitoba Fife.
Californian

Average

5-6
5'0

77
9'5

2-6
3-0

2-6
5'2

5-8
6-2

8-0
7-0

5'4

8-1

2-3

3'i

6-0

8-3

inches.

inches.

inches.

Rainfall (Oct.-Mar.) .

14-2

19-6

127

sown out of the ordinary English season, namely in April, May, or June,
that the greatest effect of time of sowing is seen.

Winter wheats, or varieties with a long vegetative period, sown at any
date after the end of April, and spring forms of short growing period,
sown after the middle of June, go on tillering throughout the year in
which they are sown into the next before sending up ear-bearing straws,
the number of short unexpanded shoots or " tillers " produced diminish-
ing more or less regularly as the date of sowing advances from the periods
named.

In a series of experiments, in which grains of four varieties of wheat
were sown at the University College Farm, Reading, every week through-
out the year, the following results, as shown on page 77, were obtained.

Another factor which very materially modifies the stem and ear pro-
duction in wheat is the space allotted to the individual plants of the crop.

Extensive tillering can only take place when each has ample room for
its development ; the roots especially must be free from competition with
the roots of other plants, and allowed to permeate the soil in all directions
without hindrance, if maximum tillering is to be attained.

Where the crop is crowded, owing to thick seeding of the grain,
branching is much reduced ; on the other hand, it is increased by thin

THE STEMS
AVERAGE NUMBER OF STRAWS AND EARS PER PLANT

77

Sown in

Winter Varieties.

Spring Varieties.

Golden Drop.

Blue Cone.

Fife.

March.

1913. April .

20

32

..

May .

14

16-4

June

II

I3-5

12-5

15-0

July . .

II

28-0

18-0

16-6

Aug. .

II'O

11-3

Sept. .

7-9

10-3

o

n-6

Oct. .

7-3

%J

97

Nov. .

77

8-6

9-9

7-1

Dec. .

6-6

12-6

9-1

97

1914. Jan.

5'5

8-0

6-6

8-8

Feb. .

4'5

6-7

7'5

4-9

1912. March .

4'3

7'4

7-0

77

April .

3-8

5'3 .

seeding. In the case of good land, especially where it is well supplied
with available plant food-materials, thin seeding often results in the pro-
duction of the same number of straws per square yard as that obtained on
a thickly sown field, the tillering process making good the loss in plants
which thin seeding entails.

Where the land is in poor condition and the grain thinly sown, how-
ever, tillering does not make up for the loss of plants, and there is reduc-
tion in the number of straws per acre.

The following results were obtained in 1914 with a dense-eared
variety of wheat sown in the previous autumn on poor soil, which had
carried a crop of wheat during the three previous seasons, without the
application of manures of any kind :

No. of Plants Area allotted to
grown. each Plant.

Average No. of
Straws per Plant.

square inches.

600

6 (6" xi")

1-6

350

18 (6"x3")

i'S

200

36 (6"x6")

r8

200

72 (12" x 6")

2-9

IOO

144 (12" x 12")

4-0

60

576 (24" x 24")

7-0

On soil in such poor condition as this, where the rows were six inches
apart, the number of straws produced by each plant was about the same,
whether they stood i, 3, or 6 inches asunder ; only when the rows were

78 THE WHEAT PLANT

12 or 24 inches apart was there any considerable increase in the tillering
of the plants.

On soil in a high state of fertility the number of straws produced by
each plant is usually twice or three times as large as these.

" LODGING "

On examining fields of cereals at harvest the crops are not infre-
quently found to be more or less " lodged " or laid, the straw being bent
down to the ground, usually, though not always, because the lower inter-
nodes are too weak to support the upper leafy portion and the heavy ears.

This " lodging " or " laying " of the crop is of serious importance to
the farmer, for it reduces the yield of grain, the ears of laid crops usually
having many of their upper and lower spikelets barren or holding only
poorly developed grains. Moreover, the straw and ears are discoloured ;
there is also much likelihood of damage to the grain by sprouting, especi-
ally in a damp season, and the labour and expense of harvesting is greatly
increased by it.

Crops may go down at any time after the ear is out of the upper leaf-
sheath, though it occurs most frequently, perhaps, in the latter half of
June or a little later, when the plants are leafy and the grain is filling
rapidly. So long as the straw is soft and green, and the cells of the nodes
and thick basal portions of the leaf-sheath retain their vitality, the ears of
a laid crop may to a greater or lesser extent regain their upright position
through geotropic stimulus, the straws being bent upwards at each node
until the uppermost internode is again vertical. Where " lodging "
occurs early in the season the crop may therefore suffer comparatively
little ; if, however, it takes place late in the season when the nodal tissue
is dying or dead, the straw and ears remain where they are laid.

Crops go down usually after heavy rain or violent winds, but the
damage is only indirectly caused by these agents, the trouble being due
to (i) specific weakness of the straw, associated in some cases with a
natural drooping habit, and (2) to weakness of the roots, or to a root-
system which has little grip on the soil. Wind and rain only reveal the
weaknesses of the stems and roots of the crop but do not cause them.
Where the straw is weak, the lower internodes are usually bent or broken
when " lodged " ; where the root only is at fault, the plant goes down as
a whole, the straw being rigid and straight.

Some varieties of wheat, especially the dense-eared forms of T.
vulgare, possess short stiff erect straws, which under normal conditions
of growth do not easily " lodge." On the other hand, many Indian and
Persian forms of T. vulgar e have equally short but slender straws which,
when the ears are ripe, assume a drooping position, and although the
stems are strong and elastic enough to keep the ears off the ground in

THE STEMS

79

dry warm seasons, they readily become laid after a heavy shower when
grown at Reading.

Some tall lax-eared European forms of T. vulgare and varieties of T.
durum and T. turgidum with slender straw exhibit this drooping habit in a
slight degree, and are correspondingly liable to go down under conditions
of weather which would not damage the stiffer erect straws of many
European winter wheats.

The peculiar arrangement of the root-system and the strength of the
individual roots, which are hereditary characters of different varieties of
wheat, have a great influence upon the " lodging " of the crop.

In short-strawed win-
ter wheats of the Square-
head type the bases of
the straws just above the
ground bend outwards in
the form of a cup, and
from their lower nodes
arises a spreading system
of adventitious roots (B,
Fig. 67) ; the first inch
or two of the roots below
the surface is somewhat
rigid and thickened con-
siderably, and the cell
walls of their tissues
strongly lignified. By this
spreading arrangement of
strong roots the plants
are firmly anchored to
the soil and prevented
from being laid except by
the severest storms.

In most spring forms,
however, the straws grow
up from the ground in a crowded more or less compact bundle, spreading
very little, and the adventitious root-system consists of much thinner, less
lignified roots closely contracted and descending almost vertically with
very little grip on the surface soil (A, Fig. 67) ; plants with these root char-
acters are very easily bent to the ground as a whole, although the straw
may be as strong and rigid as that of the best winter varieties. " Lodging "
of this kind, which may even occur among isolated well-grown plants, is due
to weak root-hold, the straw being neither bent nor broken ; it is frequently
observed among rapid-growing oriental wheats when grown at Reading.

FIG. 67. — A, Stems and roots of a " spring " wheat ;
B, a " winter " wheat.

8o THE WHEAT PLANT

In addition to the hereditary characters of root and stem which in-
fluence the " lodging " of the crops, there are external factors which
encourage " lodging," and the phenomenon, as ordinarily encountered in
farm practice in this country, is most frequently associated with weakness
of the straw due to the crowding of the plants. The weakness is chiefly
an etiolation effect caused by insufficient light. In dense closely-grown
crops the retarding action of light on growth is absent ; lengthening of
the lower internodes and of their cells occurs, and thickening and lignifica-
tion of the mechanical tissues of the straw are greatly checked in dull light.

By comparing normal straws, which have had ample space for develop-
ment and adequate exposure to light during growth, with those of closely
grown crops, it is found that the lower internodes of the latter are some-
what longer, their diameter smaller, and their tissues somewhat softer
than those of plants grown wider apart. The vascular bundles in straws
of the same order, both in crowded and uncrowded plants, are similar in
number, but their transverse sections and the cells of the hypodermal
tissue are slightly smaller and the cell walls thinner in straws of crowded
plants than in those of normally grown stems ; both the hypoderm and
mechanical tissue round the vascular bundles are reduced in weak stems,
and their individual cells comparatively small and thin-walled ; in the
weakest crowded straws, which are sometimes little more than i -5-2 mm.
in diameter, the tissue outside the zone of vascular bundles up to the
epidermal layer consists of cells with delicate walls only.

The strength of the straws of " unlodged " plants is chiefly due to
the thick lignified walls of the hypoderm.

Crowding, which results in etiolation, to which the weakness of the
straw is due, may be brought about in many ways ; the sowing of too
much seed per acre, drilling in rows too close together, early sowing,
the application of large amounts of nitrogenous manures to the soil, a
mild winter with abundant rain, and other conditions which encourage
luxuriant growth may each produce it, and when several of these factors
work together " lodging " is almost certain to follow sooner or later.

Large leafy plants when crowded together, whatever the cause of their
luxuriance, are not only liable to " lodge " because the lower internodes
are weakened by being shaded, but they go down more quickly than
smaller plants because of the greater weight of their leaves and ears.

Experiments with various fertilisers on the wheat crop, carried on for
several seasons at the College Farm, showed that the application of
phosphatic and potassic fertilisers alone, even in very large doses, had
little or no effect on " lodging " ; on the other hand, plants grown with
large amounts of nitrates of sodium, potassium or calcium, and sulphate
of ammonia, invariably go down in crops of ordinary density because the
straw is weakened by the etiolation which occurs.

THE STEMS 81

Isolated plants grown with excess of nitrogenous manures frequently
possess strong rigid straw capable of effectually resisting the strain of wind
and rain ; they are, however, very liable to fall to the ground as a whole,
their anchoring adventitious roots, which in normally grown plants bind
the plant firmly to the soil and keep it upright, being soft and weak through
the slight development, reduced thickening, and imperfect lignification of
their mechanical tissues.

In common with stems of all grasses, wheat straw contains a con-
siderable amount of silica, namely, from 2-3 to 4-6 per cent in its dry
matter or 55 to 70 per cent in the ash ; it occurs chiefly in the cell walls
of the epidermis, to which it imparts hardness. The strength and
elasticity of straw was formerly attributed to the presence of this substance,
and the application of soluble silicates to the soil was suggested by Way
and others as a means of combating the tendency to " lodging." Pierre,
however, discovered that the straw of laid wheat was as rich or richer
in silica than that of standing crops, and his researches showed that the
lower internodes of straws, on which the resistance to lodging chiefly
depends, were always poorer in silica than the upper parts of the stem
and leaves.

The investigations of Lawes and Gilbert also demonstrated that a
high percentage of silica in the ash and dry matter of wheat straw is
associated with brittleness and weakness of the latter, and occurs in
seasons of bad harvests. Well-ripened straw of good seasons, possessing
strength and elasticity, contains a high amount of lignified tissues and a
correspondingly low proportion of silica.

The parasitic fungi, Ophiobolus graminis and O. herpotrichus , invade
the lowest internode of the wheat plant, often so weakening or destroying
its tissues that the straw falls to the ground.

Erysiphe graminis is also sometimes the cause of similar damage to
the wheat crop. Where the attack is severe, large numbers of plants
may go down ; the result, however, differs in appearance from true
examples of " lodging," as, in cases of injury by fungi, the individual
straws fall in all directions, whereas when laid by the wind or rain large
portions of the crop are bent over in the same direction.

" SHOOTING " OF THE STEMS AND ESCAPE OF THE EAR

Neither the primary axis nor any of the branches arising during the
tillering period continue to produce lateral vegetative shoots indefinitely,
for sooner or later inflorescences or ears originate at the apex of the
branches, after which lateral bud formation ceases in those particular
stems.

By an examination of longitudinal sections of very young shoots in

G

82

THE WHEAT PLANT

which ears can be detected, and also of full-grown culms or ear-bearing
straws, it is seen that no buds are formed in the axils of five to seven
leaves immediately below the ear itself.

In the ordinary course of events, where wheat is sown at any date
between October and the early part of March,
the young culms with their ears begin to ex-
pand in April. In all the shoots of the " til-
lered " plant in which the six leaf-blades of the
several culms are visible, both the straw and
ear can be readily recognised within the leaf-
sheaths, although their combined length is then
only about a quarter of
an inch (Fig. 68) . Growth
advances from this stage
onwards with increasing
rapidity until the straw is
2 or 3 feet high, and the
ear, almost full grown, is
pushed out of the upper-
most leaf-sheath into the

FIG. 68. — Longitudinal sec-
tion of a young plant at open air, usually at the
the end of 'April. «, Rhi- en(j of

n

zome ; i, 2, 3, ears.

June.

The crop in which the elongation of the culms
is taking place is said to be " shooting," the process
consisting essentially in the expansion of the stems,
which have originated in the previous " tillering "
period.

While growth in length in the young stem is
effected by the activity of an apical meristem, the
straw also possesses intercalary growing-points, one
at the base of each internode, which continue to
lengthen the several sections of the culm after the
upper part of each has ceased to grow ; these
delicate growing regions are protected by the
surrounding leaf-sheaths.

The expansion of the straw proceeds from below
upwards, the lowest internode being the first to
lengthen (Fig. 69), the growth of this being fol-
lowed more or less regularly by the succeeding internodes in numerical
order to the sixth or uppermost ; growth also ceases in the same order,
the lower internodes reaching their full stature some time before the
upper ones have arrived at their maximum length.

FIG. 69. — Longitudinal
section through a you ng
straw. Lengthening of
the internodes just be-
ginning ( x 5).

THE STEMS

In the first table below is given the length of the ear, and the separate
internodes of the straw from week to week, from the commencement
of the " shooting " period to the time at which the straw reached its
greatest height ; in the second table the increase in length from week to
week is recorded. As the stems are at first hidden by the surrounding leaf-
sheaths and cannot be examined when young without destroying the
plants, measurement of the growth of a single individual straw cannot
be carried out. The figures from April n to May 30 are averages
obtained from ten straws of similar size and development ; those from
June 6 onwards, however, refer to a single stem, the parts of which were
clearly visible and easily measured after this date.

TABLE I

LENGTH OF THE STRAW AND EAR OF A SQUAREHEAD WHEAT AT
WEEKLY INTERVALS FROM APRIL n TO JUNE 27, 1914

Whole
Internodes. ist. and. 3rd. 4th. 5th. 6th. Straw Ear'

•f
April ii . -25 -25 .. .. -2 -7 -15 cm.

N- J

„ 18

•35 '7 -2 ..

'2

1-4 '2

» 25

•3 2-0 -4

C3

3'0 '3

May 2

•4 2-6 1-4 -4 .

2 . 5-0 -4

9

1-3 5-5 4-0 -6 .

2 . n-6 -8

„ 16

3-4 8-1 11-3 2-6

5 . 25-9 2-0

» 23

3-4 8-5 12-7 6-6

9 • 32-i 3'8

» 30

3-4 8-5 15-3 18-0 .

8-2

3-8 57-2 7-8

June 6

4-2 . . 16-0 21-0

21-5

8-0 . . 8-2

.. 13

4-5 11-5 16-0 21-5

27-5

13-0 94-0 8-7

» 20

4-5 11-5 16-0 21-5

30-1

21'0 104-6 8'7

„ 27

4-5 11-5 16-2 21-5

30-3

21-5 105-5 9'0

July 17

4-5 11-5 16-2 21-5

30-3

21-5 105-5 9'0

Whole

Internodes.

ist. 2nd. 3rd. 4th.

5th. 6

, vv nuie „
th- Straw. Ear-

April 11-18

•05 .45 ..

•2

•7 -05 cr

n.

„ 18-25 •

•o 1-3 -2

•I

i-6 -i

„ 25-May 2

•i -6 i-o

•3 ••

2-0 -i

May 2-9

•9 2-9 2'6 -2

•o

6-6 -4

9-i6

2-1 2-6 7'3 2-0

'3

14-3 1-2

,, 16-23

•o -4 1-4 4'O

•4

6-2 1-8

,, 23-30 .

•o -o 2-6 11'4

II'I

25-1 4-0

,, 30- June 6

•8 .. -7 3-0

. 13-3 .. 4

•2 . . -4

June 6-13

•3 '3 -o '5

• 6'O .. 5

•0 I2-I -5

„ 13-20 .

•o -o -o -o

2-6 . . 8

'O 10-6 -o

,, 20-27

•0 -0 -2 -O

•2

•5 -9 -3

84 THE WHEAT PLANT

The measurements indicate a wave of growth from the base to the
summit of the stem. The greatest increase occurred in the second
internode during the first two weeks, the greatest growth being transferred
to the third internode in the fourth and fifth weeks, to the fourth in the
next fortnight, to the fifth in the succeeding two weeks, and to the sixth
in the last fortnight, at the end of which no further addition was made
to the length of the stem or the ear.

The figures in the vertical columns of Table II. also show that in each
internode and in the ear there is an acceleration to a maximum followed
by a gradual retardation in the rate of growth.

The whole period of " shooting " occupied about eleven weeks, the
greatest increase in length both of stem and ear occurring in the same week,
namely, May 23-30.

At the time of its escape from the upper leaf-sheath in the week ending
June 6 the ear had almost reached its maximum length, being then only
i cm. shorter than the ear when full grown.

The escape of the ear from the uppermost leaf-sheath marks a definite
and easily recognisable stage in the development of the wheat plant.

The date at which it occurs is dependent upon a number of factors,
the chief of which are :

1 . The race, variety, or form of the wheat.

2. The latitude and elevation of the place of growth.

3. The rainfall, temperature, light, and general climatic conditions of
the season of growth.

4. The date at which the seed is sown.

Some Persian, Indian, Chinese, and Australian forms sown in
September or October at Reading come into " ear " at the end of the
following May, while many English wheats sown at the same time do not
reach this stage of development until the second or third week in June.

The speed in attaining the " earing " stage, i.e. the " earliness " and
" lateness " of a variety, is a constant hereditary character which exhibits
the ordinary fluctuating variability, but is not permanently modified by
sowing the grain early or late or by varying the external conditions of
growth. Varieties preserve their characters in this respect no matter
where they are cultivated ; those which are relatively early or late in
India or Australia are early or late when grown in England, and rapid or
slow developing English forms exhibit the same relative differences when
transferred to another country.

Schiibeler and others have asserted that a " late " wheat can be trans-
formed into an " early " one and vice versa by varying the date of sowing or
by transference to another climate for a time ; there is little doubt that a
mixture of sorts was sown in such cases and the conclusions were illusory.

A " late " form is later in its ear production than an " early " form

THE STEMS

when the two are sown together, no matter where or in what season the
experiment is carried out, although, as mentioned hereafter (p. 91), the
time required by a wheat to reach the " earing " stage may be 300-320
days or only 60-70 days according to the time at which its grain is sown.

Each form of wheat sown on a particular date in England shows
remarkable regularity in the production of its ears : in normal seasons
the latter frequently appear on the same day in each year, and only in
years of drought or unusual rainfall is the time of their escape from the
leaf-sheaths accelerated or retarded more than a week or ten days. The con-
stancy of this character is of use in distinguishing closely allied forms, where
these can be grown together and observations made during several seasons.

For purposes of description and classification wheats may be grouped
into " early," " mid-season," and " late " forms, the former usually
exposing their ears at Reading before the end of May, the latter during the
second or third week in June.

In the following table is given the usual date of " earing " of over
1 500 wheats when sown in September or October at the University College
Farm, Reading :

S

s

s

.

§

1

^

3
|

.to

8

s

1

Q

8

|

o

i

S

o

I

i

1

1

<j
|

I

1

Total.

f
a

s
1

1

S

-s

*§

1

&

a

"s
1

4P

a
&

t

8

i

h!

^

tl"

E-;

hi

h

E-H'

^

Ei'

h

Early —

Very early. May 14-24 .

. .

2

15

33

5 6

6l

Early. „ 24-31 .

••

2

4

120

IO

4

211

5

••

3S6

Mid-season —

Mid-season. June 1-7 .

2

••

94

5°

3

528

24

••

701

Late —

Late. June 8-15 .

4

2

4

3

IO

59

. .

240

12

. .

344

Very late. „ 15-25 .

. . 2

i

• •

5

••

70

2

••

80

Total number of forms .

4 4

10

8

239

124

7

1082

48

6

1542

Although the date of " earing " of a wheat when sown at a particular
time is almost constant, the exit of the ear from the leaf-sheath may be
arranged to occur early or late over a fairly wide period of the summer
by altering the time of sowing. If, however, it does not take place before
the first week or ten days of July there is rarely sufficient time for the grain
to ripen at Reading (see pp. 90-92).

86

THE WHEAT PLANT

In order to determine the variation which can be induced in this
manner several wheats were sown at weekly intervals throughout the
year ; in the following tables are given the results obtained with an early
and one or two later forms sown in the first week of each month :

RED FIFE

Sown.

Ear out of Sheath.

Ear ripe.

Sowing
to
Earing.

Sowing
to
Ripe

Ears.

Earing
to
Ripe
Ears.

1912.

1913-

1914.

1912.

1913-

1912

days.

days.

days.

Feb. 24 .

June 15

Sept. 5

Ill

193

82

March 2 .

» 15

„ 5

104

1 86

82

April 6 .

July 4

*

88

May 4 .

„ 18

*

74

. .

June i .

Aug. 5

*

64

» IS •

,» 29

June i

*

. .

74

July 6 .

May 19

Aug. 8

317

398

8i

Aug. 3 .

„ 20

„ 8

290

370

80

Sept. 7 .

June 7

8

273

335

62

Oct. 5 •

. .

7

9

245

308

63

Nov. 2 .

„ 10

„ 9

220

280

60

Dec. 7 .

••

„ 12

„ 12

187

248

61

1913

Jan. 4 .

. .

,, 18

„ H

165

222

57

Feb. i .

„ 18

„ 18

i3S

198

63

March i .

. .

,, 20

. .

„ 20

in

172

61

April 5 .

July i

„ 3°

77

147

70

19 •

. .

» IO

Sept. 18

72

152

80

May 3 .

„ 13

61

» IO •

„ 15

54

June 7 .

Aug. 12

June 3f

66

July 5 •

Sept. 10

? t

. .

59

,, 12 .

May 25

317

Aug. 2 .

. .

„ 25

296

Sept. 6 .

?

259

Oct. 4 .

June 3

242

Nov. i .

» 8

. .

219

Dec. 6 .

••

„ 14

190

1914

Jan. 3 .

„ 16

164

Feb. 7 .

. .

,, 20

133

March 7 .

. .

„ 25

. .

no

April 4 .

. .

July 4

91

May 2 .

„ 14

73

„ 9 •

66

„ 16 .

63

••

Average . 69-4

* Ears were produced in July and August, but these did not ripen in 1912, the summer
months in that year being dull and wet.

t Ears were produced in August and September 1913, but did not ripen : the same
plants lived through the winter and produced ears again in 1914.

This form may come into ear as early as May 19 or not until September 10, and may
ripen its ear as early as August 8 or not until September 18. Ears which did not escape
from the sheaths before July 10 had not time to ripen in that season and died off.

THE STEMS

DAWSON'S GOLDEN CHAFF

Sown.

Ear out of Sheath.

Ear ripe.

Sowing
to
Earing.

Sowing
to
Ripe
Ears.

Earing
to
Ripe
Ears.

1912.

1913-

1914.

1912.

1913-

1912

days.

days.

days.

Feb. 24 .

June 29

. .

Sept. 5

125

193

68

March 2 .
April 6 .

July 4
Sept. 3

June 2

„ i?

July 26

123
148

I98
476

75

May 4 .
June i .

» 4

5

„ 26
„ 28

395
368

448
422

53
54

July 6 .

„ 5

„ 30

332

389

57

Aug. 3 .

„ 4

„ 3i

32§

362

59

Sept. 7 •

„ 4

„ 3i

268

327

59

Oct. 5 •

5

„ 3i

241

299

58

Nov. 2 .

„ 7

Aug. 6

215

277

62

Dec. 7 •

„ 21

„ 8

194

244

5°

1913

Jan. 4 .

,, 20

. .

„ 14

165

222

57

Feb. i .

.» 21

„ 20

136

200

64

March i .

„ 25

„ 25

116

177

61

8 .

July 2

Sept. 6

116

l82

66

April 5 .

Aug. 5

129

19 •

June 2

409

May 3 .

May 30

392

June 7 .

„ 30

357

July s •

June 2

332

Aug. 2 .

May 30

301

Sept. 6 .

June i

267

Oct. 4 .

„ 3

241

Nov. i .

„ 8

218

Dec. 6 .

„ ii

187

1914

Jan. 3 .

,, 12

1 60

Feb. 7 .

,, 20

133

28 .

July 2

March 7 .

„ 18

133

14 •

» 1°

••

118

Average . 60 • 2

This form may come into ear as early as May 30 or not until September 3, and may
ripen its ears as early as July 26 or not until September 6. Ears which did not escape
from the sheaths before the end of June or beginning of July had not time to ripen in that
season and died off.

88

THE WHEAT PLANT

SWAN

Sown.

Ears out of Sheath.

Ear ripe.
1913-

Sowing
to
Earing.

Sowing
to
Ripening.

Earing
to
Ripening.

I9I3-

1914.

1912

days.

days.

days.

Sept. 14

June 1 6

. .

Aug. 1 6

275

336

61

Oct. 5

„ 19

„ 16

257

315

58

Nov. 2

„ 18

. .

„ 18

228

289

61

Dec. 7

„ 21

„ 20

196

256

60

1913

Jan. 4

„ 29

„ 30

I76

238

62

Feb. i
March i

„ 3°
July i

Sept. 10

>, J2

149
122

221
195

72
73

„ 22

» J7

. .

„ 18

117

1 80

63

April 5

Aug. 10

*

127

. .

May 3

June 7

400

» 3

, 7

372

. .

June 28

> 7

. .

344

. .

1 1

Aug. 2

. .

, 12

Sept. 6

, 12

Oct. 4

. .

Nov. i

! 16

Dec. 6

> *9

1914

Jan. 3

. .

„ 24

. .

. .

. .

, f

17

„ 24

» 3^

„ 24

Average . 63-7

Did not ripen any ears in 1913.

THE STEMS

SWAN

Sown.

Ear out of Sheath.
1916.

Ripe.
1916.

1915

April i ...

June 14*

N

8 ...

» 14*

15

„ 13*

22

» *3 *

May i ••' .

N

15

June i ...

15 • •

July i ...

June 16-22

15 . . .

Aug. i ...

15
Sept. i

• Aug. 14-19

15 •

Oct. i ...

June 22

15

» 24

Nov. i ...

„ 24

15

» 24

Dec. i ...

„ 26

15

July 3

1916

Jan. i ...

». 4

15 • •

4

,

Feb. i ...

5

Aug. 23

15 ...

„ 18

Sept. 3

March i

„ 20

t

,,15 ...

„ 24

t

April i ...
15

I

* A few ears were produced in September and October 1915 by these plants, but
most in June of the following season 1916.

t Ears were produced late in September, but did not ripen, and died off in 1916.

J These plants formed dense leafy tufts in 191 6, from which were sent up a few culms
late in September and October.

Sown before the end of February, this form produced ears which ripened in the same
season : sown later than this, few or no ears were produced or ripened until the succeed-
ing season.

9o

THE WHEAT PLANT
BLUE CONE

Sown.

Ears out of Sheath.

Ears ripe.

Sowing
to
Earing.

Sowing
to
Ripening.

Earing
to
Ripening.

.9.3.

1914.

1912

days.

days.

days.

Oct. 12

June 21

Aug. 25

252

317

65

Nov. 2

„ 21

» 30

232

301

69

9

„ 21

Sept. 2

224

297

73

23

„ 30

. .

219

Dec. 7

Sept. 12

Jan. 4

?

12

25 1

Feb. i

?

. .

» 14

. ,

225

March i

?

„ 18

. .

2O I

. .

„ 15

July 20

. .

„ 20

189

April 5

Aug. 12

12

June 7

421

May 3

. .

6

399

. .

June 7

7

365

. .

July 5

ii

341

Aug. 2

12

. .

3H

. .

Sept. 6

. .

15

. .

282

. .

Oct. 4

. .

. .

256

. .

. .

Nov. i

. .

21

232

Dec. 6

23

199

1914

Jan. 3

» 24

165

Feb. 7

» 3°

Average . 69

This wheat may come into ear as early as June 7 or not until August 12, and may
ripen as early or earlier than August 25 or not until September 20.

All wheats sown at Reading in January, February, and March produce
ears in the same season ; if sowings are made at weekly intervals later,
there arrives a time which is too advanced to allow the growth of the plant ,
to proceed as far as the " shooting " of its ears in that year. In the case
of the early or rapid-growing " Fife " wheat this period is as late as July,
while for late, slow- developing forms, such as Swan and Dawson's Golden
Chaff, it is as early as the middle of April.

If sowing occurs after these critical periods, the plants grow and tiller
extensively but do not " shoot " ; they behave as if they had cognisance
of the want of time to carry their development to the " earing " stage in
that season, and remain short leafy tufts through the summer and succeed-

THE STEMS 91

ing winter, sending up their stems and ears in the following year very
little in advance of plants sown several months later.

The critical period is nearly constant for the same form ; there is
indeed a certain degree of irregularity in the development of the plants
sown about this date, some individuals sending up a few ears, while others
refuse to do so entirely ; sown a week earlier or later, however, the plants
exhibit a decision in their behaviour which is very striking.

For a time the young plants in successive rows grow and develop
apparently at the same rate, but ultimately the earlier sown rows send up
their stems and ears to a height of 3-4 feet, the neighbouring rows in which
the sowing has been delayed but a week or ten days remaining a few inches
high ; the contrast is remarkable in view of the small difference of time
between the sowings and the apparently similar climatic conditions under
which the plants are grown.

Fife wheat sown before the critical period — say in May — produces
ears in 60-70 days, whereas it does not come into ear until about 300
days have passed when sown at the end of July or beginning of August.
Similarly, later wheats may produce ears in 115-120 days, or not before
the lapse of 400 days according to the time of sowing.

Fife wheat sown on May 10, 1913, was in ear in 54 days, i.e. on July 13.

The late form, Dawson's Golden Chaff, came into ear most rapidly
when sown in March, the time between sowing and " earing " being 116
days.

When sown at the most favourable time certain wheats come into ear
in 6 or 7 weeks, but forms with such power of rapid development are rare.
A Chinese wheat from Newchwang sown on May 25, 1911, was fully in
ear on July 8, or 44 days after sowing, and another from Fu-chia Chuang
sown on May 18, 1916, came into ear on July 6, 49 days after sowing.

On the other hand, the latest forms do not come into ear in less than
150 to 1 60 days, even when sown at the most propitious date.

For each form of wheat there is a date before which ears are never
produced when grown in the open field no matter at what time the grain
is sown : it varies with the amount of light, mean temperature, and other
climatic conditions of the place of growth.

At Reading the earliest date at which wheats can be made to come
into ear is about the middle of May, and this is the case only with very
early Chinese and Indian forms sown in autumn of the previous year.

The latest period at which ears escape from the leaf-sheath at Reading
is some time during September or October, the exact date being subject
to much more fluctuation than the time of the earliest appearance of ears.
At such late season the " shooting " or expansion of the stems is very
irregular, and only a few ears of each plant are pushed out ; these die off
as the autumn advances and do not ripen grain.

THE WHEAT PLANT

During the cold autumn months wheat makes comparatively little
growth, although the difference between plants of the same form sown in
September and those sown in November, in respect of the number of
shoots which they possess, is sometimes considerable. So far as the
" shooting " and " earing " processes are concerned, they are evidently in
much the same state of development, or very little in advance of each
other, for observations of the growth of many varieties over several seasons
show that rows of the same wheat sown during September, October, or
November come into ear within 3-5 days of each other in the following
June ; also, those sown in January and February usually send forth their
ears nearly together, although later than those sown in autumn.

Sown in January or February they send forth their ears later, the
early sorts from 4 to 7 days, the later varieties as much as 10 to 20 days later
than the same kinds sown in autumn up to November in the previous
autumn.

ANATOMY OF THE CULM AND RHIZOME

A culm or straw is composed of the following tissues :
(i) The epidermis ; (2) the hypoderm or zone of mechanical tissue
beneath the epidermis ; (3) green assimilating parenchyma ; (4) colourless

ground parenchyma ; and
(5) the vascular bundles.

In transverse section the
straw is almost circular in
most Bread wheats (T. vul-
gar e), but in other races,
especially T. dicoccoides , T.
turgidum, and T. polonicum,
the outline is wavy, the sur-
face of the culm in these
being more or less deeply
furrowed.

(i) The epidermis, which
is about 25-30 /x thick, is
formed of narrow elongated
cells 150-250 //, long, with
short square cells 9-13 /x

FIG. 70. — Lines of stomata
of the culm ( x 50).

FIG. 71. — Epidermis
of the culm ( x 210).

across intercalated among
them at intervals . In wheats
with the furrowed straws, short papillae or scabrid hairs are often
present, particularly in the furrows. Single or double lines of stomata
similar to those found on the leaf-blade and sheath are present (Fig. 70), the
stomata being 200-250 ^ apart in the row. The cells over the hypoderm

THE STEMS

93

are about 4 fj, in diameter, those covering the green assimilating tissue
being wider (10-15 /A in diameter) with more sinuous walls.

The walls are all pitted and extensively thickened, and on the parts
not hidden by the leaf-sheath a cuticle is present, the surface of which
in the case of glaucous-stemmed varieties is covered with minute flaky
particles of wax.

(2) The hypoderm is a strong elastic cylinder of mechanical tissue,
consisting of lignified fibres with narrow lumina and strong walls about
4 fj, thick : in a transverse section of the straw it appears as a continuous
zone of cells immediately within the epidermis, and is of variable thick-
ness, being more or less wavy in outline on its inner side (Figs. 72, 73).

From the measurement of a full-grown straw of " Squarehead " wheat
given on p. 64, it is seen that the average thickness of the zone is about

FIG. 72. — Transverse section of portion of the culm (lower part of the upper internode)
( x 70). a, Small vascular bundle in the hypoderm (h) • b, large vascular bundle ; c,
band of chlorophyllous tissue.

the same in the 2nd, 3rd, 4th, and 5th internodes ; in the upper internode
it is slightly stronger, and reaches its greatest development in the lowest
section of- the straw.

A similar relationship in regard to the thickness of this zone of sup-
porting tissue is seen in each of the separate internodes, being stoutest in
the basal and thinnest in the middle portions.

(3) The assimilating tissue of the stem resembles that of the leaf,
consisting of delicate parenchymatous cells, which in longitudinal view
are irregular in form, but in transverse section are almost circular and
about 12 /x in diameter (c, Fig. 73). It is imbedded in the sclerenchyma-
tous hypoderm, but a portion of its outer face is always in immediate
contact with the epidermis, through the stomata, by which it is brought
into communication with the atmosphere.

The tissue is arranged in parallel bands, which appear as narrow green
strips along the stem, with lines of colourless stereome between. The

94 THE WHEAT PLANT

bands frequently run in pairs, which sometimes coalesce to form broader
C

FIG. 73. — Transverse section of hypoderm and vascular bundle (x38o).
c, Bands of chlorophyllous tissue.

ones. Each band is wide at the apex and tapers gradually as it is traced

FIG. 74. — Transverse section of large vascular bundle ( x 260).

downwards ; beneath the leaf-sheath it diminishes rapidly, and finally
disappears before it reaches the base of an internode.

The average width and thickness of each individual band is greatest

THE STEMS 95

in the upper internode, and the bands are closer together there, giving
this portion of the straw an almost uniformly green tint ; in the three
lowest internodes the tissue is only found in the upper parts, and is much
reduced, or entirely absent, at the bases.

(4) The colourless ground parenchyma extends from the hypoderm
to the centre or to the hollow pith-cavity found in most wheats. It is
composed of thin-walled, finely-pitted cells polygonal or rounded in section,
those bordering the stereome being long and narrow (350 ^ long, 35 p in
diameter), nearer the centre they are shorter and wider (150-250/7, long
and up to 100 /u, in diameter) ; they soon die and lose their cell-contents.

In the lowest internodes the walls of the parenchyma become thick-
ened and lignified ; the tissue then materially assists the hypodermal
stereome in strengthening the base of the straw. Very thick- walled ground
parenchyma is found also in the lower part of the solid diaphragm between
the internodes, forming a lining to the upper part of the hollow pith-cavity.

(5) In a transverse section through the hollow internode the vascular
bundles are found distributed symmetrically in the wall of the straw. Im-
bedded in the hypoderm is a ring of very small bundles separated from
each other by somewhat wide intervals, each bundle being placed between
two green lines of assimilating tissue (a, Fig. 72).

Nearer the centre, in the soft parenchyma, are the large bundles of the
stem, arranged more or less regularly in a ring, with an occasional bundle
of intermediate size close to the inner edge of the hypoderm.

The large bundles are of the usual closed collateral type, with the
xylem directed towards the centre of the stem, its chief vessels arranged
in the form of a V .

The protoxylem forming the apex of the V consists of one or two
annular or spiral vessels ; two pitted vessels with wider lumina are
situated on the right and left limbs of the V , the space between the chief
vessels being occupied by small tracheids.

The phloem of the bundle lies between the open limbs of the V and
extends slightly beyond it. It is composed of sieve-tubes with their
companion-cells.

Surrounding each large bundle is a sheath of fibrous stereome, which
in the case of the small hypodermal bundles is not differentiated from the
stereome in which they are imbedded.

The bundles of the leaf-sheath curve inwards into the node and pass
through the latter as separate isolated strands without fusing with each
other or with the neighbouring bundles which have come from the inter-
node above.

In the leaf-sheath itself all are of somewhat similar diameter, but in
traversing the node approximately half of them become very much thick-
ened, and alternate in more or less regular order with the other half,

96 THE WHEAT PLANT

which are reduced in diameter : the finer strands pass into the hypodermal
stereome ring of the stem, or close to it, the larger ones arranging them-
selves nearer the centre in the pith of the straw.

Of the 30 bundles in the leaf-sheath illustrated in I., Fig. 75, 13
(Nos. i, 3, 5, 7, 10, 12, 14, 16, 18, 20, 23, 27, and 29) go through the
node into the hypodermal stereome of the stem or near it (VI., Fig. 75),
the remaining larger strands (17) going into the pith.

The small unnumbered hypodermal bundles seen in the section of
the stem (VI., Fig. 75) arise within the stereome of the leaf-sheath bundles
a short distance above the node (x, Fig. 75).

In the internodes of the stem the bundles run longitudinally in parallel
lines ; traced downwards, they are found to enter the solid node. Just
below the point of junction of the leaf-sheath the fine hypoderm bundles
curve outwards, some of them at the same time spreading more or less
horizontally and uniting with each other to form a narrow more or less
broken ring (r, II., Fig. 75). A little lower the stouter, inner, larger
cauline bundles bend outwards and insinuate themselves between the
large leaf- traces, at the same time bifurcating or fusing with each other in
an irregular manner, some of the strands twisting and curling through the
central pith in various directions, others passing out into the cortical
parenchyma on the outer margin of the zone of large bundles.

The 37 " pith " bundles of the stem (I., Fig. 75) found above the
node are reduced by fusion and anastomosis within the node to 16,
which are seen (not numbered) in the section of the stem below the
node (VI., Fig. 75), alternating more or less regularly with the 17 large
bundles which have come from the leaf-sheath.

Generally in a section of an internode of a straw all the fine bundles
in the hypoderm and a few small bundles near it come from the leaf-
sheath immediately above ; of the larger bundles within the pith, about
one half come from the leaf-sheath, the other half descending through
the node from the internode above.

In a transverse section the bundles in the leaf-sheath appear of the
ordinary collateral type with girders of stereome connecting them with
the outer epidermis (2, Fig. 57). Very near the base of the sheath,
however, the stereome strands are free from the epidermis, each becoming
oval in section (i, Fig. 57) : imbedded within them are the rudiments
of minute bundles (usually one but sometimes two in each) the first traces
of which consist of two or three leptome cells.

On entering the node the stereome strand is reduced and finally
disappears : the minute bundle within the latter becomes more clearly
defined and can be traced through into the hypoderm of the stem below.

As previously mentioned, some of the leaf-sheath bundles on entering
the node are reduced in size, but about half of them are greatly increased

9 10

10 II l2

VI

G. 75. — SUCCESSIVE TRANSVERSE SECTIONS THROUGH THE LEAF-BASE AND NODE OF A CULM>
illustrating the course of the bundles through the node ( x 13-5).

II., -4 mm. below I. ; III., -4 mm. below II. ; IV., 1-4 mm. below III. ; V., -8 mm. below IV. ;

VI., 1-6 mm. below V.

THE STEMS 97

in diameter and form a conspicuous ring of strands, each of which is
flattened laterally, becoming oval or pear-shaped in section (Fig. 75).

Around the bundle, in its course through the node, is a sheath of
parenchyma containing many green chloroplasts.

The xylem elements of these large sheath bundles are much increased
within the node, the vessels being replaced by a great many short, pitted
and reticulately thickened tracheids of small diameter distributed
irregularly with xylem parenchyma.

In the lower section the xylem elements more or less completely
encircle the bast, but this amphi vasal arrangement is lost after the bundles
leave the node, and the collateral form is soon resumed.

In the internode above and below the node the cauline bundles are
of the ordinary collateral type, but within the node they are much
changed in structure ; the sheath of sclerenchyma on the pith side is
somewhat strengthened and the vessels become more numerous, the
typical V-shaped section being altered to a Y, the stem of which is formed
of 4 to 6 vessels of more or less elliptical section. Lower, the vessels on
the arms of the V and Y are replaced by groups of short tracheids.

The number of bundles in a straw varies considerably, stout straws
of plants grown wide apart possessing more than those of crowded
crops ; in the early developed stems of tillered plants the bundles are
more numerous than in straws formed later.

The total number is generally greatest in one of the two uppermost
internodes, but sometimes in the lowest, fewer being found in the inter-
mediate portion of the straw.

In the first four internodes there are usually more in the inner zone
of large bundles than in the hypoderm, but in the portion of the stem
immediately below the ear the reverse is occasionally the case.

On the following page are given the numbers of the vascular bundles
in the internode of the 6 straws of a plant of T. vulgare (Squarehead)
from a row of plants grown 3 inches asunder in rows i foot apart.

The thin rhizomatous portion of the stem between the grain and the
tillering nodes is about -6-1 mm. in diameter. Its anatomy is somewhat
different from that of the internodes above ground, there being a clearly
circumscribed stele or central cylinder surrounded by a soft thin-walled
cortex. The epidermis consists of elongated parenchymatous cells
about 7 n in diameter, with thin walls ; hairs and stomata are absent.
Within, is the cortex of thin-walled parenchyma, generally 6 to 8 cells
deep and about -i5--i6 mm. across ; there is no zone of hypoderm, but
occasionally one or two very fine strands of stereome are seen just beneath
the epidermis. A fairly well-defined endodermis is visible, differentiated
from the rest of the cortical tissue by its thick inner walls, which are
lignified and show lines of stratification.

H

THE WHEAT PLANT

No. of
Internode.

Length of
Internodr;
(Inches).

No. of Bundles.

Total.

No. of
Internode.

Length of
Internode
(Inches).

No. of Bundles.

Total.

Hypoderm
Ring.

Inner
Ring.

Hypoderm
Ring.

Inner
Ring.

STRAW i.

STRAW 2.

6th

18-5

38

34

72

6th

18-9

34

31

65

5th

11-3

34

43

77

5th

10-9

35

35

70

4th

6-2

28

44

72

4th

6-8 -

26

36

62

3rd

3'7

22

41

63

3fd

3-6

21

4i

62

2nd

2-9

22

36

58

2nd

2-6

18

38

56

ISt

i'3

3°

33

63

ISt

1-2

19

33

52

Total 43-9

Av. 34-8

38-5

67-3

Total 44-0

Av. 25-5

35'6

61-1

STRAW 3.

STRAW 4.

5th

18-8

35

32

67

5th

I7-7

35

3i

66

4th

10-5

35

34

69

4th

10-5

3i

34

65

3rd

4-6

25

38

63

3rd

4'9

24

35

59

2nd

2-7

20

38

58

2nd

2-6

19

35

54

ISt

i'4

22

3.6

58

ISt

I'S

23

34

57

Total 38-0

Av. 27-4

35-6

63

Total 37-2

Av. 26-4

33-8

60-2

STRAW 5.

STRAW 6.

5th

IS'?

32

30

62

5th

8-7

25

26

51

4th

8-7

29

32

61

4th

7'7

24

33

57

3rd

S'O

24

34

58

3rd

5'3

19

33

52

2nd

2-8

19

36

55

2nd

2-9

17

34

51

ISt

i-o

21

4i

62

ISt

•9

19

37

56

Total 31-2

Av. 35-0

34'6

59'6

Total 25-5

Av. 20-8

32-6

53'4

The following are the numbers of the vascular bundles in the leaf-
sheaths and internodes of a full-grown straw of T. vulgare (Squarehead)
taken from an ordinary field crop.

Leaf-sheaths.

Straw.

No. of
Bundles.

Internode.

Bundles in
Hypoderm Ring.

Bundles in
Inner Ring.

Total.

5th

35

5th
(upper)

top 24
middle 27
base 24

22
24
24

46

%

4th

38

4th

top 28
middle 28
base 29

31
31
31

59
59
60

3rd

25

3rd

top 1 8
middle 18
base 1 8

31

32
32

49
50
50

2nd

23

and

top 1 6
middle 16
base 1 6

34
34
34

50
50
5°

ISt

ISt

top 22

middle 22
base 22

34

37
37

56

59
59

THE STEMS 99

The stele from -5 to 6 mm. in diameter occupies the centre of the
rhizome, and near its periphery are the vascular bundles, generally 10-15
in number, distributed somewhat irregularly in one or two zones, the
larger bundles to the outside.

Each bundle is usually of the ordinary collateral type with V-shaped
xylem and a narrow sheath of sclerenchyma. Some of them have only
one large vessel ; in others, two vessels are present, these being irregular
in size and not arranged symmetrically. The centre of the cylinder,
though sometimes hollow, is more often filled with soft pith.

CHAPTER VII

THE INFLORESCENCE

THE inflorescence or ear of wheat is a distichous compound spike, the
primary axis bearing two opposite rows of lateral secondary spikelets

and a single fertile terminal spikelet, except in
the case of T, monococcum in which the latter is
rudimentary and barren, or missing.

The main axis or rachis is a sinuous notched
structure composed of a number of short inter-
nodes (Fig. 76), which in the *' Spelt" wheats
T. aegilopoides, T. monococcum, T. dicoccoides, and
T. dicoccum disarticulate easily at each node when
the ear is ripe (Fig. 77), and in T. Spelta breaks
transversely just below each spikelet (Fig. 207).
In the majority of cultivated wheats, however,
the rachis is tough and resists disarticulation or
breaking even when thrashed.

Each short segment of the axis is narrow at
the base and broader at the apex, one side of it
being more or less convex, the other flattened or
slightly concave. Its lateral edges are fringed
with hairs of variable length, and there is usually
a larger or smaller tuft of hairs — the frontal tuft
— at the apex of the convex side between the
two lowest glumes of the spikelet. All the spike-

FIG. 76.-Rachis of an ear. lets are sessile ' the lateral ones are ranked
a, Rachilla ; b, portion of alternately on opposite sides of the rachis, each
rachis with attached spike- bdng placed at the apex of an internode) witn

its broadside towards the flat or concave surface
of the segment immediately above its point of insertion.

The terminal spikelet is arranged at right angles to the rest.

Great variety is found among wheats in respect of the number of
spikelets per unit length of the rachis. In some, the spikelets are crowded
together, the lengths of the internodes between each being very short

THE INFLORESCENCE

and hidden from view ; in others, they are widely separated and the rachis
visible between the adjacent spikelets (cf. Figs.
188 and (i) 180).

In the majority of varieties the distribution
of the spikelets along the rachis is fairly uni-
form, but in some ears the spikelets are
especially crowded at the tip and arranged in
a lax, open manner towards the base (i, Fig.
189) ; such ears are spoken of as " clubbed,"
though the name " Club wheat " is used for
the race T. compactum, in which the ears are
very short, with spikelets closely set along
the entire length of the rachis.

Viewed from the side the spikelets stand
out from the rachis, making with the latter an
angle of 2O°-8o° or more, the smaller angle
being found in lax-eared forms, the larger in
dense compact ears.

Each spikelet consists of a delicate flat-
tened and jointed rachilla (a, Fig. 76) which
bears two opposite rows of alternate solitary
flowers hidden between chaffy bracts or
glumes. The glumes are either glabrous or
covered with soft hairs, the colour when ripe
being creamy white, yellow, red, brown, or black.

The number of flowers in a spikelet varies
from 3 to 9, of which one or more of the upper
ones are usually imperfect and sterile.

At the base of the spikelet are two opposite
rigid boat-shaped bracts, the empty glumes,
which in all wheats, except T. polonicum, are
shorter than the rest of the spikelet, and in
lateral spikelets the parts right and left of the
midrib are dissimilar in size and shape. Their
size, shape, texture, form of keel, and terminal
tooth are constant characters, of high taxo-
nomic value. In some wheats they are
keeled throughout their whole length, in
others the projecting keel extends only a short
distance from the apex, the lower portion
being rounded ; each possesses several nerves,
the central strongest one dividing the glume into two unequal halves,
of which the outer is the broader.

FIG. 77. — Disarticulation of the
rachis of a form of T. aegilo-
poides.

THE WHEAT PLANT

The form of the apex of the empty glume differs considerably in the
various kinds of wheat ; in some it is broad with a blunt extension of
the midrib, in others it is narrower with the central nerve continued into
an acute point ; in certain forms of T. vulgare and T. compactum from
Central Asia it possesses a terminal scabrid awn from 2 to 5 cm. long, but
in the majority of wheats the terminal tooth or keel is not more than a
few millimetres in length.

In T. aegilopoides and T. monococcum the empty glume has a lateral
secondary tooth on its broad outside half, in addition to the primary
central one ; in most other wheats the secondary tooth is short or
-altogether absent.

The empty glumes of the terminal differ from those of the lateral
spikelets ; they are rarely keeled and always symmetrical, a line down
the centre dividing them into two similar halves. In some wheats these

glumes have single, well-defined
midribs ; in others the midrib
is missing, in which case two
strong lateral veins are present,
one on each side of the central
line, the apex being notched or
divided sometimes to near the
base of the glume (Fig. 78).

The colour of the glumes is
white, red, or black, but of each
FIG. 78.— Empty glumes (T. turgidum) ( x 3). «f these tints there are many

a and c, from terminal spikelet ; b, from a
lateral spikelet.

shades. Whatever the natural
hereditary tint, its full develop-
ment is dependent largely upon the intensity of the light, heat, and atmo-
spheric moisture during ripening. In certain varieties of T. dicoccum, T.
durum, and T. turgidum the tint is a creamy white, but in the majority of
the white-chaffed varieties it is pale yellowish ; in a few Persian forms
the glumes are a clear canary-yellow colour, and occasionally pale emerald
green.

Among red-chaffed Wheats the range is very wide, extending from the
palest shades of red through deeper tints of brick-red to dark chestnut
and sooty brown.

Black-chaffed varieties are comparatively rare, although examples are
met with in all the races of wheats. Several shades are included under
the term " black." In some glabrous forms the chaff is almost jet black,
in others a dark sooty brown ; these colours when overlaid by a pale
waxy covering or a felt of soft hairs are modified to bluish black or
mouse-grey tints.

In most black-chaffed wheats the colour is only developed to its

THE INFLORESCENCE 103

full extent on exposed parts of the ear, the portions of the flowering
glumes covered by the empty glumes being pale red or light yellow.

Among certain forms of T. dicoccoides, T. aegilopoides , and T. durum
hybrids the colour is more or less irregularly distributed in lines or patches
over the empty glume ; in others it appears only as a single dark line
along the outer margin of the glume, the latter character being con-
spicuous in some forms of T. turgidum, var. melanatherum, and in many
hybrids of T. dicoccoides, T. dicoccum, and T. durum.

It is only in hot, bright seasons that the melanotic pigment is fully
developed ; in cooler damp years the glumes of black-chaffed varieties,
especially those belonging to T. vulgare and T. Spelta, are generally an
ashy-grey tint.

The glumes are either glabrous or more or less clothed with hairs.
Sometimes the hairs are few and confined to the nerves, but in many
forms they cover the whole surface with a tomentum or felt of varying
density. The hairs are unicellular and terminate in a fine point.

In T. aegilopoides and T. monococcum the hairs are very slender, from
•16 to -32 mm. in length, and of nearly uniform diameter. Both long hairs
and short ones, about one quarter the length, are found on the glumes
of T. dicoccoides and T. dicoccum. the long ones measuring from -5 to -65
mm., the short ones averaging about -12 mm.

Similar slender silky hairs of two fairly distinct sizes occur on the
chaff of T. durum and T. turgidum, the long ones being -6- -8 mm. in
length, the shorter only -i6--2 mm. in T. durum, and -2- -32 in T.
turgidum (a, Fig. 79).

The hairs on the glumes of the pubescent varieties of typical vulgare

a b

FIG. 79. — Hairs from the glumes of (a) T. durum, (b) T. vulgare (x 40).

wheats vary also in length, but the difference between the long and the short
ones is not so great as in the races of wheats already mentioned ; moreover,
they are readily distinguished by their stouter appearance, greater width,
and thicker walls (b, Fig. 79). The long hairs usually measure from
•6 mm. to i -2 mm. in length, the short ones from -2 mm. to -3 or -4 mm.
In some vulgar es, typical stout vulgare hairs are found mixed with
fine, slender, short ones, resembling those of T. durum, a similar mixture

104 THE WHEAT PLANT

being seen also in some forms of T. polonicum. The hairs of T. Spelta
in some cases are stout, and of nearly uniform size -4- -6 mm. long ; in
a few forms, stout hairs -4 mm. long are mixed with stout short ones
•12 mm. long.

In dimensions the hairs on the glumes of T. compactum agree some-
what closely with those of T. turgidum.

Inserted alternately on opposite sides of the short rachilla are the
flowering glumes, in the axils of which the flowers arise. They are boat-
shaped, many-nerved, and without keels, the upper part notched and
ending in a point or a long awn, the length of which usually increases in
the spikelets near the tip of the ear.

The awns are tapering and triquetrous, with forward pointing scab rid

projections running longitudinally along the angles ; they are generally

straight, but may be sinuous or even bent into the form of a hook or spiral,

as in some Asiatic forms of T. vulgare. In a Persian form of T. vulgare

the awn has a pair of thin membranous and colourless outgrowths (Fig. 217).

The separation of wheat into " bearded " and " beardless " kinds was

Y made as soon as the classification of the cereal was attempted, the two

groups being characterised respectively by the presence or absence of a

long beard or awn on the flowering glumes of the ear.

In the bearded wheats, long awns occur on the flowering glumes of
all the spikelets from the base to the tip of the ear, the longest being a
little above the middle of the ear, the shortest at the base and apex. The
two lower flowering glumes of each spikelet have awns nearly equal in
length and fully developed, the third and succeeding glumes having
shorter awns which are frequently reduced to a fine point, especially when
the subtended flower produces no grain.

Strictly beardless wheats entirely without awns are of exceptional
rarity. I have met with them only in a small group of Indian, Japanese,
and Australian forms of T. vulgare, which I have little doubt are genetically
related ; in these, the flowering glume terminates in a short tooth, like
that of the empty glume

The term " beardless " is, however, used not only for those wheats
in which awns are altogether absent, but for many hundreds of forms
having short points or greatly reduced awns, usually not more than i cm.
in length, confined to the tip of the ear. In a small percentage of cases
one or two awns on the terminal spikelets reach a length of 2-5 cm., but
the difference between bearded and beardless varieties is fundamental, and
there is no difficulty in assigning any ear to one or other of these two
classes, except, perhaps, in the case of rare, unstable hybrid forms in which
awns of intermediate length sometimes appear over a considerable length
of the ear.

In truly bearded ears the awns are uniformly distributed from the top

THE INFLORESCENCE 105

to the bottom of the ear with the longest never at the apex, while in beard-
less ears whatever awns are present are longest near the apex of the ear,
the rest rapidly diminishing in length towards the base, where they are
rarely more than 1-3 mm. long.

Beardless forms are not found in T. aegilopoides, T. dicoccoides, T.
monococcum, T. orientale, and T. pyramidale, and are of great rarity in T
dicoccum, T. durum, T. turgtdum, and T.polonicum, but among the remain-
ing races, viz. : T. vulgare, T. compactum, T. sphaerococcum, and T. Spelta,
both classes are equally common, almost all the bearded varieties of these
wheats having a corresponding beardless representative.

The awns are white, red, or black, those of the two former tints being
found only on white or red glumes respectively ; black awns are met
with on glumes of all colours, white, red, or black.

Varieties with black awns are seen among all races of wheat, but are
most common in T. durum and T. turgtdum. They are exceptionally rare
in T. vulgare. I have seen but few examples of typical ears of the latter
race with jet-black awns ; these came from Asia Minor and Central Asia.
The black pigment is always most conspicuous in specimens grown in
bright hot climates.

In some wheats the black colour is almost constant under a wide
range of climatic conditions ; in others, it is dependent on the intensity
of the light, heat, and moisture at the time of ripening. In cool, damp
seasons the black pigment in the latter forms never develops, but returns
in hotter years. This variation I have repeatedly observed in the progeny
of single ears, perhaps more especially in forms of T. durum, var. melanopus,
and similar results have been recorded by Metzger, Koernicke, Howard,
and others in different varieties. Such variation greatly reduces the
taxonomic value of black awns.

The fact that external conditions of soil and climate greatly influence
the development of pigment in the awn complicates the problem of its
inheritance.

Opposite each flowering glume, but attached to the very short floral
branch with its back to the rachilla is the palea, a symmetrical thin mem-
branous glume with two prominent lateral veins, along which runs a line
of stiffish hairs ; the part between the veins is concave and the two
semi-transparent margins curve inwards round the flower.

STRUCTURE OF RACHIS, GLUMES, AND AWNS

The rachis is notched, consisting of short internodal lengths, each of
which is convex on one side and more or less flattened on the other ; the
lower internodes are somewhat semicircular in transverse section, the
upper ones flattened and spindle-shaped in section (Fig. 80).

io6

THE WHEAT PLANT

The structure resembles that of the internodes of the culm. On the
outside is an epidermis of oblong cells, their length from 6 to 10 times their
breadth, with sinuous thickened walls and oval or kidney-shaped " dwarf "
cells alternating with them. At the edges are unicellular epidermal hairs
of variable length.

Lines of stomata resembling those of the stem occur in the epidermis
overlaying the longitudinal bands of chlorophyllous parenchyma which
are found just beneath the epidermis on the convex side only. These

FIG. 80. — Transverse sections of the rachis ( x 25). A, a lower internode ; B, an upper
internode ; s, stereome ; c, chlorophyllous tissue ; r and x, vascular bundles.

green bands are most abundant in the lower internodes of the rachis (c,
Fig. 80), being reduced to two or three narrow lines near the edges in the
upper portions. Within the epidermis on the flatter side of each inter-
node of the rachis which is hidden by the back of each spikelet there is a
continuous band of stereome (s, Fig. 80).

The central part of the rachis is more or less completely filled with
thin-walled ground parenchyma, imbedded in which are numerous
vascular strands, the larger ones arranged in a circle or ellipse, the smaller
bundles being found close to the inner side of the chlorophyllous bands.

THE INFLORESCENCE

107

The rachilla is delicate and flattened with an arrangement of tissues
similar to that found in the rachis. The epidermal cells have straight
walls not greatly thickened.

Numerous hairs are usually present.

There are generally three slender vascular bundles running through
the thin-walled ground tissue.

The structure of the empty and flowering glumes resembles that of
the ordinary leaf-sheath.

The outer epidermis of the empty glume consists of rows of oblong
cells 80-100 p, long and about 25 /x broad, with oval kidney - shaped
" dwarf" cells 16x25^ between them at short intervals. The walls of
these cells have characteristic wavy thickenings and
simple pits (Fig. 81). The margins of the glumes
are fringed with hairs, and trichomes of variable
length are present on the outer surface, some being
only short thick- walled papillae about 16 /x high.

FIG. 81. — a, Epidermis of empty glume ; b, of palea ( x 210).

Hairs are also found sometimes on the inner epidermis of the glume.

Rows of stomata occur especially in the apical region of the glume.

The cells of the inner epidermis have thinner walls ; a few stomata are
seen, and numerous hairs 50-70 //, long are also found on the inner surface,
especially in the parts over the midrib and stronger nerves of the glume.

Narrow longitudinal green bands of thin-walled parenchymatous
tissue accompany the vascular strands ; the rest of the ground tissue has
thickened walls. Delicate vascular bundles traverse the glume longi-
tudinally and anastomose near the apex, the midrib strand terminating
in the apical tooth.

The tissues of the flowering glume are similar in many respects to those
of the empty glume ; stomata and hairs occur on both surfaces.

io8

THE WHEAT PLANT

In transverse section it is seen that near the margins of the glume
the space between the outer and inner epidermis is occupied by strong
thick-walled ground tissue traversed by vascular bundles and narrow
longitudinal bands of assimilating tissue which lie
close to the outer surface.

Away from the margins the glume is thinner and
the stereome extends only to the middle of the
section, the inner half consisting of thin- walled
chlorophyllous parenchyma covered by an epidermis
of larger thinner-walled cells than those on the
outside of the glume.

A number of vascular bundles run from the base
to the tip in bearded glumes ; the central one and
two of the lateral strands are continued into the awn.
The awn tapers from the base to the tip, and is
triangular in section, the sides approximately equal.

One of the sides is parallel to the flattened rachilla
and is continuous with the inner surface of the
glume -to which the awn belongs, the two other sides
corresponding to the outer surfaces of the right and
left halves of the glume.
FIG. 82.— Portion of an The epidermis is composed (i) of narrow elon-

awn of T. turgidum j n • i n i • • • i • 1

( x 35). c, Band of gated cells with walls showing wavy thickening and
chlorophyllous tissue ; numerous simple pits, (2) small oval or squarish

n, scabrid hairs. r ., , .

dwarf cells often projecting as papillae, and (3)

short thick- walled unicellular hairs with fine points which are directed
forwards and give the awn its scabrid character (h, Fig. 82). In the

FIG. 83. — Transverse sections of an awn of T. turgidum (*so). A, Near the base ; B,
at the middle ; C, near the apex ; s, stereome ; c, chlorophyllous tissue ; x, stoma ;
b, vascular bundles.

epidermis on the two outer faces of the awn are longitudinal lines of
stomata which communicate with green assimilating tissue within. The
latter consists of two separate bands of chlorophyllous parenchyma,

THE INFLORESCENCE 109

which traverse the awn from the base to near the apex, where they
unite into a single central strand.

Immediately within the epidermis, at the three angles and around the
inner face of the awn, is strongly developed stereome, the centre being
occupied by thinner- walled parenchyma.

Three vascular strands are found in each awn, namely, one large
bundle in the angle between the two bands of green tissue and continuous
with the midrib of the glume, and two smaller ones right and left of this
also entering from the glume (Fig. 83).

The palea is of simple structure and between the two keels is only
two or three cells thick (about 30 /LI). The epidermis is chiefly composed
of elongated cells and circular " dwarf " cells with sinuous walls ; over
the keels, the margins and surfaces of the apical portions of the palea it
possesses hairs 50-1 30/^1 long.

A simple vascular bundle runs through the centre of each keel ;
associated with it is a small band of green parenchyma, chloroplasts being
absent from other parts of the palea.

A few stomata are found over the green tissue.

CHAPTER VIII

THE FLOWER : FERTILISATION AND DEVELOPMENT OF THE GRAIN

IMMEDIATELY within the flowering glumes are two minute antero-lateral
scales, the lodifrules, which probably represent a small divided glume,
although they are sometimes regarded as rudimentary perianth leaves.

They are thin and colourless,
about i mm. long, -J--8 mm.
wide near the top, and -4- -5
mm. at the base, with long
hairs on the upper margin ; at
the flowering period they swell
enormously, becoming for a
short time more or less spheri-
cal, resembling pellucid drops,
pushing apart the glumes and
exposing the stamens and
stigmas of the flower ; later,
they collapse and the glumes
close again.

The perfect flower is hypo-
gynous and simple in struc-
ture, consisting of a whorl of
three stamens and a single
carpel (Fig. 84).

One or more of the upper
flowers of a spikelet are im-
perfect, the ovary being rudi-
mentary and the anthers
without pollen grains ; fre-
quently the terminal flower
is missing altogether, and the flowering glumes and pales, though present,
are poorly developed.

In many instances the entire spikelet is sterile ; especially is this the
v case in the lowest three or four of an ear. Occasionally one or two of
the apical spikelets are abortive

FIG. 84. — Flower before anthesis. a, Anther ;
style ; /, lodicule. Dehiscing anther below.

THE FLOWER

in

The stamens have slender filaments which are at first 2 to 3 mm. long,
but grow rapidly when flowering occurs and reach a length of 7-10 mm.

A

a

FIG. 85. — A, Surface view of anther
(in chloral hydrate) ; B, longi-
tudinal section of young anther ;
a, epidermis ; b, endothecium ; c,
tetrad of pollen-grains (x 175).

FIG. 86. — a and c, Epidermal cells of anther
lobe ; b, endothecial cell ( x 250).

A B

FIG. 87. — A, Transverse section of anther lobe just before dehiscence ( x 63) ; B, enlarged
view of cells at the point of dehiscence ( x 125) ; C, earlier stage of point of
dehiscence ; a, epidermis ; b, endothecium ; c, schizogenous cavity.

The anthers are bi-lobed and versatile, the filament being attached to the
connective at a point about I mm. from the base of the anther lobes.

112

THE WHEAT PLANT

They usually measure 3 to 4 mm. long and -5 to i mm. across, and are
green when young, becoming yellow or pink when ripe. Each lobe
contains two loculi, the tissue between which breaks down at the time
of dehiscence.

The filament of the stamen before anthesis is cylindrical, of uniform
thickness, and from 100 to 120 /x in diameter. It consists of long paren-
chymatous cells -5- -8 mm. long and 9-12 /x in diameter. In the centre
is a small vascular strand possessing one or two annular tracheae 8-9 /x in
diameter, with closely arranged rings or an occasional spiral thickening.
When flowering occurs the filament grows rapidly, becoming very irregular
in thickness, and the rings in its vessels greatly separated from each other.
The tissue of the connective is parenchymatous, with a slender vascular
bundle resembling that of the filament running through it ; in the

epidermis investing it on the
dorsal side are two rows of
simple stomata.

The epidermis of the anther
lobes is composed of narrow
cells with their long axis par-
allel to the long axis of the
anther. Each cell is from 50
to So/ji long and 10 to 15 /x in
diameter, its outer wall having
a cuticle which is covered with
fine wavy longitudinal lines
(c, Fig. 86).

Immediately within is the
endothecium (b, Figs. 85, 86,
87), consisting of narrow cells about 60 /x long and 10-12 /x broad with
their long axes arranged across the cells of the epidermis : a short time
before dehiscence of the anther the walls of these cells develop character-
istic short pyriform thickenings, which project at right angles from the
wall into the cell lumen (b, Fig. 86).

The pollen-grains are smooth and spherical, or slightly oval, somewhat
irregular in size and form : their diameters measured in a dry state as shed
from the anther are —

T. aegilopoides . 37-45 T. dicoccum . 48-52 T. compactum . 52-61
T. monococcum . 37-45 T. durum 48 T. vulgare . . 61-65

T. dicoccoides . 44-64 T. turgidum . 57-65 T. Spelta . . 65

In the exine is a circular or oval pore closed by a minute lid 4-5 /x in
diameter (Fig. 88), which is pushed off when the pollen tube develops.

Within the pollen-grain are large numbers of minute oval starch-
grains from i to 4 /x long ; these are more abundant in the half of the grain

FIG. 88.-Pollen-grains showing pore in the exine.

THE FLOWER

FIG

a, Transverse section of young

ovary ( x 70) ; b, portion of its wall
( x 260).

on which the pore is found and are utilised in the development of the

pollen-tube ; they are not found in the immature pollen-grain.

The gynaecium consists of a single median carpel, with an obovate

or obconical ovary, slightly trilobed in transverse section and flattened on

the ventral side. At the time of an-

thesis it measures i mm. long and is

about the same diameter across the

upper portion.

Its sides are smooth, but the broad

apex is clothed with numerous long

finely-pointed unicellular hairs about

1 6 ft, in diameter and varying in length

from -16 mm. to i mm.

The wall of the ovary averages -35 mm. in thickness. It consists of

delicate loosely-packed parenchyma 10-16 cells across, clothed on the

outside with a well-defined epidermis, its inner surface being lined with
a similar but thinner epidermal layer. Running
longitudinally through it are two very fine lateral
and one dorsal vascular strand, each of the former
possessing three to six delicate tracheae, the latter
usually only one (Fig. 89).

From the tip of the ovary arise two terminal
styles 3-4 mm. long, which curve outwards when
flowering occurs. Each has a central tapering
column from which are given off 80-100 delicate
stigmatic branches -8 to 1-5 mm. long ; the latter
extend to the base on the inner side, but only
about one-third of the way down the outer side of
each stylar column.

The two lateral vascular bundles of the ovary
are continued some distance into the styles.

Each delkate ^g™^ branch is Composed of

210) and transverse sec- four rows of elongated cells about 250 /z long and
8-10 (j. in diameter. They are joined throughout

their whole length except at the upper end of each cell, which is free,
and curved outwards into a bluntish point 24-30 ^ long (Fig. 90).

The angles between the free ends of these cells and the sides of the
branches form convenient resting-places for the pollen-grains shed from
the dehiscent anther at the time of flowering.

Within the ovary is a single sessile anatropous ovule attached to a
broad placenta on the inner ventral surface of the ovary (Fig. 91). It
is about -5 mm. long and -4 mm. broad, and possesses two integuments ;
the outer one, which is disintegrated before the seed is ripe, is shorter

i

ii4 THE WHEAT PLANT

than the inner one and does not reach the micropylar opening. Each of
the ovular coats consists of a double layer of cells 14-16 p thick.

The nucellus has a broad chalaza, and is composed of parenchyma
surrounded by a well-defined epidermis of regular closely-fitting cells,

which are almost square in transverse
section and uniformly about 8 p, in
diameter except where they abut on the
micropyle, at which point the layer is
thinner.

Imbedded in it is the embryo-sac,
•27 mm. long by -16 mm. broad, a
large oval or pear-shaped cell whose
narrow end is in immediate contact with
the thin epidermis abutting on the
micropyle ; at other points it is sur-
rounded by nucellar tissue usually 6 to
8 cells deep.

In the early stages of the develop-
ment of a spikelet of T. vulgare eight
or nine flowers can be recognised, but
under the ordinary conditions of plant
growth in the field none of the spikelets
yield this number of ripe grains. So
soon as the three or four lowest flowers
FIG. 91. — Longitudinal section of the have reached a certain stage of develop-
inttguments ment, the upper four or five cease to
grow and their tissues become depleted
of their cell - contents, the material probably going to assist in the
nutrition of the rapidly growing flowers below.

In a very young spikelet all the cells of the tissues up to the apex
contain normal cytoplasm and well-defined nuclei ; but in the early part
of June, when growth and differentiation of the parts of the flowers are
proceeding rapidly, the cytoplasm and nuclei of the cells of the upper
parts of the spikelet show signs of change. The cytoplasm loses its
staining power and the nuclei begin to degenerate : later, the cytoplasm
disappears, the nuclei remaining for a time, after which they also vanish,
leaving only the cell walls.

Depletion of the cells begins first at the undifferentiated tip of the
axis of the spikelet and gradually proceeds downwards to a point where
the three or four basal flowers are growing normally.

THE FLOWER 115

The cell - contents of the periblem disappear before those of the
plerome, and the stamens of the
aborting flowers lose their contents
and degenerate before the ovaries.

The essential floral organs and
the bracts by which they are en-
closed arise in succession as bulging
masses of meristem below the apex
of the young flower (Figs. 92, 93).

The flowering glume arises first
in the form of a thick semicircular
zone surrounding the tip on the
anterior side. Very soon the
wjiorl of stamens can be recognised
as three rounded papillae. A little
later the carpel grows up as a
crescent-shaped ridge closely en-
circling the hemispherical floral
apex on the anterior side ; about
the same time the rudiments of
the palea and the lodicules originate
almost simultaneously, the latter

FIG. 92. — Median longitudinal section of a
young spikelet (May 27). e, Empty
glume ; /, flowering glume ; p, palea ;
/, lodicule ; £ , stamen ; 9 , carpel ( x 25).

from points on the axis slightly beneath the line of insertion of the

stamens (Figs. 92, 93).

In a short time the rudiments of the styles

are visible as two bluntish conical outgrowths of
f \' ).} the apical margins of the young carpel. The

upper part of the carpel growrs over and closes

round the apex of the axis, coming into contact

FIG. 93. — Upper flowers of a young spikelet showing developing flowers.
/, Flowering glume ; 5, stamen ; o, ovary ( x 50).

with it and forming a chamber in which the enclosed portion of the axis

n6

THE WHEAT PLANT

appears as a protuberance from the posterior inner surface of the cavity.

-The enclosed tip of the axis ultimately becomes the ovule (Figs. 94, 95).

The primordia of the stamens rapidly elongate, their transverse

sections becoming oblong with rounded corners. At first they consist

of homogeneous meristem, but differentiation into dermatogen, periblem,

FIG. 94. — Stages in the development of the carpel ( x 25).

and plerome with the initials of a central vascular strand is visible
when the anther is about -15 mm. in length.

A single sub-epidermal arche-
sporial cell is soon marked out
in each corner of the anther.
Each of these divides by peri-
\ \ / \ i clinal walls into an outer primary
parietal cell and an inner primary
FIG. 95. — Successive longitudinal sections of a sporogenous cell. The former

young carpel from the centre to the outside ,. ., .. ,,

(x2s). divides radially, and usually

after two divisions the four cells

produced form with the adjoining periblem cells the first parietal layer
which encircles the sporogenous cell (Fig. 96). Two successive periclinal
divisions follow, resulting in the production of three concentric parietal
layers, the inner one in contact with the sporogenous cells becoming the
tapetum, the next surrounding zone is the " middle " layer, the third,

THE FLOWER

ii'

immediately beneath the epidermis, developing into the endothecium
of the mature anther.

For a time the sporogenous cells form a single longitudinal row. About
the period when the tapetum is formed they divide both radially and
longitudinally ; in a median longitudinal section two rows are seen at
this stage, a transverse section usually showing six radially arranged
cells, all of which on their outer surfaces are in contact with the tapetum ;
these are transformed without further division into pollen mother-cells,

FIG. 96. — i, Transverse section of a young anther, showing initial archesporial cells at
each corner ; 2, first division of archesporial cell ; 3-6, transverse sections of portions
of an anther, showing subsequent divisions ; 7, longitudinal section of 6 ; p, d, sporo-
genous cells (x 260).

which separate as the anther loculi enlarge, becoming free spherical or
oval cells about 35 /n in diameter, each with a large central nucleus.

The free pollen mother-cells, which usually form a single layer lining
the enlarged loculus (Fig. 98), undergo two successive divisions, the first
being the reduction division, the daughter nuclei receiving in some races
of wheats eight short chromosomes.

The number of chromosomes, however, is not the same for all races of
wheat, and variations have been recorded in different cells of the same
individual plant.

n8

THE WHEAT PLANT

In the few cases in which I have been able to obtain reliable counts
the haploid number is 8 in T. monococcum, and 16 the diploid number in
some forms of T. vulgare. Larger numbers, not possible to count with
certainty, I have observed in somatic cells of other forms of T. vulgare.

FIG. 97. — i, Transverse section, 2, longitudinal section of a loculus of a Jtoung anther
( x 210) ; a, epidermis ; b, endothecial layer ; c, " middle " layer ; t, tapetum ; p, pollen
mother-cells.

Overton (1893) gives 8 as the haploid and 16 as the diploid number
in T. vulgare

FIG. 98.— i, Transverse section of anther-lobe, showing single ring of free pollen mother-
cells in each loculus ( x 100) ; 2, longitudinal section of a portion of the wall of the
anther (x 260) ; 3, surface view of cells of the tapetum (x 260) ; n, endothecium ; c,
" middle " layer ; b, tapetum; a, pollen mother-cells.

Kornicke (1896) found 16 in the somatic cells and 8 the reduced number
in the embryo-sac and pollen mother-cells of T. compactum ; in one vege-
tative cell of the anther of the same plant he counted 24 chromosomes.

The number observed by Dudley (1908) in the first division of the
pollen mother-cells of T. vulgare was 8.

Diploid
No.

Diploid
No.

T. vulgare

. 42

T. turgidum

. 28

T. compactum

. 42

T. durum

. 28

T. Spelta

. 42

T. polonicum

. 28

THE FLOWER 119

Spillman (1909) states that wheat has 40 or more chromosomes.

Nakao (1911) gives 8 as the haploid number, and Bally (1912) found
the same in the wild T. dicoccoides and T. vulgare.

Sax (1918) found approximately 28 in the first division of the fertilised
egg of T. durum (Kubanka) and approximately 40 in the fusion-nucleus
of the embryo-sac.

Sakamura (1918), working chiefly with root-tips, obtained the follow-
ing results :

Diploid

No.

T. dicoccum . 28
T. monococcum . 14

This series with 7 instead of 8 as the basic haploid number is remark-
able. These numbers require further investigation.

At the first division the pollen mother-cell is bisected into two equal
hemispherical halves with a cell wall between each ; the next division
occurs in a plane at right angles to the first, and is also followed by the
formation of a partition wall. The four pollen-grains thus produced lie
in the same plane and remain for a short time within the delicate wall of
the mother-cell ; later, each develops its own wall, which is differentiated
when mature into an outer thick cutinised coat, the exine, and a thin
cellulose lining, the intine of the spore ; in the exine is the pollen-tube
pore previously described.

The pollen mother-cells when rounded off and the tetrads produced
from them are arranged in regular order as a layer round the inner wall
of the loculus ; the free pollen-grains become scattered irregularly through-
out the cavity.

The pollen-grains germinate while still in the unopened anther, and
before the protrusion of the pollen-tube producing the vegetative or tube-
cell possessing an oval nucleus, measuring 7 ^ x 8 /z, and two slender male
gametes, each of which is curved and pointed at one end and measures
10-12 /A x 2 [t, (Fig. 103).

During the development of the pollen-grains changes occur in the cell
layers constituting the walls of the anther loculi. The middle layer,
which is the thinnest, collapses, its walls and scanty contents appearing
as a narrow line round the outer boundary of the tapetum.

The tapetum consists of oblong parenchymatous cells square in trans-
verse section with abundant cytoplasm and containing at first a single
large nucleus. About the period when the sporogenous cells undergo
their first division the nuclei of the tapetum divide, the two daughter
nuclei in each cell persisting with no wall between them until the layer
disappears, when the pollen-grains are perfectly developed (Fig. 98).

120 THE WHEAT PLANT

Golinski states that these double nuclei frequently coalesce to form a
single nucleus again, but I have not been able to confirm this.

At the time of the divisions of the pollen mother-cells the tapetum
shows signs of disorganisation, its cells separating more or less as the
anther lengthens ; later, it disappears altogether, its contents breaking
down and furnishing nutritive material for the final development of the
pollen-grains.

When the latter round themselves off and their coats become cutinised,
the cells of the endothecium exhibit their characteristic thickenings and
the exothecium or epidermis acquires its wavy-lined cuticle.

The ovule at first consists of a hemispherical mass of meristem within
the carpellary cavity. Although it has the appearance of a lateral out-
growth on the inner posterior wall of the ovary, observation of its develop-
ment suggests that it is derived from the morphological apex of the floral
axis, its apparently lateral position being due to rapid growth of one side
of the axis before the closure of the carpel.

In the earliest stages the nucellus' is an almost uniform meristem,
usually six or seven cells across. A cell immediately beneath the epidermis
occupying the apex of the axial row of the mass grows into a large some-
what wedge-shaped archesporium. Division of the latter into two cells
occurs with the formation of a periclinal wall between them ; the dyad
divides again similarly, and a tetrad row of megaspore mother-cells is pro-
duced, no parietal cell being cut off the archesporium at any stage (Fig. 99).

The outer three cells of the tetrad collapse rapidly and disintegrate,
the innermost remaining for a short time imbedded in the nucellus some
distance from the epidermis. Soon, however, the surviving mother-cell
enlarges greatly, and pushing apart the cells on the sides of the narrow
channel left by the disintegrated sister cells, ultimately comes into contact
with the epidermis ; later it functions as the embryo-sac.

The germination and development of the embryo-sac up to the eight-
celled stage follow the usual course. It increases greatly in size and at
the same time absorbs some of the adjacent nucellar tissue. At the micro-
pylar end of the sac are two pear-shaped synergidae, each about 35-40 //,
long and 16-20 /j, broad. The ovum, a broader cell about 50 //, long and
32-35 {J, across, is placed by the side of these ; its nucleus is about 16 ^ in
diameter, and at the time of fertilisation generally contains two spherical
nucleoli, one about 4 /z across, the other half this size. Near the ovum
is the smaller upper polar nucleus, the lower one being placed near the
three antipodals at the chalazal end of the megaspore.

The two polar nuclei move towards each other and meet at a point
just behind the egg-apparatus (Fig. 100) ; here they remain in close con-
tact for a considerable time, and probably fuse to form the primary endo-
sperm nucleus just before the fertilisation of the ovum.

FIG. 99. — Median longitudinal sections of young ovules, a, Showing primary arche-
sporial cell ( x 260) ; b, after its first division ; c, tetrad stage ; d, inner cell of
tetrad after the collapse of the three outer cells. Outlines of the two ovular integu-
ments are shown (b, c, d x 350).

FIG. 100. — Egg apparatus and fusion-
nuclei (June 12) (><35o).

FIG. 101. — Embryo-sac before fertilisation
(June 12) (x 280).

122 THE WHEAT PLANT

Trio-three antipodals divide and increase in size at the expense of the
nucellus, producing a convex mass in contact with the wall of the embryo-
sac on the placental side a short distance from the chalazal end (Figs. 101,
105). The antipodal complex in the mature sac usually consists of 6
to 10 laige cells, although Kornicke mentions 36 or more. Each measures
from 3aTto 60 ^ in diameter and contains much vacuolated cytoplasm and
a large nucleus ; after fertilisation is accomplished the cells are gradually
disorganised, their contents being apparently utilised in the formation
of the endosperm ; their large nuclei, which may reach a diameter of
40 fj, with nucleoli 16 p or more across, are the last to disappear.

During the development of the flower, the stamens and their pollen-
sacs are for a time considerably in advance of the carpel and ovule.

At the period when the anther possesses a well-defined single longi-
tudinal row of sporogenous cells surrounded by two parietal and one
epidermal zone, the carpellary cavity is open above, and the mass of
meristem, from which the ovule is formed, shows neither integument nor
archesporium.

When the first or inner integument is beginning to develop and the
archesporium is clearly defined, the sporogenous cells of the anthers have
divided and a double longitudinal row of closely fitting cells is seen in the
pollen-sacs ; the tapetum has also been formed, so that at this stage there
are four layers round the sporogenous cells.

The midiment of the second or outer integument of the ovule appears
before the archesporium divides. The nucellus becomes more and more
enclosed by the integuments, and two successive mitoses of the arche-
sporium occur, resulting in the production of a tetrad row of megaspore
mother-cells. Pollen-grain tetrads, or rounded separate pollen-grains
with single nuclei and more or less cutinised walls, are now found in the
anthers ; the cells of the tapetum are still somewhat square in transverse
section, but the cytoplasm is reduced and much vacuolated.

When the embryo-sac is ready for fertilisation, each pollen-grain con-
tains the tube-nucleus and two gametes, the tapetum has become com-
pletely disorganised, and the endothecial layer shows its characteristically
thickened walls.

V

ANTHESIS AND POLLINATION

Anthesis, or opening of the flower, follows the escape of the ear from
the upper leaf-sheath in five or six days, although in some instances it
occurs on the first day, or may be delayed till the ninth after the appear-
ance of the ear. The development of the stamens and gynaecium in the
same flower proceeds at the same rate, and the pollen is ready to be shed
from the anthers when the stigmas become receptive.

THE FLOWER 123

At the time of anthesis the thin-walled parenchyma of the lodicules
becomes very turgid, the outer face of the tissue assuming a convex form.
The two lodicules in this swollen state act as elastic cushions which push
apart the flowering glume and palea, the separation taking place rapidly,
often attaining a maximum in five minutes or less. Very slight mechanical
stimulus either affects the turgidity of the lodicules or releases thfe edges
of the glumes from each other, for the opening of the glumes often takes
place rapidly, on gently running the ear through the hand, by touching
the awns, or by the impact of the ears against each other as ,they are
swayed to and fro in a breeze.

The angle of separation of the glumes varies considerably.! In T.
aegilopoides , T. dicoccoides , and T. dicoccum, and in some varieties of T.
vulgare, it may reach 40° or more, especially in bright warm weather ; in
the majority of cases, however, it does not exceed 2O°-3O°, and in T. Spelta,

FIG. 102. — Anthesis of lowest flower of a spikelet. a, 8.48 A.M. ; b, 8.50 A.M. ;
c, 8.56 A.M. ; d, 9.7 A.M., June 19, 1915 ( x 2).

T. compactum, and certain forms of T.polonicum and T. vulgare it is often
less than this.

As soon as the glumes begin to separate, rapid changes occur in the
floral organs. The styles, which at first lie close together and parallel
to the long axes of the ovary, quickly curve away from each other, and the
feathery stigmas spread out ; in most wheats these remain within the
glumes, but in some varieties their tips may be seen projecting on opposite
sides of the flowering glume. At the same time the filaments of the
stamens lengthen rapidly, expanding from a length of 2 or 3 mm. to 7-10
mm. in two to four minutes, and the anthers are thrust out of the opening
between the glumes, usually on the outer face of the spikelet (Fig. 102).

The pollen-sacs commence to dehisce when the filaments begin to
lengthen and a variable amount of pollen is left in each flower ; self-pollina- (_ \
tion is the rule.

Dehiscence progresses from the tip to the base of the anther, and is

124 THE WHEAT PLANT

completed after the latter is pushed out of the flower, a large proportion
of the pollen being shed into the air. In many flowers all the anthers
are carried into the open air at the time of anthesis, but quite frequently
one or more fail to escape from the glumes, being entrapped behind the
folded margin of the palea or caught between the tips of the closing glumes.
Retention of the anthers, which is most frequent in dense-eared wheats,
occurs chiefly in the lower spikelets of ears and the smaller flowers of each
spikelet, some of which do not open at all. Of flowers which open, the
glumes generally remain widely separated from 5 to 15 minutes and then
slowly close again, the lodicules meanwhile becoming flaccid ; the time
taken in opening and closing completely varies from 8 to 30 minutes or
more, the average being about 20 minutes.

Anthesis occurs in the early part of June at Reading, the individual
flowers opening at various times of the day, from dawn till an hour or two
after sunset.

Observations upon representatives of the chief kinds of wheat revealed
no characteristic specific differences ; flowering was observed to occur
during most hours of daylight, but it was especially frequent between the
hours of 5-7 A.M., 9-10 A.M., 2-3 P.M., and 8-9 P.M. Meteorological con-
ditions appear to influence the time at which it takes place. Gitkova at
Saratoff, Russia, found the flowering of a variety of T, vulgar e and T.
durum was most intense between 5 and 7 A.M., with a second maximum
between 5 and 6 P.M. Leighty and Hutcheson also noted two periods
of extensive blooming during daylight, namely, from 7 to 9 A.M., and in
the middle of the afternoon, with a secondary morning period about

II A.M.

The basal flower is the first to open in each spikelet, the others following
in regular succession upwards.

The position of the first spikelet to flower varies a little ; it is generally
in the middle third of the ear, and usually near the upper part of this
section, except in the case of T. aegilopoides, T. monococcum, and T. dicoc-
coides, in which the spikelets to open first are nearer the tip, at a point
about the middle of the upper half.

In ears possessing 16-18 spikelets the 8th to the i2th usually open first ;
in those with 28-30, the i6th to aoth. Anthesis proceeds upwards and
downwards from these points in more or less regular succession, the apical
and basal spikelets being the last to flower. Sometimes 4 or 5 spikelets
on one side open before the rest ; in other cases adjacent spikelets on
opposite sides of the rachis flower almost simultaneously.

The successive flowers of a spikelet usually open on successive days,
but occasionally two of the same spikelet open on the same day, one in
the morning, the other later, after an interval of 5 or 6 hours.

At Reading the whole ear often completes its flowering in 3 to 5

Right Side.

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Flowers of Spikelet.

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128

THE WHEAT PLANT

days when the air is warm and the sky clear : in wet or dull weather the
period is prolonged to 6 or 8 days.

The first ear to flower belongs to the primary culm and is followed by
those on the lateral stems in order of origin, the whole plant finishing the
process in normal seasons in about 8 days.

No. of Spikelets
of the Ear.

Spikelets which open
first, counting from
the Base upwards.

Days occupied in
completing Anthesis
of an Ear.

T. monococcum

28

i 7th- i 9th

T. dicoccoides

IS

ioth-i3th

6

T. dicoccum —

" Ajar "

19

i 2th- i 5th

3-5

Indian ....

17

8th- i 2th

4

European ....

3°

i 8th- 20th

T. Spella ....

21

I3th-isth

3-5

T. vulgare —

Chinese ....

20

ioth-i2th

4

Sq. head ....

24

i 2th- i 4th

6-8

Lammas ....

18

ioth-i2th

5

Karachi ....

16

7th-Qth

4

T. durum ....

25

i 2th- i 4th

4

T. turgidum ....

27

i 8th- 20th

T. compactum

17

yth-gth

5

The following results were obtained by Obermayer :

NUMBER OF FLOWERS OPENING EACH DAY ON FIVE ORDINARY
HUNGARIAN WHEAT PLANTS

1912.

Up to 7 A.M.

7-Q A.M.

Q-II A.M.

II A.M. -I P.M.

1-3 P.M.

3-5 P.M.

5-7 P.M.

After 7 P.M.

Tot?l.

June 5

9

10

3

6

28

56

,, 6

14

17

25

34

10

28

37

3

168

7

39

64

27

65

43

33

51

6

328

... 8

"3

72

98

41

30

58

35

6

453

9

112

49

57

29

3i

45

25

6

354

, 1°

75

5°

89

13

17

26

. .

270

, ii

24

48

ii

4

12

21

18

138

, 12

8

66

27

17

7

9

17

'e

157

, 13

15

9

32

8

6

7

7

84

» 14

9

2

7

1 8

, 15

3

2

5

Total

. 400

387

379

221

159

233

225

27

2031

Flowering began before 5 A.M. and continued throughout the day,
a very small number opening after 7 P.M. The period of most intense
flowering was before n A.M., the fewest flowers opening between i and
3 P.M.

He found a maximum of 23 flowers of an ear opening on one day,
and observed that the flowering of single ears of an ordinary Hungarian
wheat was completed in 4 or 5 days, the greatest number of flowers open-
ing on the third or fourth day.

THE FLOWER 129

Although self-pollination usually occurs just before or at the time
of anthesis, cross-pollination is possible, since the glumes remain separated
and the flower exposed for 15 to 20 minutes and liable to the introduction
of foreign pollen either by the agency of the wind or through transference
by small insects, especially thrips, which are frequently found in the
flowers.

More than 1 500 kinds of wheat have been grown annually at Reading
during several years, and although self-fertilisation is found to be the rule,
five or six natural hybrids are met with nearly every year. These are most
frequent among wild T. dicoccoides and some forms of T. vulgar e, but I
have observed them occasionally in T. turgidum, T. durum, T. polonicum,
and T. Spelta. I have never seen a case of natural crossing in T.
monococcum or T. dicoccum.

Flowers from which the anthers have been removed before maturity
frequently produce grain when left unprotected. Salmon in S. Dakota
found that over 76 per cent of emasculated flowers left without cover in
the field produced grain, but he obtained no grain from a similarly
emasculated ear enclosed in a paper bag.

Obermayer emasculated 29 flowers of an ear of Squarehead wheat
and removed all the rest from the ear. Twenty of the emasculated flowers
produced grain ; 15 out of 25 flowers of an ear of Hungarian wheat
similarly treated " set " grain and 28 out of 34 of an American variety.

Leighty and Hutcheson also obtained 507 grains from 1240 emascu-
lated and unprotected flowers (about 41 per cent) of 70 ears of wheat,
and only two grains from 388 similarly treated flowers (about -5 per cent)
covered with paper bags. In another series of experiments they observed
the production of grain by more than 83 per cent of the emasculated and
uncovered flowers, and less than i per cent in the case of emasculated
flowers enclosed in paraffined paper bags.

FERTILISATION AND DEVELOPMENT OF THE GRAIN

I have not succeeded in inducing the development of the pollen-tube
artificially, away from the flower, but in i^ to 2 hours after normal
pollination the pollen-tube appears, its tip turns towards the style and
pushes its way between the cells into the intercellular space in the centre
of the style, along which it grows rapidly downwards towards the ovule.
In the cases I have examined I found that the two curved male gametes
travel into the pollen-tube before the oval tube nucleus (Fig. 103), and
very frequently the latter never leaves the pollen grain.

The tube, on reaching the cavity of the ovary, finds its way to the
micropyle of the ovule, through which it passes, and finally comes into
contact with the egg apparatus. About the time of fertilisation the

K

130 THE WHEAT PLANT

synergidae degenerate, some residual debris from them remaining near
the ovum for a short period.

The fusion of one of the male gametes with the ovum was observed
in examples collected between 30 and 40 hours after pollination : the fate
of the second gamete I have not been able to discover with certainty.
The two polar nuclei of the embryo-sac remain in contact without fusing,
up to near the time of fertilisation. If they fuse and unite with the
second male gamete they do so very rapidly, since free nuclei resulting
from the division of the primary endosperm nucleus are found a few
hours after the union of the male gamete with the ovum, and before the
former has lost its individuality (Fig. 104).

Sax observed the union of the second male gamete with the fusion-
nucleus in " Kubanka " wheat (T. durum), but appearances in some cases
suggest that the double fertilisation process does not always take place,
at any rate in some forms of T. vulgar e.

(i.) Endosperm. — After fertilisation the ovum shows little sign of

FIG. 103. — Germination of pollen-grains, taken from the stigmas of a flower, showing
the transference of the gametes into the pollen-tube ( x 350).

change for a few hours. The primary endosperm nucleus, however,
begins a rapid division immediately ; free nuclei containing 3-5 well-
defined nucleoli appear in the embryo-sac in considerable numbers, even
before the male gamete is lost in the ovum (Fig. 104) ; these become
scattered irregularly in the thin cytoplasmic lining of the embryo-sac,
and are more numerous in the neighbourhood of the embryo. By the
time the embryo is 10-15 celled, 8-10 days after pollination, a single
continuous layer of cells lines the embryo-sac, and endosperm cells with
dense non-vacuolated cytoplasm fill up the narrow pocket-like cavity
in which the embryo lies, walls between the cells appearing first in this
region.

Six or seven days later the large vacuole of the sac is almost or entirely
filled with endosperm tissue which develops centripetally, the peripheral
cells being smaller than those nearer the centre.

The aleuron layer is not clearly marked out from the rest of the
endosperm until the grain has attained about half its final size.

(ii.) Nucellus. — While the embryo is developing within the embryo-
sac the surrounding nucellar tissue, which at first is several cells thick,

THE FLOWER 131

becomes gradually disorganised and absorbed, its disappearance being
most rapid in the region of the embryo and on the dorsal side of the ovule.
The epidermis of the nucellus, however, continues for a long time as
a living tissue, its cells dividing and expanding as the ovule grows ; the
position, size, and regular arrangement of the cells render it extremely
liable to be mistaken for the aleuron layer before the latter is distinctly
differentiated from the amyliferous cells of the endosperm. Later, its
cell-contents become disorganised, the remains of the cytoplasm tending

FIG. 104. — Section of the micropylar portion of the FIG. 105. — Longitudinal section of the

embryo-sac, showing the ovum with a male gamete embryo-sac (June 16) showing 2-

within it, four endosperm cells and p, tip of the celled embryo, endosperm cells and

pollen-tube with the second male gamete ( x 420). the large antipodals ( x 84).

to collect as a thin layer across the middle of the cell, where it is easily
mistaken for a cell wall dividing the epidermis into a double-celled layer.
Ultimately the radial walls of the nucellar epidermis are more or less
absorbed, and when the grain shrinks through the loss of water during
the ripening process the single-celled tissue collapses and its upper and
lower walls are crushed together, finally appearing in the ripe grain as a
highly attenuated layer, recognisable only with difficulty as a delicate
covering on the outer surface of the aleuron cells.

1 32 THE WHEAT PLANT

(iii.) Antipodal Cells. — As the growth of the embryo and endosperm
proceeds the antipodal cells slowly degenerate, the cytoplasm disappearing
first, debris of the large nuclei being visible for some time after the cells
have been stretched and flattened by the growth of the surrounding parts
of the ovule.

(iv.) Filling of the Endosperm Tissue. — The storage of the endosperm with
reserves begins when the tissue is completely formed, i.e. 10-14 days
after fertilisation, and continues for five or six weeks, till the grain is ripe.

The course of events in regard to the deposition of starch is readily
followed. The soluble carbohydrate from which it is formed is manu-
factured in the stems and leaves of the plant and sent along the conducting
strand which enters the base of the ovary, and traverses the raphe on the
furrow side of the anatropous ovule to the morphological base of the
latter in the upper part of the ovary ; from this point it diffuses into the
endosperm, and it is in this region away from the embryo that starch
grains are first observed in the centre of the two " cheeks," right and left
of the furrow of the ovary.

When the endosperm tissue has almost or quite completely filled the
cavity of the embryo-sac, minute starch grains are first seen near the nuclei
of the cells ; later, they are produced at many points in the cytoplasm
and increase rapidly in number and size, growing into thin lenticular
structures which for a time are all similar in diameter.

Not all the reserve cells are filled with starch at the same time nor at
the same rate. About three weeks after fertilisation those in the upper
part of the ovule are crowded with fully-formed grains ; half-way down
and along the furrow to near the base they are only partially filled with
smaller grains, while the cells in closer proximity to the embryo contain
but a few minute grains or none at all. In five or six weeks all the cells
of the endosperm contain starch, except those of the aleuron layer, and a
few layers in contact with the back of the scutellum ; ultimately, the
majority become densely packed throughout with starch grains of various
sizes, those formed in the later period of ripening remaining small.

The greatest amount and the largest grains are formed in the large
cells in the centre of the " cheeks " of the grain, the peripheral region
of the endosperm containing a small proportion of starch and a large
number of small grains.

The deposition of starch progresses from the upper part of the grain
towards the embryo in a diagonal plane, roughly parallel to the back of
the scutellum.

The part of the grain stored last lies round the upper margin of the
scutellum on the dorsal side, a region which is frequently opaque and
mealy in ripe grains, in which the rest of the endosperm is translucent
and flinty.

THE FLOWER 133

The slow, late deposition of the starch on the dorsal side and near the
embryo appears to be due to the difficulty of diffusion from the conducting
strand across the starch-laden cells near the furrow, and to the fact that
the developing embryo for a time uses for its growth some of the soluble
carbohydrates which would otherwise be converted into starch grains.

Immediately behind the scutellum a layer 3-5 cells in thickness
becomes filled with starch, and when the final ripening occurs it shrinks
and forms a thin band of crushed cells (Figs. 10, n).

Concomitant with the deposition of starch in the endosperm the cyto-
plasm and nuclei of the cells undergo considerable change. In the earliest
stages the cytoplasm fills the cells ; as the latter grow and expand it forms
a lining inside the cell walls and surrounds the large central vacuoles ;
the oval nuclei lie close to the walls, and there are no visible reserves in
the cells. Later, starch grains are formed in enormous numbers as
already described, and are packed closely in the cell-cavities.

In practically all the endosperm cells in which starch is accumulated
the protoplasmic contents die and become more or less disorganised.
The nuclei lose their nucleoli and clear outline and show other signs of
degeneration even before the cells are crowded with starch grains. Sooner
or later they become squeezed into the spaces between the densely
packed grains, and when stained appear in optical section as irregular
stellate or reticulate structures, coarse at first when the interstices are
comparatively wide, but in the dessication which occurs during ripening
the starch grains are forced closer together and the nuclear star or reticulum
then appears composed of thin rays of tangled strands.

In the parts previously mentioned as the last in which starch is
accumulated the nuclei are not so severely crowded, and some of them
may retain their round or oval form even up to the time of ripening of
the grain ; few, however, escape the crushing described.

The rest of the protoplasmic contents of the cells is similarly forced
into the spaces between the starch grains, the latter then appearing to be
imbedded in a matrix of protein.

(v.) The Development of the Embryo. — After a rest of a few hours the
fertilised ovum divides into two cells by a transverse wall, the large basal
suspensor cell apparently undergoing little or no further division.
Brenchley states that the first division takes place about five days after
pollination ; it is doubtless influenced by atmospheric temperature and
other factors, for I find that it frequently occurs about forty to fifty hours
after pollination. The upper cell, from which the embryo develops,
divides by a wall parallel to the first, and then by a wall at right
angles through both the cells produced, giving rise to a five-celled embryo
(Fig. 1 06).

By further division in all directions the latter rapidly increases in size,

'34

THE WHEAT PLANT

and in a few days a central core of cells with a surrounding dermatogen

layer is recognised (4, Fig. 106).

Soon the embryo becomes a club-shaped mass of cells with a narrow

elongated base, and after reaching a length of -3- -4 mm. the first sign of

differentiation is visible ; a lateral notch or depression within which the

growing -point is situated («,
Fig. 1 06) appears on the side
of the embryo opposite to the
endosperm.

From the distal portion of
the embryo above the notch
the scutellum arises ; this is a
terminal structure, which I con-
sider the cotyledon.

By the time the embryo
has reached a length of about
•5 mm. the coleoptile appears
in the form of a ring surround-
ing the stem apex, the upper
half of the ring developing
more rapidly than the lower
part, and appearing in longi-
tudinal section as a curved
scale covering the growing-
point. Later, when the embryo
is about i mm. long, the rudi-
mentary first foliage leaf arises,
the plumule consisting at this
6 stage of (i) the short and some-

FIG. 1 06.— Longitudinal sections of young em- what cone - shaped coleoptile
bryos. a, Growing-point ; 1-5 ( x 210) ; 6, with a circular terminal open-
ing, (2) the rudimentary foliage
leaf alternating with the coleoptile, and (3) the stem apex within.

Growth of the epiblast now begins.

After this stage rapid intercalary growth commences at the base of the
scutellum near the point of origin of the coleoptile. The upper part of
the scutellum which I regard as the blade of the cotyledon is lifted away
from the plumule, thus allowing for the subsequent unhindered growth
of the latter.

The rapidly lengthening portion represents the sheath of the scutellum,
the whole development being similar to that of the foliage leaves of grasses
generally.

At first the sheath of the scutellum is thin ; later it becomes thicker,

THE FLOWER

135

and a short overhanging protuberance arises at the end of the scutellum
blade (v, Figs. 10, 107) ; this structure, which is sometimes termed the
" cotyledonary " sheath, I regard as the ligule of the scutellum.

The epithelial layer of the scutellum does not begin to assume its
final form until the extension of the scutellum sheath is completed.

Differentiation of the primary root commences when the embryo is
about -5 mm. long.

Before the embryo is i mm. long the plerome, periblem, and root-cap

a

345 6 7

FIG. 107. — Longitudinal section of embryos in different stages of development ( x 50).
a, Upper portion of the scutellum (blade of the cotyledon) ; x, intercalary zone of
scutellum (sheath of the cotyledon) ; v, " ventral " scale (ligule of the cotyledon) ;
c, coleoptile ; /, first foliage leaf ; e, epiblast.

Length of embryos, i, -25 mm. ; 2, -25 mm. ; 3 and 4, about -5 mm. ; 5, -9 mm. ;
6, i-i mm. ; 7, 2 mm.

are clearly marked out and can be traced back to two or three initial cells,
situated in the centre of the undifferentiated mass of tissue which forms
the lower half of the embryo (Fig. 108). At this stage the coleoptile and
the rudiments of the first foliage leaf are visible round the stem apex.

Through division of the meristem at the root-tip the length of the root
increases and the tissue of the periblem and plerome is augmented at a
different rate from that of the root-cap and the surrounding ground tissue,
with the result that the adjacent cell walls in this region are torn asunder
and a schizogenous cavity is formed round the tip of the root (Fig. 108).

By further growth in length of the root the cavity is extended and

136

THE WHEAT PLANT

ultimately appears in the completely formed embryo of the grain as a
cylindrical space surrounding the root (Fig. 10), the ground tissue of the
lower part of the embryo, within which the root has developed, now
constituting the coleorhiza or root-sheath.

(vi.) Integuments and Pericarp. — During the growth of the ovule and
seed the two integuments undergo considerable change. The outer
coat, like its inner companion, at first consists of two cell-layers (c, d,
Fig. 109).

About the time when the fertilised ovum commences its division
it begins to degenerate ; its cells lose their turgidity, the cytoplasm
disappears, and the elongated nuclei become thinner. Later, the

x

a

FIG. 1 08. — A, Longitudinal section of young embryo ( x 175) ; c, coleoptile ; /, first
foliage leaf ; e, epiblast ; r, stem apex ; x, initial cells of the periblem of the root ;
a, schizogenous cavity ; B, longitudinal section of the root and root-cap region of
the embryo ; slightly later stage than A ( x 140).

cell-contents disappear entirely, the delicate walls shrivel, and are finally
disorganised altogether.

The inner integument, however, remains, and retains its vitality until
the grain has reached the maximum size. The cells of its two layers
up to this stage of development remain quite distinct and increase in
length with the growth of the seed, the long axes of the cells of the separate
layers crossing each other at an acute angle.

A short time before the milk-ripe stage (p. 140) the cell- contents of
the outer layer gradually disappear, a semi-permeable cuticle is developed
on the outer surface of the layer and the cells collapse (3, c, Fig. 109). The
cell-cavities of the inner layer are recognisable for some time longer, and
within them, in yellow and red-grained wheats, is formed a yellow or
reddish-brown oily or resinous substance, which is partially absorbed by

THE FLOWER

137

the cell walls, and, being visible through the more or less translucent
pericarp, gives the grain its characteristic tint.

Ultimately, the radial walls of this inner layer are disorganised or

FIG. 109. — Transverse sections illustrating the changes in the inner layer of the pericarp,
the integuments, and nucellus during the growth and ripening of the grain ( x 270). a,
" Cross layer " (p. 8) ; b, inner epidermis of the pericarp ; c, outer, d, inner integument
of the ovule ; e, epidermis of the nucellus ; /, cells of the nucellus ; g, aleuron layer.

dissolved (6, 'd, Fig. 109), and as the result of the shrinkage which occurs
during the final stages of drying and ripening of the grain the inner and
outer walls are crushed together. The two collapsed layers representing
the inner integument persist as the testa of the seed.

138 THE WHEAT PLANT

The wall of the ovary, up to the time of fertilisation of the ovum, is
composed of thin-walled parenchyma enclosed between an outer and an
inner epidermis, but changes take place in the thickness and character of
its tissues as growth proceeds.

In a half - grown grain certain portions of the wall have become
differentiated from the rest. The outer epidermis is then composed of
cells three or four times as long as broad, stretched longitudinally, with
lateral walls showing irregular beaded thickenings and a cuticle on the
external surface.

The inner epidermis consists of long thin-walled cells parallel with
those covering the exterior.

Between these outer and inner layers is loosely compacted parenchyma,
in transverse section, three or four cells across. The tissue is colourless
with the exception of a differentiated chlorophyllous " cross layer " one
cell thick, which lies immediately within the inner epidermis. The cells of
this layer are cylindrical, five or six times as long as broad, with rounded
ends, their long axes arranged at right angles to the long axis of the
grain ; each cell has an elongated nucleus and vacuolated cytoplasm, in
which are imbedded a number of chloroplasts about 4 p, in diameter,
which usually contain minute starch grains measuring, when fully grown,
about 3 /J, by i -4 p.

Chloroplasts are found also in some of the outer subepidermal cells in
the ventral furrow of the young pericarp.

In Abyssinian purple-grained wheat, the cell-sap of the cross layer
and a few of the cells of the parenchyma adjacent to it contain anthocyan.

As the grain matures the minute starch grains (1-2 p in diameter)
present in the parenchyma of the young pericarp disappear gradually
from the apex of the ovary towards the base, remaining longest in the
portion covering the embryo and in the ventral furrow : the cuticle and
thickening of the outer epidermal cell walls then become more marked.

The cells of the inner epidermis do not keep pace with the increase
in area of the inner surface of the ovary, and are torn asunder, appearing
in the ripe pericarp as a series of isolated sinuous cells — the " tube-cells "
of Vogl (Fig. 5) — which in some parts are widely separated from each
other.

The cells of the " cross layer " lose their green colour, their contents
disappear, and their walls increase greatly in thickness and develop narrow
transverse pits (Figs. 3, 109).

The rest of the parenchyma of the pericarp shrivels, and ultimately
forms an irregular crushed layer between the strong epidermis and the
thick-walled " cross layer " (Fig. 3).

THE FLOWER

RIPENING OF THE GRAIN

The ovary of the fully developed flower before fertilisation resembles
a blunt inverted cone, and is about i mm. long and i mm across the
broad stylar end (Fig. no).

After fertilisation it increases in volume from day to day, reaching

FIG. no. — Grain showing changes in size and form during development ( x 2-7).
a, June 27 ; b, July 4 ; c, July 12 ; d, August i ; e, August 15 (ripe).

its maximum size in four or five weeks, after which it shrinks through
the loss of water during the next three or four weeks until the grain is
ripe (Fig. no).

The following measurements record the changes in length, breadth,
from side to side, and thickness measured from dorsal to ventral side of
developing grains.

" SWAN " WHEAT (T. vulgare}

IQI4-

Length.

Breadth.

Thickness.

mm.

mm.

mm.

June 27 .

4

3

2

July 4 ....

6-75

3'5

2'5

12 ....

7

4'5

3'5

19 ....

7

4'5

3'5

August i .

775-8

5-25

4

„ 8. . . .

7-75-8

4-5-4-8

4

J5 (nPe)

8-6

4-1

3'5

In the early stages of ripening the length, breadth, and thickness all
increase more or less uniformly, but in the later stages the length increases
while the diameters are reduced, the grain becoming longer and narrower.

In addition to a change of volume and form there is a gradual altera-
tion in the weight, water content, and the amount of dry matter in the grain
as ripening proceeds. These variations are given below.

140

THE WHEAT PLANT

1916.

Volume of
too Grains.

Weight of
too Grains.

Water
Content.

Dry
Matter.

c.c.

gr.

per cent.

per cent.

July 7

3'36

2-84

70-61

29-39

H

4'54

4^5

68-97

3I>03

21

579

6-00

60-86

39-I4

28

6-42

7*37

52-66

47-34

Aug. 4

5-86

7-66

4075

59-25

ii

3-95

4-86*

i5'4i

84-59

18

3-47

4-63*

11-93

88-07

* Ripe grain from sheaves in the field.

The volume and weight both increase until a maximum is reached
and then decrease ; the volume was greatest in the grains examined on
July 28. The water content decreased from 70 to about 50 per cent
during July ; at the time of cutting (August 7) the amount present
was 40 per cent, that in the grain taken from sheaves two to three days
after cutting, 15 per cent, while it was reduced to 12 per cent in the grain
of the crop cut and left in the field ten to twelve days.

The dry matter increased from the beginning to the end of the develop-
ment and ripening of the grain.

The weight of the individual grains also continued to increase up to
August 4, a few days before harvest, but rapidly fell after cutting, through
loss of water.

In the week July 28 to August 4, the weight of the grain increased,
while the volume and water content decreased ; the migration of starch
and other substances into it during this period more than balanced the loss
of water.

These changes in developing grains exhibit the same general pro-
gression each season, but investigations carried on during several years
show that they do not always proceed at the same rate, the loss of water
and the accumulation of dry matter being accelerated or retarded by
variations in the amount of sunlight, and the temperature and dryness
of the atmosphere.

Although the ripening changes of the plant proceed gradually without
any definite break in continuity, four stages may be recognised, viz. the
milk-ripe, yellow-ripe, ripe, and dead-ripe stages.

In the milk-ripe stage the crop has a green appearance. The lower
leaves of the plants are dead, but the blades of the three upper leaves, the
higher internodes, and the ears are living and green ; only on the edges
or tips of the leaves are there any visible signs of ripening in the form of
yellowish spots and stripes.

THE FLOWER 141

The leaf-sheaths are green and their swollen nodal portions plump
and sappy.

The glumes and pericarp of the grain are also green, the chlorophyllous
layer of the latter showing through the outer colourless tissue.

At this stage the grain has attained its maximum volume and highest
water content ; the vacuoles of the endosperm tissue contain watery sap,
and starch grains are abundant in the cells ; from it can be squeezed a
milk-like liquid whose whiteness is due to numerous starch grains present
in it.

The several parts of the embryo are completely differentiated, but
have not quite reached their full development, and although grains
harvested now germinate readily, the young plants are somewhat weak.

In the yellow-ripe stage the crop has changed to a golden tint except
in the purple-strawed forms which are a pinkish purple colour. The
straw is smooth and shining, tough and pliable.

The chlorophyll has disappeared from all parts of the leaves, except
the thick nodal portions of the upper leaf-sheaths, which still remain
swollen and green, those of the lower leaf-sheath being shrunken and
brownish.

The glumes have assumed their characteristic ripe tint.

As ripening proceeds, the chlorophyll in the pericarp of the grain
gradually disappears, first from the upper part and dorsal side, and later,
from the lower part and the furrow ; in the yellow-ripe stage it is no longer
to be seen.

The grain can be crushed between the thumb and finger-nail, the
contents, however, are not milky, but soft, and they knead like dough.

This is the best stage of ripeness in which a crop of wheat should be
cut ; assimilation is at an end, and there is no gain in weight by leaving
it longer. The danger of losing some of the grain by " shattering " is
also reduced by harvesting at this period of development.

The ripe stage is reached in dry, clear weather thpee or four days after
the yellow-ripe stage. The thick basal portions of the leaf-sheaths are
now dry and shrunken.

The grain readily parts from the rachilla and is liable to shake out of
the glumes. It is firm, and though it may be dented by pressure of the
thumb-nail, it is not easily crushed. Its characteristic colour has become
more distinct, the yellow grains paler, the red grains somewhat darker,
and the flinty or mealy character of the endosperm is clearly emphasised.

In the dead-ripe stage the straw becomes dull, more and more brittle
and dirty the longer it is left uncut.

The axis of the ear is liable to become brittle and in some forms the
ears fall off or break in pieces ; the grain " shatters " easily, and much is
lost in harvesting the crop if left as late as this.

142 THE WHEAT PLANT

The grain is hard and breaks into angular fragments when crushed.

If damp weather prevails the plant is soon discoloured by the growth
of fungi.

The time which elapses between the escape of the ear from the upper
leaf-sheath and the ripening of the grain varies with the kind of wheat,
the date of appearance of the ear, and the climatic conditions of the locality
where the crop is grown.

It is somewhat remarkable that many late forms with a long, vegeta-
tive period of growth tend to develop their grain more quickly than the
early or rapid-growing forms, the latter also appearing to require a higher
temperature and greater intensity of sunlight for the process than the
former.

Sown in the ordinary season in autumn or spring at Reading, the time
between " earing " and ripening is usually from 55 to 63 days. By,
reference to the tables on pp. 86-88, however, it is seen that the period
may extend to 80 days or more in the case of certain wheats sown very
early or very late, especially in seasons (such as 1912) which prove to be
dull, wet or cool in late summer.

By sowing at abnormal times the production of ears may be delayed
until the end of September or later, but in such cases the grain does not
ripen, as the growth is checked by the low temperature and diminished
light of autumn.

Wheats ripen best at Reading between the last week in July and the
second week in August.

The earliest date at which ripening occurs in any wheat at Reading
is during the last ten days of July, the ears in such cases escaping from
their sheaths about the first week of June.

The latest period is about the middle of September, and to ripen by
that time the ears must appear not later than the early part of July.

Late or autumn forms of wheat at Reading must be sown before the
middle of March, or they rarely ripen grain in that season, although when
sown as late as April 5 they develop ears in due course in August or
September.

Most early or spring forms must be sown before the middle of April
or they will not ripen grain, although some of them produce ears in late
summer if sown as late as the first week of July.

Fife wheat sown at Reading between August and the first week of
December usually ripens in the first ten days of the following August ;
sown in January, February, or March, ripening occurs between the middle
and the end of August (p. 86).

The later forms of English autumn wheats sown between August and
December ripen about the middle of August ; sown in January or
February the ripening may be delayed until the end of August or

THE FLOWER

beginning of September. Some groups of wheat are intermediate between
these in their ripening period.

There is a close correlation between the point at which the first flowers
open on an ear of wheat and the region in which the heaviest and largest
grains are found. The point differs a little with the race and form of
wheat.

In T. vulgar e the heaviest spikelets and grains occur in the middle
third of the ear, in winter wheats not far from the union of the middle
and lower third, in spring wheats slightly higher and nearer the middle
of the ear.

The average 100- grain weight increases up to the point where the
heaviest grains are found, and decreases upwards to the tip.

The average weight of the grain is higher in the lower than in the upper
half.

In spikelets with more than three grains the second grain is generally
the heaviest, and the third and fourth lighter than the first.

PART II

145

CHAPTER IX

CLASSIFICATION

PRE-LINNEAN botanists of the sixteenth and seventeenth centuries
generally adopted the classification of the cultivated wheats suggested by
Columella, dividing them into two sections, namely :

I. Species of Triticum, or wheats whose ears have a tough rachis and
grain so loosely invested by the chaff that they fall out when the ears are
thrashed, and

II. Species of Zea, whose ears possess a fragile rachis, which breaks
into short lengths, and whose grains are so firmly enclosed by the glumes
that they are separated from the latter with difficulty.

Those comprising the Zea section are at the present day often spoken
of as " spelt" wheats, the term " spelt " being 'used in a generic sense
and embracing Small Spelt (T. monococcum), Emmer (T. dicoccum), and
Common Spelt (T. Spelta).

Linnaeus, however, placed all the cultivated wheats under the single
genus Triticum, of which in the first edition of his Species Plantarum
(1753) he mentions five species, viz. :

1. T, aestivum (Bearded Spring Wheat).

2. T. hybernum (Beardless Winter Wheat).

3. T. turgidum.

4. T. Spelta — Zea dicoccos vel spelta major of C. Bauhin.

5. T. monococcum — Zea Briza dicta, vel monococcus germanica of C. Bauhin.

In the second edition of his Species Plantarum (1764) he added T.
polonicum, adopting Plukenet's specific name.

Later in the Supplement (1781) his son introduced the species T.
composition, a form of T. turgidum with proliferous ears (2, Fig. 160).

In 1786 Lamarck (Encyclop. Me'thodique, vol. ii. p. 554) recognised five
species, viz. :

i. T. sativum, embracing the Linnean species T. aestivum, T. hybernum,
and T. turgidum.

2. T. compositum, L. fils.

3. T. polonicum, L,.

4. T. Spelta, L.

5. T. monococcum, L.

148 THE WHEAT PLANT

In the following year (1787) Villars, in the Histoire des plantes de
Dauphine (vol. ii. p. 153), referred all the wheats to seven species, viz. :

1. T. vulgare (T. aestivum, L.).

2. T. touzelle, Gouan hort. 57 (T. hybernum, L.).

3. T. turgidum, L.

4. T. maximum (a wheat resembling T. polonicum).

5. T. compositum, L.

6. T, Spelta, L. | 7. T. monococcum, L.

Schrank in 1789 (Baier. Fl. vol. i. p. 387) recognised only two species,
viz. :

1 . T. cereale, with the two varieties : (a) aestivum, (f3) hybernum.

2. T. Spelta.

He says that he thinks " Emmer which is cultivated in Wurtemberg
belongs to this, if it is not a distinct species ; I call it * dicoccon."

Host in 1805 (Gram. Austr. vol. iii.) extended the list to eleven species,
viz. :

i . T. vulgare ( = T. aestivum and T. hybernum).

2. T. compositum, L.

3. T. turgidum, L.

4. T. Zea ( = T. Spelta, L.)

5. T. Spelta ( = T. amyleum, Ser.).

6. T. polonicum, L.

7. T. monococcum, L.'

and added in 1809 (vol. iv.)

8. T. hordeiforme (a form of T. durum, Desf.).

9. T. villosum (a pubescent white-glumed T. durum, Desf.).

10. T. compactum.

11. T. atratum (a variety of T. amyleum, Ser., with dark brown or black

pubescent glumes).

Host was the first botanist who united the Linnean T. aestivum and
T. hybernum under one species (T. vulgare). Per soon, in his Synopsis
Plantarum (1805), united them with T. durum under the name T. sativum.

In 1809 Bayle-Barelle (Mon. agr. dei cereali) described eleven species
in two sections, viz. :

SECTION I

i. T. compositum, L. fils. j 2. T. turgidum, L. | 3. T. polonicum, L.

4. T. cerulescens ( = a variety of T. durum, Desf.).

5. T. tomentosum ( = a variety of T. durum, Desf., or T. turgidum, L.).

6. T. candidissimum, Arduini ( = a red-grained variety of T. durum, Desf.,

with white glabrous and shining glumes), which he considers the
Siligo of Varro, Pliny, and Columella.

7. T. creticum silvestre ( = T. sylvestre creticum, C. Bauhin, and awnless

T. compactum, Host).

CLASSIFICATION 149

8. T. sativum, Pers. (T. hybernum, L.) This he subdivides into four varieties :

i. T. sat. varietas mutica alba.

ii. T. sat. varietas mutica alba tomentosa (T. anglicum, Arduini).

iii. T. sat. varietas ruffa aristata.

iv. T. sat. varietas ruffa mutica.

SECTION II

9. T. farrum. He classes the Zea of Galen, Dioscorides, and Theophrastus

with this and states that it is often confused with T. Spelta, L.
10. T. monococcum.
n. T. Spelta.

with three others in an Appendix, viz. :

T. bicorne, Forskal. | T. tumonia, Beguillet. | T. Blat de Caure, Spagnuoli.

In 1816 Lagasca (Gen. et. sp.pl. 82) described sixteen species, viz. :

1. T. monococcum, L.

2. T. Cienfuegos.

3. T. Bauhini.

4. T. Spelta, L.

5. T. hybernum, L.

6. T. aestivum, L.

7. T. Linneanum.

8. T. turgidum, L.

9. T.fastuosum.

10. T. Gaertnerianum.

11. T. platystachyum.

12. T. cochleare.

13. T. Cevallos.

14. T. durum, Desf.

15. T. polonicum , L .

16. T. spinulosum.

The first four were " spelt wheats " with a fragile rachis, the last on
the list possessing long foliaceous glumes presumably like those of
T. polonicum.

Clemente, in an edition of Herr era's Agricultura general (Madrid, 1808),
described twenty species.

Eight of them (Nos. 3, 4, 12, 14, 15, 16, 17, and 19), although given as
his own species, bear the same name as those published by Lagasca two
years previously : five, namely, Nos. 2, 6, 7, 9, and n, are new.

1 . T. monococcum, L. Pequena Escafia o escana menor lampino.

2. T. Horneman. A pubescent T. monococcum. Pequena Escana velloso.

3. T. Cienfuegos. A form of T. Spelta. Escana melliza 6 de dos Carreras :

escandia ; esprilla bassona.

4. T. Bauhini. A form of T. Spelta with dense ears. Escana mazzoral.

5. T. Spelta, L. Bearded. Escanda lampina ; escana grande.

6. T. Forskal. A pubescent T. Spelta. Escanda villosa ; escana mayor

peluda.

7. T. Arias. Red and white awnless T. Spelta. Escanda mocho.

8. T. hibcrnum, L. Beardless T. vulgar e. Chamorro comun, Pelon mocho.

9. T. Koeleri. An awnless pubescent T. vulgare. T. sativum hybernum

sardonicum of Koeler and others. Chamorro velloso.

150 THE WHEAT PLANT

10. T. aestivum, L. Bearded spring T. vulgare. Candeal lampino, Treme-

sino, Hembrilla, Jeja.

11. T. Horstianum. A bearded pubescent T. vulgare. Candeal velloso.

12. T. Linneanum. A form of T. turgidum. Redondillo lampino.

13. T. turgidum, L. Redondillo velloso.

14. T. Gaertnerianum. Fanfarron lampino.

15. T.platystachyon. Chapado 6 chapado lampino.

1 6. T. cochleare. Cuchareta 6 chapado villosa.

17. T. Cevallos. Moro 6 moruno lampino.

1 8. T. durum, Desf. Moruno 6 moro velloso.

19. T.fastuosum. Fanfarron velloso ; Dicho.

20. T. polonicum, L. Trigo polonia 6 Polago.

In 1817 Roemer and Schultes (System, veg.) recognised twenty-one
species, most of them from Lagasca and Host, the only new one being
T. siculum, a species allied to T. hordeiforme, Host (T. durum, Desf.).

In his Characteristica et descriptiones Cerealium (1818) Schiibler gives
the following sixteen species, which he groups in two sections :

SECTION I. — With loosely invested grain

1. T. muticum :

(a) aestivum. (b) hybernum, L.

2. T. aristatum :

(a) aestivum, L. | (b) hybernum.

3. T. sibiricum. Siberian spring wheat, an early T. vulgar e(!).

4. T. velutinum.

5. T. compactum, Host :

(a) aristatum. Bengelweizen awned.

(b) muticum. Bengelweizen awnless.

6. T. turgidum, L.

7. T. hordeiforme, Host. (A form of T. durum, Desf.)

8. T. durum, Lagasca. Arabian winter wheat : a form of T. durum, Desf.

9. T. siculum, Schmidt. (Sicilian wheat.)

10. T. compositum, L. | n. T. polonicum, "L.

SECTION II.— With " Spelt " ears
12. T. Spelta :

(a) mutica.

a. alba.
ft. rubra.

(b) aristata.

(c) velutina.

(d) aestiva.

13. T. monococcum, L.

14. T. dicoccum.

(a) album. | (b) rufum.

15. T. atratum, Host.

1 6. T. tricoccum.

Seringe in 1818, in his Melanges botaniques and Monographic des
ceriales de la Suisse, described eight species in two sections, viz. :

CLASSIFICATION 151

SECTION I. — FRUMENTA

1. T. vulgar e, 3. T. durum.

2. T. turgidum. 4. T. polonicum.

SECTION II. — SPELTAE

5. T. Spelta. | 6. T. amyleum. \ 7. T. monococcum.

S. T. venulosum. An apparently immature specimen collected in Egypt
and sent to Seringe by Professor Desfontaines. Seringe considered
it allied to T. monococcum, but larger with anastomosing prominent
veins ; probably a form of Indo- Abyssinian T. dicoccum.

In 1824 Metzger, in his Europaeische Cerealien, adopted the species
recognised by Seringe in his Monographic des cereales de la Suisse, only
leaving out the doubtful T. venulosum, and in 1827 Link (Hort. berol.)
adopted the same classification with the addition of T. compactum.

Later, Seringe, in his Cereales europeennes (1841-42), classified the eight
species of wheats of his earlier monograph under the three genera
Triticum, Spelta, and Nivieria thus :

Genus TRITICUM —

1. T. vulgare, Willd. (This included T. compactum, Host.)

2. T. turgidum, L. | 3. T. durum, Desf. | 4. T. polonicum, L.

Genus SPELTA —

5. S. vulgaris. 6. S. amylea.

Genus NIVIERIA —
7. N. monococcum.

8. N. (?) venulosa.

Vilmorin in 1850 (Catalogue methodique et synonymique des froments)
followed the classification of Seringe and Metzger, recognising seven,
species, viz. :

1 . T. sativum.

2. T. turgidum.

3. T. durum.

4. T. polonicum.

5. T. Spelta.

6. T. amyleum.

7. T. monococcum.

Alefeld in 1866 (Landwirtschaftliche Flora) included all wheats under
the single species T. vulgare, except Polish wheat, which he placed in a
separate genus, Deina (D. polonica).

The species T. vulgare he divided into nine " varietal groups," viz. :

1. T. vulgare durum.

2. T. vulgare turgidum.

3. T. vulgare compositum.

4. T. vulgare compactum.

5. T. vulgare muticum.

6. T. vulgare aristatum.

7. T. vulgare dicoccum.

8. T. vulgare monococcum.
9 T. vulgare Spelta.

152 THE WHEAT PLANT

These groups are again subdivided into varieties, to each of which he
gives a triple name.

Kornicke in 1885 (Die Arten und Varietaten des Getreides, vol. i.)
recognised three species of wheat, viz. :

1. Triticum vulgare, Vill. . . . Common wheat.

2. Triticum polonicum, L. . . . Polish wheat.

3. Triticum monococcum, L. . . . Einkorn or small Spelt.

He divides T. vulgare into six sub-species, viz. :

I. Rachis tough : grains easily separated on thrashing.
Sub-species —

1 . Triticum vulgare, Vill. . . . Common wheat.

2. Triticum compactum, Host . . Dwarf wheat.

3. Triticum turgidum, L. . . . English wheat.

4 Triticum durum, L. Hard (or Macaroni) wheat.

II. Rachis fragile : grain firmly enclosed in the glumes.
Sub-species —

5. Triticum Spelta . . . . Common Spelt.

6. Triticum dicoccum, Schrk. . . Emmer.

E. Hackel (Engler and Prantl's Nat. Pfl. vol. ii. pp. 2, 81, 84 [1887])
separated the genus Triticum into two sections :

Section I. : Aegilops, with rounded glumes, not keeled or only faintly so.
Section II. : Sitopyros, with sharply keeled glumes.

In the latter section, which embraces the cultivated wheats, are
included three species, viz. :

i. T.monococcum. 2. T. sativum. 3. T. polonicum.

T. sativum is divided into several races as follows :

I. Rachis fragile and grain firmly invested by the glumes.

i°. Ear lax, quadrate, empty glumes with blunt keel :

(a) T. sativum Spelta.

2°. Ear denser, compressed, empty glumes sharply keeled:

(b) T. sativum dicoccum.

II. Rachis tough : grain loosely invested by the glumes.

(c) T. sativum tenax.
(c) T. sativum tenax is divided thus :

a. T. sativum vulgare . . . (T. vulgare, Vill.).

/B. T. sativum compactum . . . (T. compactum, Host),

y. T. sativum turgidum . . . (T. turgidum, L.).

5. T. sativum durum . . . (T. durum, Desf.).

CLASSIFICATION 153

A classification similar to that of Hackel is adopted by Ascherson
and Graebner (Syn. Mitteleur. Fl. vol. ii., 1901).

The classification adopted by most of the early authors is highly
artificial, being based chiefly upon the characters of the ears and grains
alone, and often had reference to comparatively few forms, or to those
confined to one or two limited wheat-growing regions.

In order to obtain a wider view of the subject, and to discover the
natural relationships of the cultivated wheats, I have critically examined,
during the last twenty years or more, living specimens from all parts of
the world.

Nearly two thousand forms have been grown annually side by side,
and their morphological characters in the young and mature states, as
well as their habit of growth, ripening period, susceptibility to the attacks
of fungi, and other characters, have been investigated and compared.

In determining the limits of the several races, I have not only taken
into account the usual morphological characters of the inflorescences,
but have given especial weight to the characters of the leaves and vege-
tative organs generally, being convinced that these have undergone little
change, and indicate natural genetic relationships more clearly than the
ears and grain, the modification of which has been the special object of the
cultivator from the earliest date.

I find, for example, that the length, amount, and arrangement of the
hairs on the young leaf-blades are much more constant and character-
istic features of a race than the size, shape, and colour of its ears and grain.

I have also taken into consideration anatomical features and the
results of hybridisation, as well as archaeological evidence, and the dis-
tribution of endemic forms in the older wheat-growing areas of the
Eastern Hemisphere.

Two wild species of wheat are known, and the cultivated wheats, none
of which are found wild, fall into eleven natural groups. To these
groups I have applied the term " race " rather than the term " species,"
although they might with equal justice be designated " cultivated species,"
for the methods used in their grouping and delimitation are the same as
those adopted in the case of wild species.

My conclusions regarding the number and classification of the races
of wheats are indicated below.

With the exception of the addition of the three new races, III., VII., ,
and X., the list and the chief distinguishing characters of the groups,
whether termed " races " or " species," agree closely with those given
by Seringe and almost all authorities since his time.

SPECIES I. T. aegilopoides, Bal. . . Wild Small Spelt.
RACE I. T. monococcum, L. . Small Spelt.

154 THE WHEAT PLANT

SPECIES II. T. dicoccoides, Korn. . . Wild Emmer.

RACE II. T. dicoccum, Schiibl. . Emmer.

III. T. orientate, mihi . . Khorasan Wheat.

IV. T. durum, Desf. . . Macaroni Wheat.
V. T. polonicum, L. . . Polish Wheat.

VI. T. turgidum, L. . . Rivet or Cone Wheat.

VII. T. pyramidale, mihi . . Egyptian Cone Wheat.

VIII. T. vulgare, Host . . Bread Wheat.

IX. T. compactum, Host . . Club Wheat.

X. T. sphaerococcum, mihi . Indian Dwarf Wheat.

XI. T. Spelta, L. . . . Large Spelt or Dinkel.

DIAGNOSTIC CHARACTERS OF THE SPECIES AND RACES OF WHEATS

SPECIES I. — Triticum aegilopoides, Bal. Wild Small Spelt.

Coleoptile, 2 -nerved.

Young shoots, erect or prostrate ; young leaves blue-green or yellow-green,
more or less hairy, with a line of long hairs on the summit of the longitudinal
lidges.

Straw, slender ; nodes clothed with white deflexed hairs.

Ear, bearded, compressed, very narrow across the face, broad across the 2-
rowed profile ; spikelets i- (or 2-) grained ; terminal spikelet minute and abortive ;
rachis very fragile and fringed along the margin with long straight hairs.

Empty glume, long, narrow, keeled to the base with stout, acute, apical tooth
often turned slightly outwards ; on the outer face a prominent nerve ending
in a distinct secondary tooth some distance from the base of the apical tooth.

Palea, in ripe ears divided longitudinally into two halves.

Grain, small, rice-like, flinty, pointed at both ends, compressed from side to
side, furrow indistinct ; tip with few hairs.

RACE I. — Triticum monococcum, L. Small Spelt : Einkorn : Engrain.

Coleoptile, 2-nerved.

Young shoots, erect or semi-erect ; young leaves yellowish-green, with very
short hairs or scab rid projections on the longitudinal ridges.

Straw, slender ; nodes clothed with deflexed hairs.

Ear, bearded, compressed, much narrower across the face than the 2-rowed
profile ; spikelets i- (or 2-) grained ; terminal spikelet minute and abortive ;
rachis fragile, glabrous or edges fringed with short hairs.

Empty glume, long, narrow, keeled to the base with a stout, acute, apical
tooth, straight or turned slightly inwards ; the prominent nerve on the outer
half ends in a distinct secondary tooth some distance from the apical tooth.

Palea, divided longitudinally to the base when the ear is ripe.

Grain, as in T. aegilopoides, but shorter.

SPECIES II. — Triticum dicoccoides, Korn. Wild Emmer.
Coleoptile, usually 4-nerved.

CLASSIFICATION 155

Young shoots, prostrate ; young leaves blue-green, clothed with soft hairs of
nearly equal length ; culm leaves yellowish -green.

Straw, slender, solid or hollow with thick walls ; nodes clothed with de-
flexed hairs.

Ear, bearded, compressed, much narrower across the face than the 2 -rowed
profile ; spikelets i- to 2 -grained ; terminal spikelet large and frequently fertile ;
rachis very fragile, usually fringed along the margin with long straight hairs.

Empty glume, long, narrow, keeled to the base ; apical tooth blunt or acute ;
•the strong nerve on the outer half converging towards the base of the apical tooth
and ending in a short secondary tooth.

Palea, not divided longitudinally.

Grain, very long (9-12 mm.), narrow, and flinty, somewhat triangular in
section, with a conspicuous tuft of white hairs at the apex.

RACE II. — Triticum dicoccum, Schiibl. Emmer.

Coleoptile, in Indo-Abyssinian Emmers usually 4- to 6-nerved ; in European
Emmers 2 -nerved.

Young shoots, usually erect ; young leaves with soft hairs of nearly uniform
length.

Straw, solid or hollow with thick walls ; Indo-Abyssinian forms short,
European forms taller.

Ear, bearded, compressed, narrower across the face than the 2-rowed pro-
file ; spikelets generally 2-grained ; rachis fragile or tough, narrow, fringed
with short hairs.

Empty glume, long and narrow, the outer face flat ; keel prominent from tip
to base ; apical tooth in Western European forms acute and curved as in T.
durum ; in the Russian. Indian, and Abyssinian forms short and blunt.

Grain, flinty or semi-flinty, 7-9 mm. long, narrow, pointed at both ends,
ventral side flat or slightly concave ; cross section more or less triangular.

RACE III. — Triticum orientale, mihi. Khorasan Wheat.

Coleoptile, 2-nerved.

Young shoots, erect ; young leaves narrow, pubescent, hairs of nearly uniform
length.

Straw, of medium height, solid or hollow with thick walls.

Ear, bearded, very lax, almost square ; rachis tough ; awns scabrid to the
base, more or less deciduous.

Empty glume, long and narrow ; apical tooth blunt.

Grain, white, very long (10-5-12 mm.), narrow, flinty.

RACE IV. — Triticum durum, Desf. Macaroni Wheat.

Coleoptile, 2-nerved.

Young shoots, erect ; young leaves quite glabrous or nearly so.
Straw, tall, solid or hollow with thick walls.

Ears, usually bearded, square in section or narrower across the face than the
2-rowed profile ; bearded, stiff, usually erect ; rachis more or less fragile ; the

156 THE WHEAT PLANT

awns of great length, diverging slightly, and almost smooth at the base ; spikelets
longer and narrower than those of T. turgidum or T. vulgar e.

Empty glume, long and narrow, the outer face flattish ; keel curved, pro-
minent from tip to base ; apical tooth stout, generally acute and usually curved
inwards ; glumes somewhat readily detached from the rachis.

Grain, long and narrow, hard, more or less pointed at both ends, with a
prominent dorsal ridge, endosperm flinty ; somewhat triangular in section.

Transitional forms are met with having slightly hairy leaves and approximat-
ing towards T. turgidum, T. vulgare, and T. dicoccum, and one or two hybrids
are known differing only from typical T. durum in having beardless ears.

RACE V. — Triticum polonicum, L. Polish Wheat.
Coleoptile, 2-nerved.

Young shoots, erect ; young leaves blue-green, glabrous or nearly so.
Straw, tall, solid or hollow with thick walls.

Ear, bearded ; spikelets large, usually i - to 2-grained ; rachis somewhat fragile.
Empty glume, as long as or longer than the rest of the spikelet, narrow, 3-4
cm. long, keeled, with two small apical teeth.

Grain, very long (11-12 mm.), narrow, flinty, somewhat triangular in section.

RACE VI. — Triticum turgidum, L. Rivet or Cone Wheat.

Coleoptile, 2-nerved.

Young shoots, erect or prostrate ; young leaves clothed with soft hairs of
nearly uniform length.

Straw, tall, solid or hollow with thick walls.

Ear, usually bearded, square in section or narrower across the face than the
2-rowed profile ; bearded, heavy, often pendulous, and less rigid than T.
durum ; rachis tough ; awns stout, very scabrid from tip to base, frequently
deciduous ; spikelets about as long as broad, often ripening 3-4 grains.

Empty glume, short and broad, the outer face convex, keel prominent from
tip to base ; apical tooth stout, usually acute and curved ; glumes more firmly
attached to rachis than in T. durum.

Grain, generally mealy, though sometimes flinty or semi-flinty ; large,
plump, and somewhat short, with truncate apex and high dorsal hump behind
the embryo.

RACE VII. — Triticum pyramidale, mihi. Egyptian Cone Wheat.

Coleoptile, 2-nerved.

Young shoots, erect ; young leaves pubescent, hairs somewhat short ; culm
leaves yellowish-green.

Straw, very short, solid or hollow with thick walls.

Ear, bearded, dense, short, usually tapered towards the apex and oblong in
section, wider across the 2-rowed side than across the face ; rachis tough ; awns
scabrid to the base and sometimes deciduous.

Empty glume, keeled to the base.

CLASSIFICATION 157

Grain, usually mealy, short and narrow, somewhat pointed at the apex ;
dorsal hump prominent.

RACE VIII. — Triticum vulgare, Host. Bread Wheat.

Coleoptile. 2-nerved.

Young shoots, erect, semi-erect, or prostrate ; young leaves more or less hairy,
the hairs of unequal length, with a single row of long ones along the summit of
some or all the longitudinal ridges.

Straw, hollow with thin walls, though forms are occasionally found with
solid upper internodes.

Ear, bearded or beardless, square in section or more commonly broader
across the face than the 2-rowed profile, bearded or beardless ; the awns shorter
than those of T. durum or T. turgidum ; some entirely beardless, others classed
as beardless have awns of variable length on the upper spikelets ; spikelets
usually about as long as broad ; rachis tough.

Empty glume, broad, the outer face convex, keeled from tip to base or in the
upper half only ; apical tooth in the bearded forms short, acute, or sometimes
prolonged into an awn (1-4 cm. long), in the beardless forms usually short and
blunter.

Grain, very varied in form and size ; flinty or mealy ; usually plump with
bluntish apex, rounded on the dorsal side without prominent hump or ridge.

RACE IX. — Triticum compactum, Host. Club Wheat.

Coleoptile, 2-nerved.

Young shoots, leaves, straw, and glumes as in T. vulgare.

Straw, hollow, very variable in length.

Ear, bearded or beardless, short and dense from 3-5 to 6 cm. long, density
40-50 ; spikelets broad and short, often containing 3-4 grains ; rachis tough.

Empty glume, as in T. vulgare.

Grain, generally soft and mealy, plump, small, variable in shape, some with
prominent dorsal hump like those of T. turgidum.

RACE X. — Triticum sphaerococcum, mihi. Indian Dwarf Wheat.

Coleoptile, 2-nerved.

Young shoots, erect ; young leaves as in T. vulgare.

Straw, very short, stiff, erect, and hollow, usually not more than 65-70 cm.
long, except where irrigated ; culm leaves rigid and somewhat erect.

Ear, bearded or beardless, stiff, erect, 4-6 cm. long, not so dense as T. com-
pactum ; awns of bearded forms shorter and stouter than those of T. com-
pactum ; rachis tough ; length and breadth of the spikelets about equal.

Empty glume, broad and short, inflated, with strong curved scabrid apical
tooth.

Grain, very short (4-5 mm. long), flinty, often angular on account of pressure
of the glumes.

158 THE WHEAT PLANT

RACE XI. — Triticum Spelta, L. Large Spelt or Dinkel.

Coleoptile, 2-nerved.

Young shoots, erect or prostrate ; young leaves dark green with few hairs
arranged as in T. vulgare.

Straw, stout and hollow.

Ear, very lax, bearded with short awns or beardless ; rachis broad and stout,
convex on one side, flat or concave on the other, fragile, breaking transversely
below each spikelet ; spikelets narrow with 2-3 grains.

Empty glume, firm, with broad truncate apex ; apical tooth short and blunt ;
prominent lateral nerve ending in a blunt projection.

Grain, long, usually flinty, somewhat pointed at both ends, apex with tuft
of hair, ventral surface flattened or hollowed slightly, furrow shallow.

VARIETY. — The several races of wheats are subdivided into smaller
groups or varieties, these being founded upon a number of obvious
hereditary morphological differences of the ears and grain.

The presence or absence of awns, the colour, smoothness, and hairiness
of the chaff, and other characters were utilised in subdividing the races
by Seringe, Metzger, Vilmorin, and others. The scheme adopted by
Kornicke I have found most convenient and clearly defined for taxonomic
purposes, and it carries the classification further without loss of clearness
than any previous scheme.

The following are the characters on which varieties are based :

1 . Presence or absence of awns.

2. Colour of glumes (white, red, or black).

3. Colour of the awn (white, red, or black).

4. Glabrous or pubescent glumes.

5. Colour of grain (white or red).

In the races T. turgidum and T. dicoccum in which branching of the
rachis is hereditary the division into simple and compound ears is made.

While all the characters mentioned are hereditary, the extent of their
development is affected in greater or lesser degree by external con-
ditions, a fact to be borne in mind when a decision has to be made in
respect of the variety under which a particular individual is to be classed.

Beardless wheats, as explained in another place (p. 104), are either
truly awnless or the awns of the flowering glume are not more than one
or two cm. long, the bearded forms of the same race having awns three
or four times this length. In practice there is no risk of confusing the
bearded with the beardless condition.

The glume colour is constant, but in some seasons the black tint is
greatly subdued, or missing altogether, and the pale red varieties are
sometimes difficult to separate from those with white chaff.

The awns of varieties with white and red chaff are either white or red
like the glume, or black. The black pigment, however, is so variable in

CLASSIFICATION 159

its appearance in this country that plants belonging to varieties differing
only in the colour of the awn cannot be accurately classified except after
observations extending over two or three seasons at least.

Grain colour is a hereditary character subject to little variation and
usually determined without much trouble. Only in Macaroni Wheats
is it sometimes extremely difficult to distinguish the pale red flinty grains
from those which are white with yellowish translucent, endosperm.

The amount and character of the hair on the glumes is of good
diagnostic value and in most cases is readily seen. Care, however, is
needed where the hair is short and sparsely distributed.

FORM. — Wheats belonging to the same variety as defined by reference
to the scheme just mentioned differ among themselves in the length of
the ear and straw, the density of the ear, the shape of the empty glumes,
the earliness and lateness of ripening, and in other hereditary features
both morphological and physiological. For example, the variety of
T. vulgar e with beardless ears, white glabrous chaff, and red grain (var.
lutescens) includes the wheats known among farmers as Purple straw,
Red Fife, Squarehead, Briquet Jaune, Warden, and many others.

These names represent, or ought to represent, the progeny of a single
individual, the origin of which may, or may not, be known with certainty.
To each of these I apply the term form, in place of the terms variety,
type, or sort used by others.

Many attempts have been made to group these forms for purposes of
easy reference and identification, but none of the various schemes pro-
posed are of more than limited application.

In the case of the rarer varieties in which only a small number of forms
are known the task is not insuperable, and simple keys for the identifica-
tion of the several forms may be readily devised ; but in the varieties
erythrospermum (bearded ears, with white glabrous glumes and red grain),
ferrugimum (bearded ears, with red glabrous glumes and red grain),
lutescens, and milturum (the corresponding beardless varieties) of T.
vulgar e, and in lesser degree the varieties leucurum (ears with white
glabrous glumes and white grain) and hordeiforme (ears with red glabrous
glumes and white grain) of T. durum, the existing forms are so numerous
and intergrade in almost all their characters between one extreme and
another that the formation of clearly defined groups or classes is a problem
of the greatest difficulty, if not practically impossible.

The most useful way of dealing with such extensive varieties is to make
a separate classification for the forms of it cultivated in each country.
In order to accomplish this, and for purposes of identification of the
separate forms, detailed descriptions are needed. These should be made
after a study of the living plants grown side by side during several seasons
and should take cognisance of the following characters :

i6o THE WHEAT PLANT

Young Tillered Plant. — Habit of shoots, erect, sloping, or prostrate ;
colour of the leaves ; leaf-surface, glabrous or pubescent, and distribution
of the hairs on the longitudinal ridges.

Straw. — Its average height ; whether hollow or solid, rough or
smooth.

In the description of wheats given later I have adopted the following
general table of straw lengths :

Very short . . . Below 24 in.

Short .... 24-34 m-

Medium . . . 34-44 ,,

Tall . . . 44-54 „

Very tall . . . Over 54 in.

Ear. — Average length ; breadth across the face and width of the two-
ranked side : whether specially glaucous or non-glaucous and yellowish-
green. In awned forms the length of the awns ; in beardless forms
whether " tip-bearded " or truly awnless.

The general form, whether uniform in diameter, clubbed at the apex,
or tapered either from base upward, or from middle upward and down-
ward.

Ear " density " and number of spikelets.

It is usual to define the " density " (D) of an ear as the number of
spikelets per 10 cm. length of the rachis. The total length in centimetres
(L) of 10 normally developed ears is measured and the number of spikelets
(N) counted ; from these determinations the density is calculated thus :

In the descriptions of wheats given later the following table of densities
is adopted :

Lax . . D below 22.

Medium . . . 22-28.

Dense .... 28-34.
Very dense . . . above 34.

Very dense ears are characteristic of the Club Wheats ; a few are seen
in the Bread Wheats, but they are rare among the Macaroni and Rivet
Wheats and unknown in Emmer and Large or Small Spelt.

The density of the ear is subject to the ordinary fluctuating variation
and is influenced to some extent by climatic changes and alterations in the
water-supply and nutrition of the plant.

The application of large amounts of nitrogenous fertilisers tends to
lengthen the internodes of the rachis, but I have failed to produce by the
most drastic manurial treatment a lax ear in plants of T. compactum or
in any Squarehead vulgare wheat.

CLASSIFICATION 161

I find that the ear-density of some forms of Bread Wheat from India
and Australia is more readily influenced by external conditions than that
of any other wheats.

Although the " density " in some countries appears to be so variable
that it is of little or no value there for purposes of classification, in other
regions it is practically constant under all ordinary conditions of cultiva-
tion, and of the greatest service in assisting in the identification of the
various forms.

Empty Glume. — The form of the empty glume is one of the most
constant characters of wheats, and of the greatest value in distinguishing
nearly related forms. In some cases the glumes at the base, middle, and
tip of the ear are slightly different in shape and length of the apical or keel
tooth, but the glumes from the middle spikelets of ears belonging to plants
of the same form are almost exactly alike, and different from those taken
from the same parts of the ears of different forms. In some the upper
part of the glume is broad, in others it is narrow ; the apical tooth has a
characteristic form and size, and the amount of its curvature inward or
outward is also a constant feature. These forms unfortunately cannot
be adequately described, but must be illustrated.

Grain. — Its colour, flintiness or mealiness, shape and size.

Other characters which it is important to notice are —

(1) Time at which the ear appears ;

(2) The tillering capacity ; and

(3) Resistance to frost, drought, and attacks of insects and fungi.
Earliness and Lateness. — Wheats are often classified by the practical

agriculturist into early, mid-season, and late varieties, the terms referring
to the time when the grain is fully developed and ripe. The period at
which this stage is reached is chiefly dependent upon latitude and climatic
conditions and subject to very considerable variation. For example, the
same variety of wheat sown at the same date in the same locality may
ripen its grain from 1 5 to 20 days or more earlier or later in one season than
another, the grain ripening off quickly in a bright dry hot summer, the
process being very much delayed by damp dull weather.

A much more accurate classification is made by referring the terms
early, mid-season, and late to the time at which the ear escapes from the
upper leaf-sheath. This time for any particular variety grown at any
definite locality in England does not vary more than four or five days
in any season even when the sowing is carried out during the period
extending from September to December. When the sowing is later than
this, say in January or early February, the time of appearance of the ear
is rarely delayed more than about a week.

For details of the time usually elapsing between " earing " and
ripening of the grain, see p. 142.

M

1 62 THE WHEAT PLANT

In the descriptions of wheats in the succeeding chapters I have
adopted the following classification based on observations of more than
1500 forms grown for several years at Reading :

Date of Earing.

Very early . . . before May 24. \
Early .... May 24-31.
Mid-season . . . June 1-7.
Late .... June 8-15.
Very late . . . after June 1 5 .

This system of classification holds good for all districts and an early
or late variety will bear the same relationship to each other in regard to
the time of earing, wherever grown, though the actual dates to which
these terms correspond will depend upon the latitude and climate of the
place of growth.

Early forms always have the erect habit, while late forms are prostrate
(see p. 69).

Spring and Winter Wheats. — At Reading, where the average minimum
winter temperature rarely falls below - 3° or - 4° C., I have always sown
all kinds of wheats in autumn and have rarely observed any damage by
frost, even among the most delicate kinds.

There is, however, considerable difference among wheats in regard
to their resistance to frost, some being killed outright by temperatures
which others will withstand without damage.

Many wheats are little injured at - 10° to - 15° C., but suffer when
the temperature falls much below this.

In cold climates the differences are readily determined, and farmers
term the sorts which can be sown in autumn " Winter " wheats, applying
the term " Spring " to those which are delicate and must be sown after
the winter has passed.

Varieties of T. dicoccum, T. orientate, T. durum, T. polonicum, T.
turgidum, and T. pyramidale are usually delicate ; on the other hand,
T. sphaerococcum, T. Spelta, and T. monococcum are hardy races. Some
forms of T. compactum and T. vulgar e are also hardy, while others are
tender and die out in severe continental winters. Of these, the rapid-
growing forms with the erect habit (p. 69) and broad leaves are usually
delicate, while the slower-growing, late-ripening sorts with narrow leaves
which lie close to the ground are hardy.

Sinz found that Winter wheats showing great resistance to frosts
transpire less, have firmer tissues and higher dry-matter content than
Spring forms.

I have frequently observed that hares and rabbits pick out and eat
typical Winter wheats before touching the Spring forms when both are
grown in the same field.

CLASSIFICATION 163

Nomenclature of Forms. — The nomenclature of wheats in all countries
is in hopeless confusion ; the same form is frequently found under
many different names, and totally different forms are often given the
same name. The trouble is not modern, for mediaeval names for the
same form of wheat are abundant : it has, however, increased in recent
times, and there is no prospect that it will ever cease.

Some of the evil effects of the confusion might be mitigated by detailed
and illustrated descriptions of the wheats of each country, and the use of
the name of an old-established form for another form belonging to a
different variety should be prohibited. For example, the name " Velvet
Chaff " sometimes given to " Preston," a smooth-glumed form, should
not be allowed, nor should the name " Browick," originally given to a
red-chaffed form, be permitted for a white-glumed wheat as at present
in England.

The application of a new name to an old form cannot be prevented,
but official lists of synonyms and names of forms found after extended
trials to be closely similar in morphological characters would be of much
service to the present generation and of value in the future.

CHAPTER X

WILD SMALL SPELT

Triticum aegilopoides, Bal. ex Korn. (aegilopodioides, sphalm.). Handb.
d. Getreideb. i. 109 (1885).

Crithodium aegilopoides. Link in Linnaea, ix. 132 (1834).
Triticum boeoticum, Boiss. Diagn. ser. i. fasc. 13, p. 69 (1853).
Aegilops Crithodium, Steud. Syn. Gram. 355 (1855).
Triticum monococcum, fi. lasiorrachis , Boiss. Fl. Or. v. 673 (1884).
T. monococcum, A. aegilopoides, Asch. u. Graeb. Syn. ii. 701 (1901).

THIS wild species of Triticum was first described under the name
Crithodium aegilopoides by Link, who found it in Greece between Nauplia
and Corinth in 1833.

It is a widely distributed grass on the sides of low hills in Thessaly,
Boeotia, and Achaia in Greece, in South Bulgaria, and on loamy soils in
vineyards in Southern Serbia. Three varieties of it were discovered by
Larionow in 1909 near Balaklava in the Crimea, and the variety of T.
monococcum mentioned by Marschall Bieberstein (Fl. Tauro-caucas.
vol. i. p. 85) as occurring wild in the Crimea and Eastern Caucasus was
probably the same species. The species is also met with throughout
Asia Minor extending in Kurdistan to the western border of Persia.

There is no doubt that it is from some of the varieties of this wild
species that the cultivated Small Spelt (Triticum monococcum, L.) has
been derived.

GENERAL CHARACTERS OF T. aegilopoides, Bal.

The coleoptile is usually purple, the shoots of the young plants in some
forms erect, in others prostrate. The culms are erect, solid or hollow
with thick walls, elastic and slender, from 40 to 160 cm. (15-60 inches)
high, smooth except at the nodes, which are covered with white deflexed
hairs ; the upper internode is exceptionally long, in some forms reaching
a length of 65 cm. in culms whose total length is not more than 100 cm.

The blades of the young plants are narrow (3-6 mm. across), those of

164

WILD SMALL SPELT 165

the culm leaves broader (5-15 mm. across), their surfaces in most plants
being clothed with soft short hairs, among which are longer hairs sparsely
distributed along the summit of the longitudinal ridges («, Fig. in) and
on the margins of the leaves. The leaf-
sheaths,' which are striate, are pubescent in
some cases and glabrous in others. The
auricles of the lower leaves are fringed with

long hairs.

,, • , r . FIG. in. — Diagrammatic trans-

Shooting, OF the escape Ot the ears verse sections of young leaves

from the upper leaf - sheaths, occurs at of (a) T. aegilopoides, (b) T.

. monococcum.

Reading about June 20, in this respect

resembling the cultivated " Engrain double " (T. monococcum).

The ears are flat, consisting of two rows of spikelets closely arranged
on opposite sides of a flattened and jointed rachis. They are from 5 to n
cm. long, '45--9 cm. broad, and -2--25 cm. thick, and possess from 12 to
30 or more spikelets. The density (D) or number of spikelets per 10 cm.
of rachis varies from 25 to 50, the Asiatic varieties having the laxest ears.

The rachis is smooth, but its edges are fringed with a line of longer .or
shorter hairs, and a tuft of long whitish hairs (from 1-5 to 3 mm. long)
is attached to it in front of the base of each spikelet. When ripe it is
very brittle, and separates at the joints into short pieces about 3 mm. long
and i mm. broad to the upper end of each of which is attached a single
spikelet. Disarticulation of the ears proceeds from the tip towards the
base, the apical spikelets often falling while the basal ones are still green.

The terminal spikelet of the ear is abortive. The lateral spikelets
are two-flowered, the lower flower fertile, the upper one usually sterile ;
they measure about 9 mm. long, 2-2-5 mm- broad, and i mm. thick.

When two caryopses ripen in the spikelet the upper one is the larger
and germinates more rapidly than the lower one, even when both are
stripped from their enclosing glumes.

The glumes are yellowish, reddish, or black, usually pubescent, but
in some forms they are glabrous.

The empty glume (7-8 mm. long) is ovate, the outer half somewhat
broad and truncate, with a scabrid keel terminating in a triangular tooth
about 1-5-2 mm. long, which is usually bent outwards especially in unripe
ears ; it possesses a strong lateral nerve which ends in a small tooth -5
mm. long (Fig. 112).

The flowering glume of the lower flower bears a scabrid awn varying
from 6 to 1 1 cm. long, that of the upper flower in robust forms having an
awn of similar length, while in smaller forms the glume is awnless or with
an awn rarely exceeding 1-2 mm. in length. When young the palea is
entire, but in ripe ears it is divided longitudinally into two halves.

The caryopsis is pointed at both ends, very much laterally compressed,

1 66 THE WHEAT PLANT

and olive-yellow, with a flinty endosperm. It measures from 6 to 7-5 mm.
long, 1-1-5 mm- fr°m side to side, and 2-1-2-6 mm. from back to front.
The furrow is seen as a shallow groove along the narrow edge of the grain ;
the radicle of the embryo is prominent, the opposite end having a " brush "

231 123

FIG. 112. — Empty glumes ( x 2) and grains (front, back, and

side views) of T. aegilopoid.es (nat. size),
i, var. Larionowi. 2, var. boeoticum. 3, var. Thaoudar.

of hairs. The weight of 100 caryopses varies from
i-o to 1-3 grams.

T. aegilopoides may be subdivided into two sections
as indicated below :

SECTION I. — Young shoots and culms erect and caes-
pitose (" spring type ") ; young leaves broad (10-15 mm-
across), yellowish-green.

1 . Flowering glume of the lowest flower of the spikelet
with a long awn (9-11 cm.), the second flowering glume
usually with a short point only (2-5 mm.), var. boeoticum.

2. Flowering glumes of both flowers generally with
equally long awns (9-10 cm.) . . . var. Thaoudar.

T. aegilopoides, var. boeoticum, mini.

T. boeoticum, Boiss. Diagn. ser. i. fasc. 13, p. 69 (1853). FlG- IJ3-

This is the most widely distributed European variety Grains from the spike-
of T. aegilopoides (i, Fig. 114), being frequent in parts of lets of on^side.,of
southern Serbia, Bulgaria, and Greece. It is found poides, var. boeoti-
throughout Asia Minor and has been recorded by Larionow cum (nat. size).
from the Crimea, although from Vavilov's account (Bull.
App. Bot. vol. vi. p. 691) it is probable that the Crimean form should be placed
in Section II. with varieties Pancici and Larionowi, which are winter forms with
narrow leaves and prostrate young shoots.

The young shoots of the forms obtained from M. Kornicke and grown at
Reading are erect, the leaves pale yellowish-green, pubescent, with long white
hairs sparsely distributed along the summit of the longitudinal ridges.

WILD SMALL SPELT 167

The ears of wild plants are 5-6 cm. long possessing 24-26 spikelets ; those
of cultivated specimens are often 8 or 9 cm. long with 30-32 spikelets each
about 10 mm. long.

The yellowish-white glumes are usually clothed with soft inconspicuous
hairs, but glabrous forms are met with : Flaksberger's var. Zuccariorii appears
to be the pubescent form.

T. aegilopoides, var. Thaoudar, mihi (2, Fig. 114).

T. Thaoudar, Reuter (in Bourgeau, PI. Exs., 1860).

In 1854 Balansa collected wild strong-growing forms of T. aegilopoides
at Balamaut Kaive between Smyrna and Magnesia in Asia Minor. Similar
robust wild plants were discovered later in Lycia by Bourgeau (1860) ; near
Amasia in 1889 by Bornmiiller ; in Northern Syria by Aaronsohn ; and in 1909 in
Kurdistan on the borders of Persia by Strauss.

The Turkish name for these wild forms of Triticum as well as for rye (Secale
cereale) and Darnel (Lolium temulentum) is " Thaoudar," and Bourgeau's speci-
mens were labelled Triticum Thaoudar by Reuter. I have critically examined
and compared authentic specimens of all the recorded varieties of T. aegilopoides,
and find that the Asiatic types usually have slightly broader and thicker ears
with fewer spikelets than those of Serbian and Greek plants ; the glumes of the
former are sometimes tinged a pinkish colour, and in addition, many, though
not all of them, have a long awn (9-10 cm.) on the upper as well as on the lower
flowering glume.

This is the common variety of Asia Minor, although plants indistinguishable
from the European variety boeoticum and intermediates appear with it.

The glumes, which are yellowish-white, are generally pubescent, and the outer
part between the strong lateral nerve and the margin of the empty glume is
sometimes broad and rounded as in cultivated T. monococcum.

The ears are shorter and laxer and the spikelets fewer and stouter than those
of var. boeoticum, and in wild specimens usually 4-5 cm. long and -8--9 cm. wide,
having 16-18 spikelets each 10-15 mm. long. When cultivated the ears attain
a length of 8 or 9 cm. with 30 or more spikelets.

SECTION II. — Young shoots prostrate ; culms decumbent at the base
(" winter type ") ; young leaves narrow (4-5 mm. across) and dark green, the
habit of the young plants in winter resembling that of T. hermonis.

1. Glumes and awns black .... var. Paricici, mihi.

2. Glumes reddish-yellow, awns black . . var. Larionozvi, mihi.
T. aegilopoides, var. Pandci, mihi.

T. monococcum, var. Pantid, Flaksb. Bull. App. Botany, Petrograd, vi. 682

A wild black-glumed variety (4, Fig. 114) discovered by D. Larionow in
1909 near Balaklava in the Crimea, growing with vars. boeoticum and Larionozvi
(see below), possibly the same as T. nigrescens, PanCici (in PL Exsicc. et lift.,
Korn. Handb. d. Getr. i. 109, 1885).

In the Fielding Herbarium at Oxford are four specimens of this variety

1 68 THE WHEAT PLANT

collected near Smyrna by Sherard, who was British Consul there from 1703
to 1718 : they are labelled " Hordeum distichum spica nitida arratis et glumis
nigricantibus. Comp. Tab. Bot. 38."

The following description is from plants grown at Reading, the original ears
being sent me from Moscow by Dr. Vavilov :

The coleoptile is purple.

The shoots of the young plants are prostrate, the leaves narrow (4-6 mm.
broad) and a dark blue-green tint. The leaves are clothed with short velvety
hairs, interspersed among which are some 1-1-5 mm- l°ng> arranged on the
ridges and along the margins as in a, Fig. in. The blade of the upper leaf
of the straw is shorter than the sheath.

The culms at first spread outwards close to the ground, becoming erect later.
They possess 4 or 5 internodes and vary much in length on the same plant, some
reaching a height of 130-150 cm. (about 50-60 inches), while others are not
more than 30-45 cm. (12-18 inches) long.

The upper internode is solid or hollow with thick walls and of exceptional
length, being often as much as 80 cm. long.

A conspicuous muff of white reflexed hairs surrounds the nodes, below which
-| to \ inch of the straw is brownish.

The ears without the awns are 9-11 cm. long and -y-'S cm. across the two-
rowed side : they possess 28-32 spikelets and are moderately dense (D = 30-32).

The rachis is smooth, but its edges are fringed with whitish hairs
3 mm. long, and there is a frontal tuft about 3 mm. long at the base of each
spikelet.

The spikelets are somewhat larger than those of var. Larionowi, measuring
12-14 mm. long and 4 mm. broad. They are two-flowered and Dually ripen
two grains in each spikelet ; the lower is the smaller grain and germinates
more slowly than the upper one or not at all.

The empty glumes are jet black or more or less covered with dark streaks,
and the awns are black. The keel and lateral nerves are scabrid, the keel tooth
about 2 mm. long and straight or bent outwards slightly.

The caryopses are small, narrow, and pointed at both ends, greenish-grey
or purplish in colour with flinty endosperm. The smaller ones measure about
6 mm. long, 1-3 mm. from side to side, and 1-45 mm. from front to back, the
larger ones being 7 mm. long, 1-45 mm. broad, and 1-8 mm. thick ; 100 weigh
from i to 1-2 grams.

T. aegilopoides, var. Larionowi, mihi.

T. monococcum, var. Larionowi, Flaksb. Bull. App. Bot., Petrograd, vi. p. 681

A wild form with glabrous yellowish-red glumes and black awns found in
1909 by Larionow near Balaklava in the Crimea (3, Fig. 114).

The following description is from plants grown at Reading :

The coleoptiles are purplish. The young shoots are prostrate, the foliage
leaves dark blue-green, 4-5 mm. broad, with short blades and clothed with soft
hairs as in var. Pancici.

The straws are slender, solid or hollow with thick walls, green or purplish

FIG. 114. — WILD SMALL SPELT (T. aegilopoides, Bal.).

1. var. boeoticum.

2. var. Thaoudar.

3. var. Larionowi.

4. var. Pancici.

WILD SMALL SPELT 169

with a muff of white deflexed hairs : they grow to a height of 120-150 cm.,
the upper internode often 60-70 cm. long.

The ears are narrow, without the awns 8-10 cm. long and -5 -'7 cm- across
the two-rowed side. They possess 30-36 spikelets and are of moderate density
(0 = 30-32).

The rachis is smooth, its edges fringed with white hairs 2-3 mm. long ; the
frontal tufts of hairs at the base of each spikelet 2-3 mm. long. The spikelet
is 11-13 mm- l°ng and 3 mm. broad, two-flowered, generally producing a single
ripe grain, though spikelets with two caryopses are sometimes found.

The empty glumes are glabrous, reddish-yellow in colour, the keel and strong
lateral nerve scabrid ; the apical tooth is about 2 mm. long and bent outwards
slightly.

Both flowering glumes of the spikelet possess slender black awns, one of them
8-9 cm. long, the other 1-2 cm. long ; occasionally both awns are 8-9 cm. long.

The caryopsis is laterally compressed, pointed at both ends, olive green
especially the apical half, the endosperm flinty. It measures about 7 mm.
long, 1-6 mm. through from side to side, and 2-2-5 mm- thick from dorsal to
ventral surfaces ; 100 weigh from 1-2 to 1-3 grams.

CHAPTER XI

SMALL SPELT

T. monococcum, L. Sp. PI. 86 (1753).

Nivieria monococcum, Ser. Cer. Eur. in, 114 (1841).

T. vulgare bidens, Alef. Landw. FL 334 (1866).

T. monococcum, B. cereale, Asch. u. Graeb. Syn. ii. 702 (1901).

SMALL Spelt, sometimes termed one-grained Wheat, St. Peter's corn,
or Einkorn, is one of the most primitive of the cereals, exhibiting little
differentiation from the wild T. aegilopoides from which it has no doubt
been derived.

Its cultivation was established in prehistoric times, and evidence
points to its having been one of the chief wheats grown in Mid Europe
in the Neolithic period. Grains of it have been discovered in Hungary
in the Stone Age deposits of Fels5-Dobzsa, the Neolithic subterranean
dwellings and store-rooms of Lengyel, and in the cave deposits of Aggetelek ;
they have also been found in deposits of similar date in Bosnia and
Denmark.

Heer states that a single ear — now lost — was obtained from the pile-
dwellings, referred to the same era, at Wangen in Switzerland. Car-
bonised grains dug up by Schliemann at Hissarlik on the site of ancient
Troy (3000-2500 B.C.) have been attributed to Small Spelt, and specimens
belonging to the Bronze Age have been found at Toszeg in Hungary.

The grains of Neolithic Age are somewhat smaller than those of T.
monococcum of the present day, resembling the caryopses of the wild
T. aegilopoides of Serbia and Macedonia. It is possible that prehistoric
man in Mid and South-Eastern Europe first selected for cultivation the
wild species of this region and at the early period mentioned had made
little progress with its improvement. The increased size of the grain
of some of the varieties of this race at present cultivated may be the result
of a long period of selection applied to the European plant ; on the other
hand, it is perhaps more likely that the modern forms of T. monococcum
have been derived from the larger robust variety — T. aegilopoides, var.
Thaoudar of Asia Minor.

170

SMALL SPELT 171

There is no proof that T. monococcum was known to the ancient
Egyptians, nor is there any reference to it among the Roman writers on
agriculture, but ri(f>r] mentioned by Aristotle, Theophrastus, and other
early Greek authors appears to have been this cereal.

On account of its poor quality, lateness in ripening, and insignificant
yield in comparison with other wheats, its cultivation is now very re-
stricted. Nevertheless the awns and thick glumes protect it from the
attacks of birds, and its power of resisting frost and rust, and the fact that
it will grow without manure on poor sandy, chalky, and rocky soils where
better wheats fail, adapt it to the needs of the inhabitants of barren
mountainous districts.

From very early times it has been cultivated by primitive peoples in
remote districts of Europe and Western Asia, and it is still grown in many
parts of Spain and to a lesser extent in Switzerland, France, Italy,
Germany (Wilrtemberg and Thuringia), Herzegovina, Greece, Macedonia,
and in the Eastern Caucasus. It has also been met with in cultivation in
Asia Minor, but there are no records of its occurrence in India, China,
or on the continent of Africa, and it does not appear to be cultivated in
Serbia or Bulgaria, of which countries its wild prototype is a native.

Small Spelt is mainly utilised in its husked state instead of barley
as fodder corn for cattle and horses. In lesser amount the true grain
freed from the tough glumes is used in soup and gruel in place of " pearl "
barley or groats.

It has also been employed in the manufacture of beer and vinegar.

The flour obtained from it is yellow or brownish in colour, and makes
a good flavoured' though dark brown bread.

Both winter and spring forms are known, the former being perhaps
the most frequently cultivated. In Europe the winter forms are sown in
September, spring forms early in March, about 72 Ibs. of husked grain
per acre being needed. The crop is harvested in August or the first half
of September, the yield is very variable, from 8 to 16 hectolitres per hectare
(about 320-640 Ibs. per acre) being obtained in poor mountainous regions ;
on good soils more than 80 hectolitres per hectare have been recorded.

The ears of T. monococcum are very brittle, and it is stated that the
plant sometimes sows itself among other crops from which it is difficult
to extirpate it when once established ; it was reported in 1871 as a
troublesome weed in cornfields round Montpellier in France.1

GENERAL CHARACTERS OF T. monococcum, L.

Small Spelt is strikingly distinct from other cultivated wheats in its
stiff erect habit, slender straw, and characteristic pale yellowish-green tint.

1 Bull. Soc. Bot. France, p. 173 (1871).

172 THE WHEAT PLANT

The leaves of young plants in winter forms lie spread out on the surface
of the soil, in spring forms they are more erect ; the blades are narrow,
from 4 to 10 mm. across, bluish-green at first, yellowish-green later, and
covered with very short hairs, 50-80 ^ long, on both surfaces.

Tillering is extensive, particularly so in the slow-growing winter forms.

The straw, which, according to variety, reaches a height of 60-120 cm.
(2-4 feet), is smooth and thin, but tough and elastic, usually hollow in all
the internodes except the basal one, which in some kinds is solid : the
walls of the straw are thin.

On each there are four or five nodes, the upper ones being clothed with
somewhat conspicuous deflexed hairs, those next the ground often destitute
of hairs or nearly so. Just below each node the straw in the common
forms is a brownish-purple colour.

In straws with four internodes the average successive lengths of the
latter from below upwards were found to measure 6, n-6, 22, and 42 cm.
respectively, straws with five internodes measuring 4, 12-6, 16, 25-5,
and 42-7 cm.

The combined sheath and blade of the fully developed leaves on the
straws are from 25-5 to 35-5 cm. (10-14 inches) long, -6--J cm. broad,
the hairs on them being fewer and longer than on the young leaves of
autumn and spring ; the overlapping edge of the sheath is generally
fringed with soft hairs.

The ligule is short and truncate, with irregular teeth ; a pair of small
auricles with accompanying long hairs are present at the base of the
blade.

The ears are bearded, erect, laterally flattened and thin, superficially
resembling those of two-rowed barley. They vary in length from 4-5
to 9 cm., having a width of 7 to 10 mm. across the two-ranked side and
3-5 mm. across the narrow face of the ear.

The spikelets are packed closely in an imbricate manner, small ears
having 22-25 spikelets, the largest 45 or more ; the density (D) = 50-55.

The rachis is flattened (1-2-1-5 mm. wide and -3~-4 mm. thick) and
very brittle, breaking into pieces 1-5 mm. long, to the upper end of which
is attached a single spikelet. In all forms the rachis is smooth, but
in some the edges are fringed with straight white hairs, especially at
the point of insertion of the empty glumes, and a tuft of similar hairs is
present on the rachis in front of the base of each spikelet between the
glumes.

The spikelets generally possess three flowers, the upper one of which
is always imperfect and sterile ; in some varieties only the lower flower
is fertile, while in other kinds the first and second flowers of the spikelets
produce ripe grains. The terminal spikelet of the ear is very minute and
barren.

SMALL SPELT 173

Each fertile spikelet is about 10 mm. long, 4-6 mm. wide, and 3 mm.
thick, the front being convex, the back flatter.

The glumes are glabrous or slightly pubescent.

The two empty glumes are shorter than the flowering glumes, boat-
shaped and unsymmetrical, the halves of each right and left of the midrib
differing considerably in form and width. They are strongly keeled from
apex to base and usually 5 -nerved, one small nerve being found on the
inner narrow side of the glume close to the midrib and three others on the
broad outer half, the outermost of these nerves being strong and forming
a conspicuous rib.

The exposed part of the glume turned away from the rachis between
the keel and the third strong nerve is thick and leathery, but its
edges and the whole of the inner half nearest the rachis are thin and mem-
branous.

The apex of the empty glume is notched in a characteristic manner,
the keel terminating in an acute point or tooth 1-2 mm. long, the strong
secondary nerve of the broad exposed half ending in a shorter tooth -5-
i mm. long (Fig. 115), while a third blunter projection is seen on the narrow
inner half.

The flowering glume is boat-shaped, rounded on the back, and without
a keel. It is thin, white, and semi-transparent, with 9 to u nerves, the
middle and two outermost being the strongest. The apex is divided
into two short points between which arises a long straight scabrid awn of
triangular section.

Well-developed awns which are only associated with the lowest flower
of a spikelet vary in length from 3 to 8 cm., those in the upper part of the
ear being longest. The second flowering glume, whether belonging to a
barren or a fertile flower, has a short awn usually not more than 2-5 mm.
long, or is awnless.

The palea is a thin membranous glume equalling the flowering glume
in length, with two nerves and terminating in a blunt apex. At first it
is entire, but shows a slight inward fold down the centre, along which it is
torn into two separate halves when the fruit ripens ; the latter character,
together with the compressed fruit, led Seringe to place Small Spelt in a
separate genus, Nivieria.

The flowers are of the ordinary gramineous type with yellow anthers
4-4-5 mm. long and lodicules which are pubescent in the upper part.

The abortive second flower of the spikelet possesses a pistil whose
stigmas are rudimentary, and three stamens, the anthers of which do not
produce pollen-grains.

Anthesis begins in the upper third of the ear.

The caryopsis, which is closely invested by the glumes, is yellowish
and flinty, oval and tapered at both ends, the outline of both the dorsal and

M

H

t

THE WHEAT PLANT

ventral surfaces being curved when the grain is viewed from the side.

The radicle of the embryo projects prominently, and the opposite end of

the grain is clothed with a " brush " of short hairs.

It is compressed laterally, being usually of considerably greater

diameter from front to back than from side to side.

The furrow which
extends to the centre of
the grain is almost com-
pletely closed and only
visible as a narrow
groove on the ventral
part of the caryopsis

((Fig- "5)-
The grains of differ-
ent varieties measure
from 7 to 8-5 mm. in j&

, length, 1-8-3 mm- m &

FIG. 115. — Empty glumes (x 2), and r «j

grains (front, back, and side views), breadth from Side to ft

of T. monococcum (nat. size). g^e, an(J 3.3.5 mrrii ^

;:r*£&.. f«»« d^1 to ventral *

surfaces.

The ratios length : breadth : thickness = about 100 :
26 : 43 in some varieties, in others 100 : 35-3 : 41.

The produce of the thrashed ears, i.e. the " husked "
grain or " vesen," weighs from 40 to 50 kg. per hectolitre.
T. monococcum is a comparatively homogeneous
species, the morphological diversity of the different ,

/ . Grams from the

forms being small. spikelets of an

The hairs on the glumes of pubescent varieties are ear of ,T- mon°-

coccum (nat. size).

inconspicuous and never abundant ; when present they

are most readily seen on the upper part of the flowering glume. In

some cases the hairs are reduced or disappear altogether under good

cultivation.

Both winter and spring forms are known, the former having narrower
leaves, shorter culms, and smaller ears than the latter ; they also ripen
later than the spring varieties.

In most wheats the length of the axis and number of spikelets in the
ear is subject to little variation, but in this race these characters are very
extensively influenced by the fertility of the soil, the date of sowing, and
by climatic conditions.

On poor soils the ears are often not more than 3 or 4 cm. long, but
when grown on good soils the ears may reach a length of 9 cm. (3^ inches)
or more and possess 40 or 50 spikelets.

SMALL SPELT 175

In general facies T. monococcum closely resembles the wild T.
aegilopoides ; it differs from the prototype in having somewhat broader,
closer-set, and larger ears, and grains of slightly larger size ; its awns are,
however, shorter, being usually not more than 4-8 cm. long, while those
of the wild species reach a length of 10-11 cm.

The leaves are covered with very short hairs (b, Fig. in), there being
no long ones on the ridges as in T. aegilopoides. The leaf-sheaths, auricles,
and rachis of the ear in the cultivated plant are much less hairy, and the
empty glumes broader with rounded and more membranous margins than
those of T. aegilopoides. The terminal keel tooth of the empty glume of
the latter is generally bent outwards ; in most varieties of T. monococcum
it is either straight or curved slightly inwards.

VARIETIES OF T. monococcum, L.

1 . Glumes pale reddish-yellow or brownish, glabrous, shining.

var. vulgare, Korn.

2. Glumes pale yellow, scabrid, dull . . . var. flavescens, Korn.

3. Glumes reddish-brown or brownish-yellow, pubescent, dull.

var. Hornemanni, Korn.

Glumes pale reddish-yellow or brownish, glabrous, shining.

T. monococcum, var. vulgare, Korn. Handb. d. Getr. i. 112 (1885).

This variety (2, Fig. 117) is a winter form of slow growth, but often ripens
grain at Reading in the same season, even when sown as late as the beginning
of March.

When sown in autumn the ears appear in the last week of the following June
or the first week in July.

The plants tiller extensively, producing 10-15 straws per plant when the
grains are sown 6 inches apart in rows 6 inches asunder.

The coleoptile is purple, the young foliage leaves narrow, about 4 mm.
across, bluish-green, and lying close to the surface of the ground in winter ;
the fully developed culm leaves are 6 to 8 mm. across, of a yellowish-green tint.
The leaf-sheaths are generally smooth, the lower ones and the fringed auricles
usually pinkish.

The straw is very short, from 65 to 75 cm. (about 25-30 inches) in height,
slender, elastic, and hollow, with thin walls, the nodes covered with fine more
or less adpressed and deflexed hairs.

The ears are thin, smooth, and reddish-yellow, with more or less polished
surfaces, the keel and veins of the glumes being non-scab rid or nearly so ; they
average 5-6 cm. in length, about 8 mm. in width, and 3-4 mm. across the face
of the spikelets, each of which usually contains only one small compressed
caryopsis. D = about 54.

Only the flowering glume of the lower flower has an awn, 3-8 cm. long, that
of the second flower being awnless.

1 76 THE WHEAT PLANT

The produce of the thrashed ears, i.e. the husked grain or " Vesen,"
consists of about 80 per cent of caryopses and 20 per cent of chaff (pieces of
rachis and glumes). From 450 to 480 caryopses weigh 10 grams.

The caryopses are yellowish and flinty, flattened from side to side ; they
measure about 7 mm. long, 1-8-2 mm. through from side to side, and 3 mm.
from front to back.

The average ratio of length, breadth, and thickness = 100:27-1 : 42-8
respectively.

Glumes pale yellow, scabrid, dull.

T. monococcum, var. flavescens, Korn. Handb. d. Getr. i. 112
(1885).

T. monococcum, a. alba spica glaberrima, Lam. Enc. ii. 560 (1786).

The commonest representative of this variety (3, Fig. 117) is known as
Engrain double in France, into which country Heuze says it was introduced
from Spain about 1850. It is a spring form with caespitose young shoots. The
plants ripen from 15 to 20 days earlier than the preceding variety when both
are sown at the same time either in autumn or spring. The coleoptile is
colourless and the young foliage leaves much broader than those of other
varieties, being from 10 to 12 mm. wide. The auricles are fringed with a few
long hairs and, like the leaf-sheaths, usually pale green ; the leaf-sheaths have
long hairs sparsely distributed over their surfaces. The straw is strong and
hollow, with thick pithy walls and of medium height, reaching a length of
100-115 cm. (40-45 inches) or more. On the nodes are spreading deflexed
hairs.

The ears are pale straw colour, usually 5-6 cm. long, less densely packed
on the rachis and thicker than those of var. vulgar e. D = about 50.

The spikelets are broad, 4-5-5 mm. across, and in some ears many of them
contain two well-developed caryopses. The " Vesen " have a less proportion
of " husk " than those of the Common Small Spelt, averaging 16 per cent husk
to 84 per cent of true grain.

The coarse empty glumes have characteristic rough dull surfaces, on which
are minute papillae ; the keel and outer nerves are strong and scabrid ; their
apical teeth not so acute as in other varieties.

The flowering glume of the second flower possesses a short awn 2-5 mm.
long, that of the first or lower flower having an awn 3-6 cm. long.

The caryopses are longer and broader than those of var. vulgar e. The
ventral surface of the grain is broader and flatter and the furrow more easily
seen than in those of the previously mentioned varieties. They measure 8-8-5
mm. in length, 2-5-3 mm. in width, and 3-3-5 mm. from front to back ; from
340 to 350 weigh 10 grams.

The average ratio of length : breadth : thickness = 100 : 33-3 : 39'3-

Kornicke's variety laetissimum appears to be merely a small pale-eared form
of \ar.flavescens.

FIG. 117.— SMALL SPELT (T. monococcum, L.).

i. var. Hornemanni. 2. var. vulgare.

3. var.flavescens (Engrain Double).

SMALL SPELT 177

Glumes pale reddish-yellow or brownish, pubescent, dull.

T. monococcum, var. Hornemanni,Korn. Handb. d. Getr. i. 112 (1885).

T. Horneman, Clemente (in Herrera, Agric.gen. i. 3, 1818).

T. monococcum, b. spica pallide rubra pubescente, Lam. Encyclop. ii. 560
(1786).

This is the largest and most commonly cultivated variety of T. monococcum
(i, Fig. 117), ripening at Reading about the middle of June, resembling in this
respect v&r.flavescens.

It is grown in small quantity in Switzerland and Spain and the Eastern
Caucasus (Tersk Government), and was formerly cultivated by Tatars in the
Crimea.

The commonest representative of this variety is a spring form with
caespitose young shoots, and stout straw of medium height (110-120 cm.).
The leaf-blades are about 10 mm. across and yellowish-green.

The ears are pale reddish brown from 6 to 10 cm. long, 10 to n mm. across
the 2-ranked side and 4 to 5 mm. across the narrow face. The number of
spikelets 25-40. 0=40-50.

The spikelets are 11-12 mm. long, 5 mm. broad, usually ripening one grain
in each, although spikelets with two fully developed caryopses are frequent.

The glumes near the tips are clothed with short inconspicuous hairs and the
flowering glume of the lower flower bears an awn about 9 cm. long, that of the
second upper flower having only a short awn 3-6 mm. long.

The caryopses are 8-9 mm. long.

N

CHAPTER XII

WILD EMMER

T. dicoccoides, Korn. in lift, ex Schweinf. in Ber. d. Deutsch. Bot. Ges. 309
(1908).

T. vulgare, Vill., var. dicoccoides, Korn. Verh. des naturhist. Vereins d. Pr.
RheinL, Jahrg. 46, Bonn (1889).

T. hermonis, Cook. Bureau of PI. Ind. (U.S.A.] Bull. No. 274, 13, 52 (1913).

IN 1873 Kornicke discovered in the National Herbarium at Vienna a
portion of an ear of a species of Triticum among specimens of Hordeum
spontaneum collected in 1855 by Theodor Kotschy at Rasheyya on the
north-western side of Mount Hermon in Syria. In 1899 he referred to
the plant at a meeting of the Niederrheinischen Gesellschaft fur Natur-
und Heilkunde, and according to the report of the meeting he named it
T. vulgare, Vill., var. dicoccoides, considering it closely allied to Emmer and
the prototype of most of the cultivated wheats.

The plant was rediscovered first at Rosh Pinar at the foot of Jebel
Safed in Syria by Aaronsohn in 1906 and later at Rasheyya and other parts
of Mount Hermon as well as on the plateau of Es-Salt east of the Jordan
valley. Specimens were also collected in 1910 by Theodor Strauss in
the mountainous region of Western Persia near Kerind, between Kerman-
shah and Bagdad.

T. dicoccoides grows in the crevices of limestone rocks in dry situations
from 300 to 500 feet below to over 6000 feet above the level of the Medi-
terranean, and is generally associated with Hordeum spontaneum and often
with forms of Triticum aegilopoides .

The reports of Aaronsohn, Cook, and others who have observed T.
dicoccoides in its native habitat leave no doubt that it is truly wild, and
different from any kind of wheat at present cultivated in Palestine.

For some time I have grown it from ears obtained from El Hadr and the
eastern slopes of Mount Hermon. I have also examined specimens col-
lected by Strauss in Western Persia and others from various parts of Syria.

The plants in their native habitat exhibit great diversity in the form of
the glumes, the size and prominence of the keel and secondary teeth, and

178

WILD EMMER 179

in the colour and pubescence of the glumes and rachis (Fig. 119). There
is no doubt that some of these forms are hybrids of T. dicoccoides with the
cultivated wheats T. durum and T. vulgare, and probably with the wild
T. aegilopoides which is often found growing with it. However, investiga-
tion of the pure lines established at Reading from ears collected on Mount
Hermon indicate that T. dicoccoides is a good species. The larger size of
the ear, the form and size of the grain, and the character of the pubescence
of the leaves separate it from T. aegilopoides, while the exceptionally easy
disarticulation of the ear, the length and form of the spikelets, the striking
abundance of hair on the rachis, and the shape and size of the grain
distinguish it from cultivated Emmer (T. dicoccum), to which wheat it has
the closest affinity.

T. dicoccoides is an early species coming into ear at Reading about the
last week of May when sown in October or November, in this and other
respects very closely resembling the cultivated varieties of T. dicoccum
from India and Abyssinia.

GENERAL CHARACTERS OF T. dicoccoides, Korn.

The coleoptiles are generally deep purple, though in some cases they
are colourless or pale green. They are remarkable in possessing four
vascular bundles like those of the Indian and Abyssinian forms of T.
dicoccum, which I have no doubt have been derived from this wild species.

The leaves of the young plant are narrow, the first being 2-5-3 mm"
the succeeding ones about 4 mm. broad, some-
what thick, bluish green in colour, the margins
inclined to curve inwards, and the surface
clothed with soft velvety hairs similar in char-
acter and arrangement to those of T. dicoccum
and T. turgidum, arid without the specially long FlG- " 8. — Diagrammatic

. ° • r i transverse sections or

hairs which are seen on the summits or the young leaves of T. dicoc-
ridges of the leaves in T. aegilopoides (Fig. 118). ^fentah' dicoccum> T'

The plant in its early stages of growth re-
sembles couch grass (Agropyrum repens) in colour and general appearance,
and from autumn to late in spring its shoots and leaves are prostrate, lying
quite close to the surface of the soil.

The culm leaves are somewhat rigid and narrow, 4-6 mm. across,
with well-defined, longitudinal, hairy ridges on the upper surface ; the
auricles are long, often purple, and the sheaths are frequently more or
less tinged with purple.

The sheaths in some forms are quite glabrous, or with a fringe of hairs
on the outer edge ; in others they are covered with hairs i mm. or more
long. The thickened nodal portion in many forms is from -8 to i cm. long
and clothed with deflexed hairs as in T. monococcum.

i8o THE WHEAT PLANT

The plants tiller freely and the first few straws expand to full length
a week or ten days before the rest, which are much shorter and remain
green some time after the first ears are ripe. At the time of harvest each
plant exhibits ears dead ripe with the terminal spikelets shed, others just
ripe, some green, as well as a few still enclosed in their leaf-sheaths.

The culms are slender but strong, from 65 to 100 cm. long (about 26-40
inches), with 4 or 5 internodes above ground, the upper one often reaching
a length of 50-60 cm. (about 20-30 inches). In some varieties they are
hollow throughout, in others the upper internode is solid or nearly so.

Before the ears escape from the upper leaf-sheaths the shoots with
the culms enclosed lie close to the surface of the soil, but towards the
middle of May at Reading the shoots rise rapidly, becoming more or
less vertical in about a week ; later they incline again, the ripe ears often
touching the ground.

The bases of the culms of mature plants curve away from each other
in a characteristic decumbent manner, the plants appearing to occupy a
considerable area of ground.

The ears are bearded, stiff, and laterally compressed, measuring 4 to 6
mm. across the face of the spikelets, and 7 to 9 mm. across the two-
ranked side of the ear. They vary in length from 8 to 10 cm. without the
awns and consist of 12 to 17 spikelets ; D = 17-20.

The surface of the flattened rachis is smooth and shining, but along
its edge is a conspicuous fringe of white, yellow, or dark-brown hairs which
are often 5 mm. long, and visible when the ear is viewed from the side ;
in front of the base of each spikelet is a tuft of similar hairs which some-
times extend across the rachis from one margin to the other at that
point.

The lateral spikelets usually have three flowers, the first only, or first
and second of which are fertile ; they are from 15 to 20 mm. long, 5 to 6 mm.
broad, and 3-5 mm. thick.

The terminal spikelet is usually, though not always, sterile, and its
glumes are generally well developed.

The lateral empty glumes, which may be yellowish or pinkish white,
red, uniformly black, striped along the margins or spotted irregularly with
dark brown, are from 10 to 15 mm. long, the outer half 2-4 to 3 mm. across,
the inner half about i mm. broad. They are coriaceous and rigid, usually
5-7 nerved, long and narrow, with a scabrid keel and either glabrous
or covered with silky hair. The apical tooth varies much in length and
form ; in some cases it is bluntish, in others acute and 1-5 mm. long ;
it is usually straight but may curve slightly inwards or outwards (Fig. 1 19).

The strongest nerve on the broad exposed half of the glume converges
towards the base of the apical tooth, where it ends in a secondary tooth
which in some specimens is -5 mm. long, in others very short.

WILD EMMER

181

The empty glumes of the apical spikelet are almost symmetrical with
two strong nerves which terminate in blunt points.

The flowering glume is boat-shaped, 10-15 mm. long and 5-7 mm.
broad, rounded on the back, mostly with 9 nerves, membranous, in colour
like the empty glume, and slightly divided near the apex, the free apical
points being about i mm. long. It possesses a very scabrid and slightly
dorsal awn. Awns are always present on the two lowest flowering glumes
of the spikelet, and are often of nearly equal dimensions, generally 10-12
cm. long, but in some cases reaching a length of 18 cm. or more ;
occasionally the awn of the upper of these two flowers is only about half
the length of that of the lower one, small ears then resembling some
forms of T. aegilopoides .

The palea, which is about as long as the flowering glume, is smooth

1 23456789

FIG. 119. — Empty glumes of forms of T. dicoccoides ( x 2).

and membranous, divided at the tip, two-nerved and bicarinate, the two
ciliate keels often slightly scabrid near the tip.

The flowers resemble those of other wheats and generally have purple
anthers, though in a few forms the latter are yellow.

Anthesis first occurs in the spikelets nearer the apex of the ear than is
the case among cultivated wheats.

The caryopses are reddish, long and narrow, pointed at both ends and
compressed laterally, the dorsal side forming a ridge ; the ventral furrow
is a narrow V-shaped groove, the flanks right and left of the furrow angular.

In profile the ventral side of the grain is a straight line, thus differing
from T. aegilopoides, whose outline is strongly curved on the ventral as well
as the dorsal side.

At the apex of the grain is a tufted " brush" of whitish hairs 1-1-5
mm. long (Fig. 120).

The endosperm is flinty.

182

THE WHEAT PLANT

When two caryopses are present in a single spikelet the one produced
by the lower flower is the smaller.

The grains of the small forms which I consider typical T. dicoccoides are
about 9 mm. long, 17 mm. broad, and 1-85 mm.
thick, the ratios length : breadth : thickness = about
100 : 19 : 20-5 ; 100 grains weigh from 2-2 to 2-8 gr.
Those of the form called " large seeded " by Cook
are among the longest of all wheat grains, equal-
ling in this respect those of
T. polonicum ; they measure
11-12 mm. long, 2-9 mm.
broad, and 27 mm. thick, the
ratios length : breadth : thick-
ness = about 100 : 25 : 23 ;
100 grains weigh from 5 to

FIG. 1 20. — Grains, front,
of T. dicoccoides ; front,
back, and side views
(nat. size).

I am of the opinion that

most of these " large-seeded "

forms are of hybrid origin,

and contain a trace of T.

durum.

The rachis of the ripe ear becomes disarticulated
at its joints on the slightest shake, the spikelets near
the apex becoming detached first, the others break-
ing off in orderly succession towards the base ; the
basal spikelets are frequently green and unripe when
the apical spikelets begin to fall. The short inter-
nodes of the rachis form beaks at the base of the
detached spikelets.

Each flattened internodal beak is from 4 to 5 mm.
long, somewhat convex on one side and correspond-
ingly concave on the other, the base or free end slightly
curved to one side, so that the plane of the disarticu-
lation-scar is almost parallel to the flat side of the
rachis ; the scar is shaped like a narrow segment
of a circle. The detached spikelets after falling to
the ground creep into crevices and bury themselves
in the soil, the hairs on the rachis and the scabrid
awns allowing free motion in a forward direction, but effectually preventing
any backward movement.

In the small-grained forms where two caryopses are present in a
spikelet the upper grain germinates very soon after sowing, but the lower
one remains dormant for a time or refuses to grow at all ; this difference

FIG. 121. — Grains from
the spikelets of one
side of an ear of T.
dicoccoides (nat. size).

FIG. 122. — WILD EMMER (7\ dicoccoides , Korn.).

1. var. Kotschyanum.

2. var. fidvovillosum.

3. var. spontaneonigrum.

4. var. Aaromohni.

WILD EMMER 183

is observed even when the germinating capacity is tested on grains free
from the glumes.

In the large-grained forms both grains germinate at the same time
when freed from the glumes.

VARIETIES OF T. dicoccoides, Korn.

1. Glumes white, glabrous .... var. Kotschyanum, mihi.

2. Glumes white, pubescent .... var. fulvovillosum, mihi.

3. Glumes reddish, glabrous . ... var. Aaronsohni, mihi.

4. Glumes black or striped, glabrous . . var. spontaneonigrum, mihi.

5. Glumes black or striped, pubescent . . var. spontaneovillosum, mihi.

Glumes white, glabrous.

T. dicoccoides, var. Kotschyanum, mihi (Figs. 122, 123).

This is one of the commonest varieties on Mount Hermon. It possesses
glabrous yellowish-white glumes, a conspicuous frontal tuft, and long hairs on the
edges of the rachis. The ears are 6-8 cm. long and have 12-15 spikelets.

T. dicoccoides forma Straussiana, Schulz (Ber. d. Deutsch. Bot. Ges. xxx.
226 (1913), is the western Persian representative of this variety ; forma
Kotschyana, Schulz (!), has a less hairy rachis and is probably a cross or a form
produced by cultivation.

Glumes white, pubescent.

T. dicoccoides, var. fulvovillosum, mihi (Figs. 122, 123).

A common variety received from Mount Hermon. It is similar to var.
Kotschyanum, but with soft white hairs on the glumes and long yellowish or
orange-brown hairs on the rachis. The apical tooth of the empty glume is
generally short and blunt.

I received from Dr. M. Kornicke a very late form which must be classed
with this variety (4, Fig. 123). It has large coarse spikelets 1-5-1-8 cm. long,
the outer glumes clothed with long silvery hairs and terminated by a curved
tooth 4-5 mm. long (9, Fig. 119); the awns are exceptionally stout and 18-20 cm.
long. Figured also by Aaronsohn (Bur. PI. Ind. Bull., No. 180, PI. vii. Fig. i).
It is probably a hybrid of T. dicoccoides and T. durum.

Glumes reddish, glabrous.

T. dicoccoides, var. Aaronsohni, mihi (4, Fig. 122).

T. dicoccum, var. Aaronsohni, Flaksb. Bull. App. Bot., Petrograd, vii. 767
(1914).

This variety resembles var. Kotschyanum, but its glumes are pale brownish
or pinkish-red.

A common large-seeded form of it from El Hadr very closely resembles the
cultivated Emmers (T. dicoccum) of Abyssinia and India in its erect straw, pale
yellowish-green leaves, comparatively glabrous rachis, and early ripening period.

1 84 THE WHEAT PLANT

Glumes black or striped, glabrous.

T. dicoccoides, var. spontaneonigrum, mihi (3, Fig. 122).

T. dicoccum, var. spontaneonigrum, Flaksb. Bull. App. Bot., Petrograd, vii.

768 (1914)-

A frequently occurring variety with smooth shining glumes, which are
uniformly black or striped, with dark brown lines or spots on a yellowish or
reddish-white ground.

Usually the hairs on the rachis are pale yellow or white, but sometimes the
black or deep brown pigment present in the glumes is also found in many of
these hairs.

Glumes black or striped, pubescent.

T. dicoccoides, var. spontaneovillosum, mihi.

T. dicoccum, var. spontaneovillosum, Flaksb. Bull. App. Bot., Petrograd, vii.
768 (1914).

This differs only from the previous one in having villose or pubescent glumes.

All the varieties of T. dicoccoides are found in the neighbourhood of
Mount Hermon ; many of them are found also in other parts of Syria
and Palestine, and var. Kotschyanum extends to Western Persia, but the
distribution of the different varieties is not yet fully known.

As already mentioned, T. dicoccoides is a well-defined species, commonly
self-fertilised, but at the time of anthesis its glumes remain widely separated,
with the stigmas exposed for several hours during the day in bright
weather, and crossing also occurs among the several varieties.

In addition natural hybridisation certainly takes place between this
species and forms of the cultivated wheats T. durum and T. vulgar e ; cross-
ing probably occurs with T. aegilopoides and possibly also with Aegilops
ovata.

To such hybridisation is doubtless due the numerous intermediate
and extraordinarily diverse forms met with in its native habitat, and
among the progeny of plants cultivated at Reading and elsewhere.

Aegilops ovata is a common indigenous species in the wild wheat
region of Palestine, and Cook records the finding of spikelets intermediate
between this species and T. dicoccoides, suggestive of a natural hybrid
between these two plants. I have not seen these.

Triticum aegilopoides, var. Thaoudar, is a common wild species, often
associated with T. dicoccoides, and Aaronsohn mentions the occurrence
of forms morphologically intermediate" between the two. I have also
observed small plants of T. dicoccoides which in glume characters suggest
a relationship with T. aegilopoides. Schulz considers that these are
hybrids of the two species, but without further knowledge of the normal

FIG. 123. — WILD EMMER (T. dicoccoides, Korn.).

i. \zr.Kotschyanum. 2-4. var.fulvovillosum.

WILD EMMER 185

variation of the plants and the experimental production and cultivation
of the hybrid the matter must remain unsettled.

While the majority of ears collected on Mount Hermon, when grown
at Reading, have given forms of T. dicoccoides with constant characters,
many of them have produced a mixed progeny of typical dicoccoides and
a small number of vulgar e and durum-\ike forms, most of which in the
first season are either completely sterile or produce only two or three
fertile grains in an ear.

During the last ten years I have grown T. dicoccoides and found that
these natural hybrids are produced in bright, hot seasons. Apparently
typical ears of T. dicoccoides sometimes give rise to offspring like them-
selves, together with forms of T. durum and T. vulgar e and T. Spelta ;
in such cases one of the grains of a spikelet often produces the typical
wild wheat, while the other grain of the same spikelet gives rise to one or
other of the strikingly different cultivated wheats.

CHAPTER XIII

EMMER

Triticum dicoccum, Schiibler. Char, et Descr. cerealium in Hort. Tub.
29 (1818).

T. Spelta * dicoccon, Schrank. Baier. Fl. i. 389 (1789).

T. Spelta, Host. Gram. Austr. iii. 21, t. 30 (1805), (non Linn.).

T.farrum, Bayle-Barelle. Mon. de Cer. 50, t. 4, Figs, i and 2 (1809).

T. amyleum, Seringe. Melanges botaniques, 124 (1818).

T. Zea, Wagini. Anb. d. Getreid. 33 (1819).

Spelta amylea, Seringe. Cer. Eur. 76 (114), (1841).

T. vulgare dicoccum, Alef. Landw. Fl. 331 (1866).

T. dicoccum, Korn. Handb. d. Getr. i. 81 (1885).

T. sativum dicoccum, Hack. Nat. Pfl. ii. 2, 81, 84 (1887).

THE word Emmer is found in old glossaries of the eighth-thirteenth
centuries as Amer, Amar, and Emmer. Its derivation is obscure, but it
is probably of Celtic origin, for in a glossary to Prudentius it is stated that
" far is a genus of corn which is called ' emerum ' by the Celts of Gaul."

In Germany in mediaeval times this cereal was termed Amelkorn or
Ammelkorn, a name which Bock (1539) and Fuchs (1543) say was given
to it because Amylon or Amylum (starch) was prepared from it ; its
French name Amidonnier is similarly connected with Amylon or Amylum.

Emmer is one of the most ancient cultivated cereals. A beardless ear
of it is described by Heer from the Neolithic pile-dwellings at Wangen
(Switzerland), and Schulz states that remains of it from the Neolithic
period have also been obtained in Denmark, Germany (Bruchsal and
Heidelberg), and Bohemia (Kl. Czernosek).

Portions of ears were found in prehistoric pile-dwellings at Ripac
near Bihac" in Bosnia, and grains attributed to this species by Wittmack
and Buschan were found among the vegetable remains from prehistoric
pile-dwellings of the Bronze Age at Auvernier and Peterinsel (Switzerland).
Fragments of carved models of ears (natural size) in ivory and wood from
the tomb of King Zer-Ta (third king of the 1st Dynasty 5400 B.C.) are
good representations of ears of T. dicoccum, although described as ears of

186

EMMER 187

" bearded barley" in the memoirs of the Egypt Exploration Fund (aist
Memoir, 1901, Plates v. Fig. 16 ; vi. Fig. 17 ; xxxiv. Figs. 82 and 83).

Portions of ears, spikelets, and grain exhumed from the foundations
of the tomb of King Sashu-Re of the Vth Dynasty (3000-4000 B.C.) have
been identified and described by Schulz. Excellent specimens of spikelets
(Fig. 124) with enclosed grain and portions of culms were given to me by
Sir Ernest Budge, who found them in a box in a tomb of the XVIIIth
Dynasty (about 1400 B.C.).

Ears and grain were also discovered in a tomb at Gebelen (Xlth-
XXIst Dynasty), and other examples have been obtained from graves of
the Xllth Dynasty.

Hrozny states that, next to barley, Emmer was the chief cereal culti-

FIG. 124. — Spikelets and grain of T. dicoccum. Upper row recent ; lower row,
ancient Egyptian specimens (1400 B.C.)

vated in ancient Babylon (4000 B.C.) down to the beginning of Persian
rule over the country.

There is no doubt that Emmer was widely grown by prehistoric man
in various parts of Europe and Asia, and was one of the chief cereals
cultivated in Egypt from the earliest times to the Graeco-Roman period
and later, when it appears to have been largely supplanted by macaroni
and ordinary bread wheats.

The bote of the Ancient Egyptians, oAvpa (olura) and £ea or £eta (zea
or zeia) of early Greek authors, and the far of ancient Roman writers
on agriculture, were forms of this " spelt " wheat, and not T. Spelta, with
which it is so often confused by translators of the Classics.

Although an important cereal in the earliest times, its cultivation is
now restricted to comparatively small dimensions, and is of little moment

1 88 THE WHEAT PLANT

in comparison with the area devoted to ordinary bread and macaroni
wheats. In districts in South Germany in which T. Spelta is extensively
grown small areas are devoted to T. dicoccum, in place of barley as a
spring-sown cereal, and a small amount is raised in Switzerland, France,
Italy, Serbia, Spain, and the United States. In some of these countries a
little of the free grain is utilised for human food, but the greater portion
grown there is fed in the husk to horses and other stock in place of oats
and barley. It is cultivated on a much larger scale for human food in
Russia and Abyssinia, and is also grown for the same purpose in moderate
quantity in India, especially on irrigated land in Madras, Mysore, Bombay,
Berar, and the Central Provinces.

It was apparently grown in Egypt in small quantities about the begin-
ning of the nineteenth century, but is not found there in cultivation now\
There are reports of it as a cultivated crop among the Kurds in the pro-
vince of Luristan in Western Persia (Haussknecht, 1868) and in Yeman,
Arabia (Glaser, 1891).

I have no records of its cultivation at the present day in Asiatic Turkey
nor in China.

Emmer will grow on soils which are too light to yield a good crop of
T. Spelta. The method of cultivation adopted for the latter is applicable,
however, to Emmer (see p. 326). It is more or less immune to rust fungi
and suited to warm, dry climates ; the majority of forms are rapid-growing
spring varieties, with erect caespitose habit and little power of resisting frost.

One late variety with black glumes, however, is more hardy, and is a
typical winter form whose young shoots lie close to the surface of the
ground.

The yield per acre varies between twenty-five and fifty bushels of
" spelt " grain.

The grains are starchy and the flour obtained from them is especially
white and highly esteemed for the manufacture of the finest pastry and
cakes. As an addition to soup, they are superior to " pearl " barley or
Dinkel grain. A fine starch can be prepared from the flour.

GENERAL CHARACTERS OF T. dicoccum, Schiibl.

The young plants of all varieties except Black Winter Emmer have an
erect habit, with comparatively broad leaves, measuring 7-8 mm. across.
The wheats belonging to this race fall into two groups, viz. :

1. Indo- Abyssinian Emmers ; all early forms, with four- to six-nerved

coleoptiles, yellowish-green leaves, short straw, ears with brittle
or tough rachis.

2. European Emmers, later in growth, with two-nerved coleoptiles,

glaucous leaves, taller straw, and fragile rachis.

EMMER 189

In the European Emmers the leaf-blades are covered with soft hairs
(Fig. 118), and are bluish-green in tint with large fringed auricles. The
Indian and Abyssinian forms possess paler yellowish-green culm leaves,
which have fewer and shorter hairs on the ridges, and the auricles are
frequently quite glabrous.

The leaf-sheaths and auricles of most Emmers are pinkish. The
plants tiller moderately. The straw is strong and somewhat slender ;
in some varieties it is hollow, in others the upper internode is solid. Most
European forms reach a height of 100-125 cm- (about 39-50 inches) ;
the Indian and Abyssinian forms are rarely taller than 55-65 cm. (about
22-26 inches) ; the full-grown culm leaves have a breadth of about
16 mm.

The ears, which are white, red, or black, are usually bearded, erect, and
laterally compressed, although in a few kinds the cross section of the ear
is almost square. Those of different varieties vary in length from 5 or 6
cm. to over 10 cm., with fifteen to thirty or more spikelets ; the density
varies between wide limits (0 = 24-40).

The rachis is flattened and smooth, more or less hairy along the margins
with a tuft in front between the empty glumes ; each individual segment
is narrow at the base, widening upwards in some cases to about twice the
breadth at the top. In some forms it is fragile and the grains firmly
enclosed in the glumes ; in others of the Indo-Abyssinian sub-race the
rachis is tough and not so readily disarticulated, and in these the grains
are more loosely invested by the chaff.

In the " spelt " forms the rachis becomes disarticulated at the narrowest
basal end of each internode, each separate spikelet carrying with it the
internodal portion immediately below it (Fig. 124).

The spikelets, which are more or less closely imbricate on the rachis,
are oval, from 5 mm. to 8 mm. across, and twice as long, containing three
or four flowers, the two lower ones usually producing grain, those above
being rudimentary.

The empty glumes are oval and boat-shaped, narrow towards the apex,
seven-nerved, with a prominent keel running from base to apex and
ending in a straight or curved tooth, which varies in length and form in
different varieties : in one form the empty glume has a long awn (Fig.
125). The outer half is somewhat flat and about twice as wide as the
inner half of the glume ; upon it is a lateral nerve, which divides the outer
half almost equally, and ends near the base of the apical tooth, sometimes
in a short point.

The flowering glumes are boat-shaped, rounded on the back, with
nine to eleven nerves, but without a keel ; they are usually terminated
by scabrid three-angled awns from 5 to 14 cm. in length. The awn of the
lowest flowering glume of a spikelet is longest, that of the second slightly

THE WHEAT PLANT

shorter, a very short one or none at all being found on the glumes enclosing
abortive flowers.

The palea is equal to, or slightly longer than, the flowering glume,
ovate-lanceolate, with a narrow bifid apex, and an inward fold between
its two keels, the latter ciliate, especially near their tips. The two margins

123 456 7

8 9 10 ii 12

FIG. 125. — Empty glumes of Emmer (T. dicoccum) ( x 2).

of the palea, which curve inward round the grain, are transparent, with a
single nerve running through each.

The caryopses of the " spelt " forms are firmly enclosed by the glumes,
and do not thrash out as naked grains ; in other forms the grain is readily
separated from the chaff on thrashing.

EMMER 191

Two, or occasionally three, are present in each spikelet.

The colour of the grain is chiefly white, yellowish, or red as in other
races, but in recently harvested grains of the Abyssinian varieties Arraseita
and Schimperi it is a deep purple tint, the anthocyan pigment being present
in the " chlorophyll layer " of the peri-
carp, and to a lesser extent in some of the
adjacent parenchyma.

The grains are comparatively narrow

Mtt
MM
M M

FIG. 126. — Grains of Emmer (T.
dicoccum). Front, back, and
side views (nat. size).

and pointed at both ends and more or less
laterally compressed, with a flattish or hollowed
ventral surface and a narrow furrow (Figs.
126, 127) ; each has a conspicuous " brush "
of hairs. A cross section of the grain is some-
what triangular, with fairly distinct basal angles.
In some varieties the endosperm is flinty, in
others mealy.

Well-developed grains measure from 7-2 to
9 mm. in length, 2-85 to 3-4 mm. in width, and
2-6 to 3-1 mm. through from front to back. They
are slightly more compressed laterally than the
grains of Dinkel or Large Spelt (T. Spelta) ;
the average ratio of length, breadth, and thick-
ness = 100 : 37-7 : 34-1.

The produce of the thrashed ear of the
" spelt " forms, i.e. the " husked " grain or
' Vesen," consists of 73 to 75 per cent of cary-

FIG. 127.— Grains of the opses and 2I to 2r per cent of chaff (glumes
spikelets or one side or , _ . * . • i r

an ear of Emmer (T. and pieces or the rachis), and weighs from 40

dicoccum) (nat. size). to ^ fcg per hectolitre.

192

THE WHEAT PLANT

VARIETIES OF T. dicoccum, Schubl.
GROUP I. — INDO- ABYSSINIAN EMMERS. Coleoptile 4- (3-6) nerved.

SECTION A. — Speltae. Rachis fragile,
i . Ears bearded —

(i) Glumes white, glabrous ; grain red .... var. Ajar, mihi

SECTION B. — Tenaces. Rachis tough,
i . Ears bearded —

(1) Glumes white, glabrous ; awns white ;

grain red ......

(2) Glumes white, glabrous ; awns black ;

grain red ......

(3) Glumes white, glabrous ; grain purple

(4) Glumes white, pubescent ; grain white

(5) Glumes red, glabrous ; grain red

(6) Glumes red, glabrous ; grain purple

(7) Glumes dark brown or mottled, pubescent

var. uncinatum, mihi.

var.pseudo-uncinatum, mihi.
var. Arraseita, mihi.
. var. tomentosum, mihi.
var. rufescens, mihi.
var. Schimperi, mihi.
var. persicum, mihi.

GROUP II. — EUROPEAN EMMERS. Coleoptile 2-nerved.
i . Ears bearded —

(1) Glumes white, glabrous.

i. Awns short .....
ii. Awns long.

a. Ears simple * narrow

** broad and dense

b. Ears branched ....

(2) Glumes white, pubescent.

i. Awns short.

a. Ears simple

b. Ears branched .
ii. Awns long.

a. Narrow lax ears.

* Awns straight
** Awns bent, often black .

b. Broad dense ears ; black awns
(3) Glumes red, glabrous.

i. Awns short.

a. Ears simple

b. Ears with double spikelets .

c. Ears branched .
ii. Awns long.

a. Ears simple.
* Narrow
** Broad and dense .

var. tricoccum, Korn.

var.farrum,
. var. liguliforme,
var. albiramosum,

var. submajus,
var. Metzgeri,

var. semtcanum,

var. flexuosum,

var. majus,

var. Fuchsit,
var. Dodonaei,
var. cladurum,

var. rufum,
var. pycmirum,

Korn.
Korn.
Korn.

Korn.
Korn.

Korn.
Korn.
Korn.

Korn.
Korn.
Korn.

Korn.
Korn.

FIG. 128 — EMMER (T. dicoccum, Schiibl.).

var. Ajar.
Indian Emmer (i, non-irrigated ; 2, irrigated plant).

EMMER 193

b. Ears with double spikelets . . . var. Schubleri, Korn.

c. Ears branched ..... var. erythrurum, Korn.

(4) Glumes red, pubescent.

i. Awns short.

a. Ears simple ..... var. Bauhini, Korn.

b. Ears with double spikelets . . . var. Tragi, Korn.

c. Ears branched ..... var. Krausei, Korn.
ii. Awns long.

a. Ears simple .... var. macratherum, Korn.

b. Ears with double spikelets . . . var.'Mazzucati, Korn.

c. Ears branched .... var. rubriramosum, Korn.

(5) Glumes black, pubescent.

i. Awns short . . . . . . var. subatratum, Korn.

ii. Awns long.

a. Ears simple ..... var. atratum, Korn.

b. Ears branched ..... var. melanurum, Korn.

2. Ears beardless or with short awns —

(1) Glumes white, glabrous.

a. Ears simple ...... var. inerme, Korn.

b. Ears with double spikelets . . var. bispiculatum, Korn.

c. Ears branched .... var. leucocladon, Korn.

(2) Glumes white, pubescent.

a. Ears simple ...... var. muticum, Korn.

(3) Glumes red, glabrous.

a. Ears simple ...... var. novicium, Korn.

b. Ears branched . . . . var. subcladurum, Korn.

(4) Glumes red, pubescent.

a. Ears simple ...... var. hybridum, Korn.

GROUP I. — INDO- ABYSSINIAN EMMERS

SECTION A. — Speltae. Rachis fragile.

Ear bearded ; glumes white, glabrous ; grain red.

T. dicoccum, var. Ajar, mihi.

T. Arras, Hochst. Flora, 31, 450 (1848).

i. Abyssinian Emmer (2, Fig. 129). — This variety I obtained under the
native Amharic name " Ajar " or " Agga," from Abyssinia, where it is grown
at an elevation of 5000-8000 feet. It is sown in June and harvested in
October, giving a six- to eight-fold return ; where irrigated it is sown in
November and December and reaped in May.

It is closely related to the wild T. dicoccoides and, like it, is exceptionally early,
coming into ear at Reading about May 20-25 when sown in autumn (October-
December).

Coleoptiles, 4- to 6-nerved, sometimes pinkish.

O

i94 THE WHEAT PLANT

Young shoots, erect ; young leaves pubescent ; the blades of the culm leaves
less pubescent and a yellow-green tint ; lower leaf-sheaths pink ; auricles
glabrous or with a few long hairs only.

Straw, very short, rarely more than 60 cm. (23-24 inches) long.

Ears, short, 6-7 cm. long, on non-irrigated land ; flat, never glaucous, but a
characteristic yellow-green colour becoming white or pinkish, like some glabrous
forms of T. hermonis.

Rachis very fragile, thin, fringed along its edges with short hairs ; frontal
tufts short. Spikelets 16-20 ; 10-12 mm. long, 5-7 mm. broad, containing one
or two pale reddish flinty grains ; D = about 30 ; awns 9-12 cm. long.

Empty glume, 10 mm. long, narrow, 2-5-3 mm. across, keeled, apex more or
less truncate with a short blunt apical tooth ; secondary tooth and nerve distinct.

Grain, pale reddish, flinty, 9-10 mm. long, 3 mm. broad.

2. Indian Emmer ; Khapli (Fig. 128). — This variety, sometimes erroneously
described as T. Spelta, is grown in the Central Provinces, Madras, Bombay,
and Mysore. It agrees in all its characters with Abyssinian Emmer, and the
two forms I have no doubt have had a common origin.

SECTION B. — Tenaces. Rachis more or less tough.
Ear bearded ; glumes white, glabrous ; awns white ; grain red.

T. dicoccum, var. uncinatum, mihi.

An early variety received from Abyssinia under the name Nach Sinde.
Two forms appeared, viz. :

1. FORM i (2, Fig. 131).

Young shoots, erect ; young leaves pubescent.

Straw, slender, short, solid or hollow with thick walls.

Ear, 5-9 cm. long, square, 10 mm. across the sides ; spikelets 14-20, some-
times 3-grained ; D = about 25 ; awns slender, 3-5 cm. long.

Empty glume, 8-9 mm. long, keeled, apical tooth strong, curved inwards (i,
Fig. 125).

Grain, flinty or semi-flinty, compressed laterally with prominent dorsal
hump ; apex bluntish ; " brush " short, 6-5-7-4 mm. long, 2-6-3-3 mm.
broad, 2-9-4 mm. thick.

2. FORM 2. — An early variety received from Abyssinia in a sample of the
violet-grained var. Arraseita (see below), from which it differs only in the colour
of its grain.

Young shoots, erect ; young leaves pubescent.

Straw, short, slender, 56-66 cm. (about 22 inches) long, upper internode
hollow with thick walls.

Ear, slender, 8-9 cm. long, square, 10-1 1 mm. across the sides ; spikelets 23 ;
D = 3i ; awns 10-13 cm. long (Ear type, Fig. 130).

Empty glume, 9-10 mm. long, apical tooth narrow, straight or slightly curved,
acute (2 and 3, Fig. 125).

Grain, flinty, with pointed apex 8-8-6 mm. long, 3-4 mm. broad, 3-6 mm.
thick.

FIG. 129. — EMMER (T. dicoccum, Schiibl.).

i. var.farrum.
(White Russian Emmer.)

2. var. Ajar.
(Abyssinian Emmer.)

EMMER 195

Ear bearded ; glumes white with dark margins, glabrous ;
awns black ; grain red.

T. dicoccum, var. pseudo-uncinatum, mihi.

This variety is similar to Form 2, var. uncinatum, but usually has a black
stripe along the outer margin of the empty glume ; in some seasons the whole
glume is a dark brown tint. It was obtained from the same source as Form 2,
var. uncinatum.

Ear bearded ; glumes white, glabrous ; awns white ; grain violet.

T. dicoccum, var. Arraseita, mihi.

T. durum, var. Arraseita, Hochst ex Korn. Handb. d. Getr. i. 70 (1885).

T. durum, var. Hildebrandti, Wittmack. Ber. Naturf.-Vers. Baden-Baden,
211(1879)?.

T. " Eloboni," Caporn. Journal of Genetics, vii. 259 (1918).

Hochstetter's plants were grown from a sample of violet-coloured grains sent
from Abyssinia by W. Schimper under the name " Arraseita." From the same
sample var. Schimperi (p. 196) was also obtained.

Tukur Sinde (Fig. 130).— This variety is a very distinct, early short-strawed
form with violet-coloured grain, received from Abyssinia under the name
' Tukur Sinde."

Young shoots, erect ; young leaves more or less pubescent.

Straw, short to medium height, 76-90 cm. (about 30-36 inches) long, upper
internode solid.

Ear, slender, narrow, 8-5-9 cm- l°ng> compressed, 5-6 mm. across the face,
8-9 mm. across the side; spikelets 21, 2 -grained ; density variable = 20-30 ;
awns 10 cm. long.

Empty glume, narrow, 9 mm. long ; sometimes with a dark marginal stripe,
apical tooth narrow, acute (2, 4, Fig. 125).

Grain, narrow, a- rich purple tint when fresh ; 8-5-8-9 mm. long, 3-5 mm.
broad, 3-2 mm. thick.

Ear bearded ; glumes white, pubescent ; grain white.

T. dicoccum, var. tomentosum, mihi.

A characteristic variety with somewhat dense narrow ears, received from
Abyssinia in a sample named Nach Sinde (i, Fig. 131).

Young shoots, erect ; young leaves pubescent.

Straw, slender, stiff, hollow with thick walls.

Ear, small, dense, 5-6 cm. long, oblong in section, 8-9 mm. across the face,
ii mm. across the two-rowed side ; spikelets 19-20 ; D =35-36 ; awns 6-8 cm.
long.

Empty glume, narrow, 9-10 mm. long, 2-2-5 mm- broad, apical tooth blunt,
short, curved outwards, secondary notch and vein prominent (6, Fig. 125).

Grain, white or amber, flinty, compressed laterally, apex blunt, brush very
short ; 7-7 mm. long, 2-4-2-7 mm. broad, 2-9-3-1 mm. thick.

196 THE WHEAT PLANT

Ear bearded ; glume red, glabrous ; grain red.

T. dicoccum, var. rufescens, mihi.

Received from Abyssinia in a sample of Nach Sinde.

Young shoots, erect ; young leaves pubescent.

Straw, very slender, short, hollow with thick walls.

Ear, narrow, lax, 7-8 cm. long, square, 8-10 mm. across the sides ; spikelets
14-18 ; D = about 25. Awns slender, spreading somewhat, 9-10 cm. long (Ear
type 2, Fig. 130).

Empty glume, 9-10 mm. long, keeled throughout ; apical tooth short, blunt ;
secondary notch distinct (5, Fig. 125).

Grain, flinty, with blunt apex, " brush " short, dorsal hump prominent ;
7-75 mm. long, 3-45 mm. broad, 3-3-3-4 mm. thick.

Ear bearded ; glumes red, glabrous ; grain purple.

T. dicoccum, var. Schimperi, mihi.

T. durum, var. Schimperi, Korn. Handb. d. Getr. i. 71 (1885).

Kornicke states that this purple-grained variety was sent from Abyssinia by
Schimper in a mixed sample of var. Arraseita, which it resembles in all its
characters except the colour of its glumes.

Ear bearded ; glumes dark brown or mottled, pubescent.
T. dicoccum, var. persicum, mihi.

Persian Black Wheat (Ear type i, Fig. 130). — A remarkable wheat obtained
from Haage and Schmidt, Erfurt, Germany ; very late, not coming into ear at
Reading until the second or third week in June. It has been described as a
variety of T. vulgare, var. fuliginosum, but its 3- to 4-nerved coleoptile,
pubescent, grass-green leaves, slender thick-walled or solid straw, narrow rachis,
and general form of ear place it near the Abyssinian Tukur Sinde (p. 195).

Vavilov found it perfectly immune to Erysiphe graminis, and in this respect
quite unlike 580 sorts of T. vulgare which he tested at the same time.

The hybrid (Fx) with T. vulgare is almost sterile.

Young shoots, erect ; young leaves pubescent.

Straw, thin, of medium height, 104 cm. (about 42 inches), solid or hollow
with thick walls ; nodes hairy.

Ear, very lax, narrow, 9 cm. long, square, 10 cm. across the sides ; spikelets
2- to 3-grained, 19-20 ; D = about 20.

Empty glume, with a long awn 3-4 cm. long (7, Fig. 125).

Flowering glume, white below, dark brown above, with awn 8-n cm. long.

Grain, flinty, reddish ; apex blunt ; 6-5 mm. long, 3 mm. broad, 2-9 mm.
thick.

GROUP II. — EUROPEAN EMMERS.
Ear bearded ; awns short ; glumes white, glabrous.

T. dicoccum, var. tricoccum, Korn. Handb. d. Getr. i. 86 (1885).

T. tricoccum, Schubl. Char, et descr. Cer. 33 (1818), and Flora, 445 (1820).

FIG. 130. — EMMER (T. dicoccum, Schtibl.).

var. Arraseita.

Tukur Sinde : Abyssinian Purple-grained Emmer.
(i, lax ; 2, dense-eared form.)

EMMER 197

T. amyleum, G., Metzger. Eur. Cer. 33, Taf. 7 B. (1824) ; Landw. Pfl. i. 114

T. amyleum, Ser., Krause. G^r. Heft V. 13, t. 5 D. (1837).
T. vulgar e subtricoccum, Alef. Landw. Fl. 332 (1866).

Egyptian Spelt ; Egyptian Winter Wheat. — A somewhat hardy form,
described by Schiibler, Metzger, and others. I have not seen it, but think
it probable that it is similar to Form i, var. uncinatum, mihi, and should be
placed in Group I.

Straw, 120-150 cm. high ; leaves yellowish-green.

Ear, 10-12 cm. long, with short awns. D = 28. The spikelets are about i
cm. broad and contain two or three reddish flinty or mealy grains 9 mm. long
and 4 mm. broad.

Now rare, but formerly grown in Egypt, Italy, south of France, and the Jura
in Switzerland.

Ear bearded, narrow ; awns long ; glumes white, glabrous.

T. dicoccum, var. farrum, Korn. Handb. d. Getr. i. 87 (1885).

T.farrum, Bayle-Barelle. Mon. d. Cer. 50 (1809).

T. dicoccum album, Schiibler. Char, et descr. Cer. 29 (1818), and in Flora
Jahr. 3, ii. 450.

The commonest European variety of T. amyleum. Three forms of it are
distinguished.

1. White Emmer (T. amyleum, A., Metzger, Eur. Cer. 30) (i, Fig. 132). —
A late form found in small quantities in France, Germany, Austria, Switzer-
land, Italy, and Spain. It comes into ear at Reading from June 10 to 20.

Young shoots, erect ; young leaves pubescent.

Straw, 100 cm. (40 inches) long.

Ear, dense, 7-9 cm. long, narrow, 9 mm. across the 2-rowed side ; spikelets
27-28 ; 0 = 38 ; awns 6-8 cm. long.

Empty glume, 8-9 mm. long, 3 mm. broad, keeled to the base, apex with long,
curved, claw-like tooth (9, Fig. 125) ; secondary lateral nerve prominent, ending
in a short tooth, near the base of the keel tooth.

Grain, flinty, brownish-red ; 7-8 mm. long, 3 mm. broad.

2. Large White Emmer (T. amyleum, B., Metzger, Eur. Cer. 31) (2, Fig.
132). — A taller form found with the preceding variety ; it is about a week later
in ripening.

Straw, 130 cm. (50 inches) long.

Ear, larger and laxer, 10-12 cm. long, 1-1-2 cm. broad, keeled, spikelets about
30 ; 0 = 31. The edges of the rachis are fringed with long hairs ; spikelets
12-13 mm. long, 7 mm. broad ; awns 7-9 cm. long.

Empty glume, 9-10 mm. long, 4 mm. broad ; apex narrow ; apical tooth short,
straight, acute ; secondary nerve ending in a blunt tooth (8, Fig. 125).

Grain, semi-flinty, reddish-yellow, 8-9 mm. long, 3-5 mm. broad, with pro-
minent dorsal ridge.

198 THE WHEAT PLANT

3. Slav Emmer (i, Fig. 129). — A small-eared form cultivated in Serbia and
Russia, chiefly in the provinces adjacent to the Volga and to a lesser extent in
the Caucasus, Siberia, and Turkestan. It is somewhat earlier than the preceding
form, coming into ear at Reading about ten days sooner.

Young shoots, erect ; young leaves pubescent.

Straw, 90-95 cm. (36-38 inches) long.

Ear, small and short, 5-7 cm. long, i cm. across the 2-rowed side ; spikelets
23-25 ; 0 = 38-40; awns 6-8 cm. long.

Empty glume, 8 mm. long, 3 mm. broad, apex with short blunt tooth (12,
Fig. 125).

Gram, red and flinty, pointed at both ends, " brush " long, dorsal ridge
prominent ; 7-5 mm. long, 2-5-2*7 mm. broad, 2-4-2-6 mm. thick.

Ear bearded, broad, dense ; glumes white, glabrous.

T. dicoccum, var. lig-uliforme, Korn. Handb. d. Getr. i. 90 (1885).

T. amyleum, Ser., Krause. Getr. Heft V. 9, t. 4 C. D. (1837).

A variety with short dense ears 5 cm. long and 1-8 cm. broad, frequently bent
over on the flat side ; the awns 12-15 cm. long. Kornicke states that he
received this variety from Jena.

Ear bearded, branched ; glumes white, glabrous.

T. dicoccum, var. albiramosum, Korn. Arch. f. Biontologie, ii. 411
(1908).

Apparently a hybrid discovered by Kornicke in the Botanic Garden at
Poppelsdorf in 1901.

Ear. with short awns ; glumes white, pubescent.

T. dicoccum, var. submajus, Korn. Arch.f. Biontologie, ii. 409 (1908).

Ears like var. majus, but with short black or white awns.

Found by Kornicke in a mixture in the Botanic Garden, Poppelsdorf.

Ear bearded, branched ; awns short ; glumes white, pubescent.

T. dicoccum, var. Metzgeri, Korn. Handb. d. Getr. i. 91 (1885).

T. amyleum, Ser., Krause. Getr. Heft V. 15, Taf. 6 C. (1837).

T. amyleum, D., Metzger. Eur. Cer. 32 (1824), and Landw. Pfl. i. 116 (e)
(1841)-

T. vulgare Metzgeri, Alef. Landw. Fl. 332 (1866).

This variety is not constant ; its simple ears resemble those of the preceding
variety.

Ear bearded, lax ; awns long, straight ; glumes greyish-white, pubescent.

T. dicoccum, var. semicanum, Korn. Handb. d. Getr. i. 89 (1885).
T. amyleum, C., Seringe. Mtl. hot. 128 (1818).
T. amyleum, C., Metzger. Eur. Cer. 32 (1824).

FIG. 131. — EMMER (T. dicoccum, Schiibl.).

I. var. tomentosum.
(Abyssinia.)

2. var. uncinatum.
(Abyssinia.)

EMMER 199

T. amyleum, Ser., Krause. Getr. Heft V. 47 (1837).

The ears of this variety are short, narrow, and lax, 5-6 cm. long, 1-2 cm.
broad, and the awns blackish. The grains are pale red, flinty, and loosely
invested by the glumes. Received by Kornicke from Jena, and believed by
him to be descended from Krause's stock.

Ear bearded, narrow, lax ; awns long, usually black and bent ;
glumes white, pubescent.

T. dicoccum, var. flexuosum, Korn. Handb. d. Getr. i. 88 (1885):
A white velvet-chaffed form, with very lax fragile ears and awns 8-10 cm.
long, generally black and bent at the base. The grain is large, red, and flinty,
7-5 mm. long, 3-5 mm. broad, and loosely invested by the glumes.

Kornicke received it from Jena mixed with the straight-awned var. semi-

Ear bearded, broad, dense ; awns long, black ; glumes white, pubescent.

T. dicoccum, var. majus, Korn. Handb. d. Getr. i. 89 (1885).

T. amyleum, Ser., Krause. Getr. Heft V. 8, t. 3 A. (1837).

Received by Krause from a Botanic Garden under the title " T. vulgar e or
turgidum cochleare."

A very tall, strong-growing variety 140-150 cm. high, with broad ears 10-12
mm. long, tapering upwards, and consisting of 22-24 spikelets containing pale
reddish-yellow mealy or semi-flinty grains 8 mm. long, 3-5 mm. broad, loosely
held in the glumes. The awns are dark brown or black and 15-17 mm. long.

Received by Kornicke from Hohenheim.

Ear with short awns ; glumes red, glabrous.

T. dicoccum, var. Fuchsii, Korn. Handb. d. Getr. i. 87 (1885).

T. vulgar e Fuchsii, Alef. Landw. Fl.^2 (1866).

A variety obtained in 1824 by Metzger from the white-chaffed var. Metzgeri.

Ear with double spikelets and short awns ; glumes red, glabrous.
T. dicoccum, var. Dodonaei, Korn. Arch.f. Biontologie, ii. 410 (1908).

Ear branched ; awns short ; glumes red, glabrous.

T. dicoccum, var. cladurum, Korn. Handb. d. Getr. i. 90 (1885).

T. amyleum, Ser., Krause. Getr. Heft V. ii. t. 5 C. (1837).

T. vulgar e cladura, Alef. Landw. Fl. 333 (1866).

Alefeld obtained this variety from var. Fuchsii.

Kornicke received it from a Botanic Garden and states that only a certain
proportion of the ears are branched, and in these the branching is not pro-
nounced.

The commonest form of it has very tall straws, 130-140 cm. long, branched
ears with a toughish rachis and awns 4-5 cm. long. The grains are reddish-
yellow and mealy, small, 6 mm. long, 3~3'5 mm. broad.

200 THE WHEAT PLANT

Ear bearded, narrow ; awns long ; glumes red, glabrous.

T. dicoccum, var. rufum, Korn. Handb. d. Getr. i. 88 (1885).

T. dicoccum rufum, Schvibl. Char, et descr. Cer. 29 (1818).

T. amyleum, Ser., Krause. Getr. Heft V. 5, Taf. 2 A. B. (1837).

T. amyleum, E., Metzger. Eur. Cer. 32 (1824), anc^ Landw. Pfl. i. 114 (b)
(1841).

T. vulgar e brunneum, Alef. Landw. Fl. 331 (1866).

This variety has red glumes, but in other respects resembles the white-
glumed var.farrum, with which it is generally found mixed.

I have met with two forms, viz. : (i) a small-eared, short-strawed Serbian
and Russian form with blunt-ended empty glumes (i, Fig. 133), and (2) the
Western European Red Emmer (2, Fig. 133), with taller straw, larger ears, later
habit, and empty glumes, with a claw-like apical tooth similar to that of White
Emmer (p. 197).

Ear bearded, broad, and dense ; awns long ; glumes red, glabrous.

T. dicoccum, var. pycnurum, Korn. Handb. d. Getr. i. 90 (1885).

T. vulgar e pycnur a, Alef. Landw. Fl. 333 (1866).

Two forms are recognised, viz. :

FORM i. (T. amyleum, I., Metzger, Eur. Cer. 34) with pale red short ears
4-5 cm. long, 1-6 cm. broad, tapering towards both ends ; awns 16-18 cm. long ;
spikelets occasionally with three grains, which are less firmly enclosed in the
glumes than in most Emmers.

Sent to Kornicke from Hohenheim as T. Cienfuegos and from some Botanic
Gardens as T. Hisbu.

FORM 2 has dark red, longer ears 5-7 cm. long, 1-4 cm. broad, and widest at
the apex ; awns 12-16 cm. long (figured by Krause, Getr. Heft. V. 9, t. 4
A. B.).

Ear bearded with doube spikelets ; awns long ; glumes red, glabrous.

T. dicoccum, var. Schiibleri, Korn. Arch.f. Biontologie, ii. 410 (1908).

Found by Kornicke in 1899 in the Botanic Garden at Poppelsdorf among
var. erythrurum ; he states that the grains are loosely held by the chaff, and pre-
sumes it is a hybrid with some form of T. vulgar e.

Ear branched ; awns long ; glumes red, glabrous.

T. dicoccum, var. erythrurum, Korn. Handb. d. Getr. i. 91 (1885).

T. amyleum, F., Metzger. Eur. Cer. 33 (1824).

T. vulgare phaeocladus, Alef. Landw. FL 332 (1866).

Kornicke 's examples were derived from a cross.

Ear with short awns ; glumes red, pubescent.

T. dicoccum, var. Bauhini, Korn. Handb. d. Getr. i. 87 (1885).
T. amyleum, H., Metzger. Eur. Cer. 33 (1824).
T. vulgare Bauhini, Alef. Landw. Fl. 332 (1866).

FIG. 132.— EMMER (T. dicoccum, Schiibl.).

var. farrum.

i. White Emmer. 2. Large White Emmer.

(European forms.)

EMMER 201

The straw of this variety is very tall, 140-150 cm. (55-60 inches) high, ears
10-12 cm. long, with a toughish rachis and small reddish-yellow mealy grains
7 mm. long and 3-5 mm. broad, loosely invested by the glumes.

Obtained from a Botanic Garden by Kornicke.

Ear with short awns and double spikelets ; glumes red, pubescent.

T. dicoccum, var. Tragi, Korn. Handb. d. Getr. i. 90 (1885).
Kornicke's type was obtained from a Botanic Garden.

Ear branched ; awns short ; glumes red, pubescent.

T. dicoccum, var. Krausei, Korn. Handb. d. Getr. i. 91 (1885).

A common form of this variety has very tall straw, 140-150 cm. (55-60 inches)
high ; the ears are dense and branched at the base, with small reddish-yellow
mealy and flinty grains 7 mm. long, 2-5 mm. broad, which are loosely enclosed
by the glumes. The short slender awns are pale red and are only found on the
flowering glumes of the lower of the two flowers of each spikelet.

Obtained from a cross.

Ear bearded ; awns long ; glumes red, pubescent.

T. dicoccum, var. macratherum, Korn. Handb. d. Getr. i. 89 (1885).
Kornicke's type was obtained from a Botanic Garden.

Ear with double spikelets, bearded ; awns long ;• glumes red, pubescent.

T. dicoccum, var. Mazzucati, Korn. Arch.f. Biontologie, ii. 410 (1908).
Originated in 1895 from a form with branched ears. Kornicke suggests
that it arose from the crossing of a branched Emmer with T. vulgare.

Ear branched, bearded ; awns long ; glumes red, pubescent.

T. dicoccum, var. rubriramosum, Korn. Arch.f. Biontologie, ii. 411
(1908).

Appeared in the Botanic Garden, Poppelsdorf among a sowing of var.
albiramosum in 1904.

Ear bearded ; awns short ; glumes blue-black, pubescent.

T. dicoccum, var. subatratum, Korn. Arch.f. Biontologie, {1.409 (1908).
Discovered by Kornicke in 1884 in the Botanic Garden, Poppelsdorf.

Ear bearded ; awns long ; glumes blue-black, pubescent.

T. dicoccum, var. atratum, Korn. Handb. d. Getr. i. 89 (1885).

T. atratum, Host. Gram, austr. iv. 5 and 8 (1809).

T. atratum, Roem. et Schult. Syst. vegetab. ii. 766, No. 15 (1817).

T. atratum, Schubl. Char, et descr. Cer. 32 (1818).

T. amyleum, D., Ser. Msl. hot. 129 (1818).

202 THE WHEAT PLANT

T. amyleum, K., Metzger. Eur. Cer. 32 (1824) ; Landw. Pfl. i. 117 h. (1841).
T. amyleum, Ser., Krause. Getr. Heft V. 3 PI. I. A. B. (1837).

Black Emmer (3, Fig. 133). — A common representative of this variety is
Black Winter Emmer, a very late form, the ears of which do not escape from
the upper sheath until about June 20 at Reading.

Kornicke states it is of African origin, and Heuze says it is believed to have
come from Abyssinia.

Although much more hardy it yields less grain than the white-chaffed forms
of var. farrum.

Small plots of Black Winter Emmer appear to be grown in France and
Germany.

It was introduced from France in 1904 into the United States, where it is
sometimes grown as food for stock in place of oats and barley.

Young shoots, prostrate in winter and early spring.

Straw, of medium height to tall, about 102-115 cm. (40-46 inches) long.

Ear, very dense and flat, broad at the base, tapering towards the apex ; 7-8
cm. long, about i«6 cm. broad, with 26-28 spikelets ; D=46. The spikelets
are 10-12 mm. long, 5 mm. broad, each with two small reddish-yellow, opaque,
mealy grains 6-7 mm. long, 2-5 mm. broad.

Empty glume, 9-10 mm. long, 3 mm. broad, bluish-black or brownish with
a glaucous hue and clothed with fine hairs, the colour of the hidden portions
of the flowering glumes being a pale reddish-white. The keel terminates in a
short bluntish tooth (n, Fig. 125).

The awns of the flowering glume are slender and brownish, those of the
lower flower of the spikelet being 8-9 cm. long, those of the upper flower 3-4
cm. long.

Grain, red, mealy, pointed at both ends, with prominent dorsal ridge ; 8-3
mm. long, 3 mm. broad, 3 mm. thick.

Ear branched, bearded ; awns long ; glumes blue-black, pubescent.

T. dicoccum, var. melanurum, Korn. Handb. d. Getr. i. 91 (1885).

T. amyleum, L., Metzger. Eur. Cer. 35 (1825) '•> Landw. Pfl. i. 118 bb. (1841).

T. vulgare melanura, Alef. Landw. Fl. 333 (1866).

The ear is slightly branched or bears only double spikelets at the notches on
the lower half of the rachis. Some of the ears are simple but produce branched
examples when their grain is sown. Metzger states that the variety is the
branched form of the Black Winter Emmer (var. atratum), but Kornicke found
that the simple ears differed somewhat from the latter.

Ear beardless ; glumes white, glabrous.

T. dicoccum, var. inerme, Korn. Arch. f. Biontologie, Bd. ii-4o8 (1908).

Discovered by Kornicke in the Botanic Garden at Poppelsdorf, and derived
from var. semicanum. It differs from typical T. dicoccum in having ears broader
across the face than across the 2-rowed side.

FIG. 133.— EMMER (T. dicoccum, Schiibl.).

var. rufum.

1 . Russian Red Emmer.

2. European Red Emmer.

var. atratum.
3. Black Emmer.

EMMER 203

Ear with double spikelets, beardless ; glume' white, glabrous.

T. dicoccum, var. bispiculatum, Korn. Arch./. Biontologie, ii. 410 (1908).
Derived from var. semicanum ; apparently of hybrid origin and discovered
by Kornicke in 1883 in the Botanic Garden, Poppelsdorf.

Ear branched, beardless ; glume white, glabrous.

T. dicoccum, var. leucocladum, Korn. Handb. d. Getr. i. 90 (1885).
T. vulgare leucocladus, Alef. Landw. Flora, 332 (1866).
Alefeld states that this variety originated from var. tricoccum and is not
constant.

Ear beardless ; glumes white, pubescent.

T. dicoccum, var. muticum, Korn. Handb. d. Getr. i. 86 (1885).
Formerly cultivated extensively in the Trentino (Kornicke).

Ear beardless ; glumes red, glabrous.

T. dicoccum, var. novicium, Korn. Arch.f. Biontologie, ii. 408 (1908).
Apparently a hybrid derived from var. cladurum. Discovered in the Botanic
Garden at Poppelsdorf.

Ear branched, beardless ; glumes red, glabrous.

T. dicoccum, var. subcladurum, Korn. Arch. f. Biontologie, ii. 410
(1908).

Apparently of hybrid origin and discovered by Kornicke in the Botanic
Garden at Poppelsdorf in 1892.

Ear beardless ; glumes red, pubescent.

T. dicoccum, var. hybridum, Korn. Arch.f. Biontologie, ii. 409 (1908).
A hybrid discovered by Kornicke in 1881 in the Botanic Garden at Poppels-
dorf.

CHAPTER XIV

KHORASAN WHEAT

Triticum orientale, mihi.

A SMALL but distinct race of wheat sent by Sir Percy Sykes from Khorasan,
Persia, where it is cultivated on irrigated land. At Reading it is among
the earliest wheats, coming into ear about the end of May.

In its narrow pubescent leaves, very lax ears, scabrid awns, very long
grain, and early habit it is clearly related to T. dicoccoides and T. dicoccum
and unlike T. durum.

In form of ear and length of grain it closely resembles some of the
varieties of T. polonicum figured by Seringe.

The plants at Reading are often cleistogamous and their fertility
much reduced, especially in damp seasons, when the anthers remain
minute and the filaments of the stamens do not lengthen.

GENERAL CHARACTERS OF T. orientale, mihi.

The coleoptile is 2-nerved.

The young shoots are erect, with very narrow pubescent leaves. The
plants tiller very little and the straw is thin, reed-like, short or of medium
height, 66-1 10 cm. (26-44 inches) long, the upper internode solid or hollow
with thick walls.

The ears are long, narrow, very lax, with a tough rachis and 10-11-5
cm. long, almost square, 10-11 mm. across the sides or 10-11 mm. across
the face and 8-9 mm. across the 2-rowed side ; density, 16-18.

The spikelets (16-20) are 2- to 3-grained, with more or less deciduous
awns, 12-13 cm. long, scabrid to the base. The sides of the rachis are
fringed with white hairs, and there is a conspicuous frontal tuft below
each spikelet ; the internodes are narrow and wedge-shaped, each about
i -5 mm. wide at the base and 3 mm. at the top.

The empty glumes are white, pubescent, 12-15 mm. long and 4 mm.
broad, keeled to the base, with a short apical tooth ; lateral nerve prominent
(Fig. 134).

The flowering glume bears a coarse awn, scabrid to the base, 14-16 cm.
long.

204

'

FIG. 136. — KHORASAN WHEAT (T. orientale, mihi).

i. var. notabile.
(Siah Das.)

2. var. insigne.
(Shutar Dandan.)

KHORASAN WHEAT

205

The grains are very long, narrow, white, and flinty, resembling those
of T.polonicum and the largest-grained forms of T. dicoccoides, being 10-5-
12 mm. long, 3 mm. broad, 3-2-3-4 mm. thick (Figs. 134, 135).

VARIETIES OF T. orientate, mihi.
Two varieties are found differing only in the colour of the awns.

1. Ear bearded ; glumes wrhite, pubescent ; awns white

2. Ear bearded ; glumes white, pubescent ; awns black

Ear bearded ; glumes white, pubescent ; awns white ;
T. orientale, var. insigne, mihi.

Shutar Dandan(= Camel's
tooth) (i, Fig. 136). — A
distinct very early variety,
with long lax ears, and long
narrow glumes and grains.
It comes into ear at Read-
ing during the last week of
May.

Young shoots, erect ;
young leaves very narrow,
pubescent.

Straw, thin, reedy, of
medium height, 95-110 cm.
(38-44 inches long) ; upper
internode solid or hollow
with thick walls.

Ear, long, very lax, nar-
row ; 10-11-5 cm. long,
almost square, 10-11 mm.
across the sides or 10-11

mm. across the face and 8-9 mm. across the side ;
spikelets 16-20, 2- to 3 -grained ; awns 13-17 cm.
long, deciduous, scab rid to the base ; D = 16-18.

Empty glume, white, pubescent, long and narrow ;
keeled to the base, 12-15 mm. long, 4 mm. wide ;
apical tooth short (Fig. 134).

Grain, very long, white, flinty, 10-5-12 mm. long,
2-75-3-0 mm. broad, 3-2-3-4 mm. thick.

var. tnsigne.
var. notabile.

grain white.

FIG. 134. — Empty glume
( x 2) and grain (front,
back, and side views,
nat. size) of Khora-
san wheat (T. orien-
tale).

FIG. 135. — Grains of the
spikelets of one side of an
ear of Khorasan wheat
(T. orientale) (nat. size).

Ear bearded ; glumes white, pubescent ;
awns black ; grain white.

T. orientale, var. notabile, mihi.

Siah Das ( = Black Hand) (2, Fig. 136). — This variety was sent by Sir Percy
Sykes from Khorasan with the preceding variety, from which it differs only in
the possession of jet black awns.

CHAPTER XV

MACARONI WHEAT

Triticum durum, Desf. Fl. Atlant. i. 114 (1798).

T. sativum, y, durum, Pers. Syn. PL i. 109 (1805).

T. vulgare durum, Alef. Landw. Fl. 324 (1866).

T. sativum durum, Hackel. Nat. Pfl. ii. 2, 85 (1887).

T. tenax, B. II., durum, Asch. und Graeb. Syn. ii. 692 (1901).

T. alatum, Peterm. in Flora, xxvii. 234 (1844) is often given as a synonym
of T. durum, although Petermann specifically states that it is different from T.
durum ; it has some characters of T. turgidum, and its place cannot be deter-
mined at present.

MACARONI Wheat is frequently named Hard Wheat on account of the
hard flinty nature of its grain ; as the term " hard " is, however, often
applied in the cereal markets to samples of North American Bread Wheats
(T. vulgare), it is advisable to use the name Macaroni Wheat for this race.

No authenticated specimens of ears or grain of T. durum of prehistoric
age are known, and there are no trustworthy data regarding its cultivation
in early historic times.

Grains discovered by Schliemann in a " pithos " when excavating
the walls of Hissarlik have been referred to T. durum by Wittmack and
Ko'rnicke. Seringe states that grains of this race have been obtained
from the wrappings of Egyptian mummies ; these were probably modern
and accidentally or fraudulently introduced.

In excavations of the cemetery at Hawara in Egypt Professor Flinders
Petrie found a number of well-preserved ears and grains of wheat in a
grave of the Graeco-Roman period dating from about the first century
B.C. Examples of these in the British Museum (Nat. Hist. Dept.) closely
resemble forms of T. durum, some of them with glabrous, others with
pubescent glumes, strongly keeled to the base ; the grains are small and
plump, with pointed base, narrowed apex, and high dorsal ridge, the
average dimensions being 7-1 mm. long, 3-1 mm. broad, and 3-25 mm.
thick.

Ears with keeled pubescent glumes and grains resembling those of

206

MACARONI WHEAT 207

T. durum were found at Kahun in Egypt in tombs of the Xllth Dynasty

(2OOO B.C.).

The earliest historical reference to this wheat is made by Dodoens in
his Historia frumentorum, published in 1566. Some grains found by him
among the grains of Canary grass (Phalaris canariensis) obtained from
Spain and the Canary Isles were sown, and gave rise to a fbrm of wheat
which he described and figured under the name Triticum Typhinum,
apparently connecting it with the ri^t] (tiphe) of ancient Greek authors.
The latter is now regarded as Triticum monococcum, but the account and
figure given by Dodoens leave no doubt that his plant was a form of
Macaroni Wheat.

Not until near the end of the eighteenth century was T. durum dis-
tinguished from the Mediterranean forms of T. turgidum and T. vulgare
by Desfontaines, who described it from Barbery in his Flora Atlantica
(vol. i. p. 114) in 1798, mentioning solid straw, pubescent glumes, and
long flinty grain as its specific characters.

Next to T. vulgare the various forms of T. durum are the most widely
cultivated wheats of the present day. They are grown throughout the
Mediterranean region, in Portugal, Spain, Italy, Morocco, Algeria,
Egypt, Abyssinia, Greece, Bulgaria, Turkey, and Asia Minor.

The greatest amount is produced on the " Tchernozem " or black
soil of Russia, especially in the region of the Volga. It is a common
cereal in Turkestan, the Transcaucasus, and the southern portion of
East Siberia, and is found in lesser amount in Bokhara, Persia, and India,
but it does not extend to China or Japan.

In the Western Hemisphere it is cultivated in the United States,
Canada, Mexico, the countries of Central America, Chili, and Argentina.

Small quantities are grown also in South Africa and Australia.

T. durum does not tiller much, but grows rapidly and succeeds best as
a spring crop, although at Reading it ripens later than many common
forms of T. vulgare. I have not met with any very early forms.

In regions possessing mild winters it may be sown in autumn, and
in such localities often provides useful green herbage, which may be fed
off lightly with stock during winter without damaging the yield of grain
at harvest.

Macaroni Wheat requires a dry hot climate for satisfactory growth,
the plants being easily damaged by frost. It possesses great power of
resisting drought, giving fair yields of grain in regions where the rainfall
is not more than 10-18 inches per annum. In districts too dry to allow
of the cultivation of T. vulgare, T. durum will yield 10-20 bushels per
acre, and crops of it have been raised in several semi-arid parts of the
world where no rain has fallen between seed-time and harvest. Its growth
is most prosperous, however, where a considerable proportion of the annual

208 THE WHEAT PLANT

rainfall occurs during the early part of the growing season, a few good
showers alternating with bright sunshine and a dry atmosphere resulting
in the highest yield of grain of fine quality. In addition to a suitable
climate, a deep rich soil containing an abundance of humus and an
adequate supply of lime, potash, and phosphates is necessary to secure
the best returns.

The Macaroni Wheats are highly resistant to rust and smut fungi.

In Australia, where wheat is frequently utilised as a hay crop, some
forms of T. durum have proved of especial value for this purpose. The
Macaroni Wheats, however, are grown chiefly for their grain, the best
samples of which are yellowish, translucent, and very hard and difficult
to grind. In modern mills the grains undergo a process of " conditioning "
before being ground.

In Russia and some other parts of the world a large amount of durum
is used for bread-making, either by itself or mixed with 10-20 per cent
of flour of Red-grained Bread Wheat (T. vulgar e). The bread made from
Macaroni Wheat is yellowish in colour, but of excellent flavour, and keeps
fresh longer than that from Bread Wheat.

The high gluten- content and the physical quality of the protein of
these wheats make them specially suited to the manufacture of macaroni,
spaghetti, and other edible pastes, and large quantities are employed for
this purpose, especially in France, Italy, and Spain. For the preparation
of these pastes the grain is milled only as far as the " semolina " stage, a
finely grained flour not being required. The dough formed from the
semolina is subsequently forced through tubes or openings of various
forms and sizes and then dried.

GENERAL CHARACTERS OF T, durum

The young leaf-blades and shoots have almost always the erect
" spring " habit (Fig. 65) ; I have seen none with the low spreading
" winter " habit, and a few only in which the young leaf-blades are semi-
erect, making an angle of 45°-6o° with the soil surface in the early stages
of their growth.

The leaves, which may be yellowish-green or bluish-green, are all
glabrous in most varieties. In a small number of forms, which are
probably hybrids, short hairs are present, especially on the surfaces of
the lower leaves, the character of the hairs in some caseg suggesting an
affinity with T. vulgar e, in others a relationship with T. dicoccum or T.
turgidum (Fig. 137).

The culms are striate, from 75 to 150 cm. or more (about 30-60 inches)
high, the surface somewhat dull and slightly rough ; in some cases they
are hollow with thick walls, but in most varieties they are solid throughout

MACARONI WHEAT

209

or in the upper internodes. When well developed they possess 5 or 6
internodes above ground.

Except in the case of one or two varieties, which appear to be hybrids,
the ears are always bearded, and either square
in section or compressed laterally and oblong
in section, in which latter case the 2-ranked
side is generally wider than the face of the ear.
They are from 4 to n cm. (about i| to 4^
inches) in length without the awns, having an

FIG. 137. — Diagrammatic
average of 21 spikelets per ear, the density vary- transverse sections of

ing from about 20 to 47 per 10 cm. length of young leaves of T. durum

0 . and 1 . polonicum .

rachis.

The rachis is usually tough, though in some varieties it disarticulates
more or less easily, especially near the base of the ear ; it is fringed

8 9 10 ii

12 13 14 15 l6 17 l8 IQ 20

FIG. 138. — Empty glumes of Macaroni wheat (T. durum) ( x 2).

210

THE WHEAT PLANT

along the margins with
hairs, and bears a frontal
tuft at the base of each
spikelet.

The spikelets are 5- to
y-flowered, those of square
ears ripening three or four
grains, those of com-
pressed ears usually only
two ; they are from 10 to 15
mm. long, 8- 1 5 mm. broad,
and 4-5 mm. thick.

The empty glumes are

FIG. 139. — Grains of Macaroni wheat (T. durum),
front, back, and side views (nat. size).

glabrous or hairy, the length and amount of
the hair varying considerably. The colour
is yellow, red, or blue - black in various
shades. Those of the lateral spikelets are
8-12 mm. long, unsymmetrical, the outer face
somewhat flat and measuring from the midrib
to the outer edge 4-5 mm., the inner portion
being 2-3 mm. wide. A prominent keel runs
from the base to the tip. The apical tooth,
which may be acute or blunt, is of variable
length ; the lateral secondary tooth in which
the strong nerve on the outer face terminates
being usually short or missing : the various
forms of empty glumes are illustrated in Fig.

138-

The flowering glumes are thin and pale,
rounded on the back with 9-15 nerves, the
two lowest of each spikelet having long awns,
measuring in some cases as much as 20-23
cm., those of the upper flowers of the
spikelet usually having a length of -5 to 4 cm.
long.

The awns are white, red, or black, the
latter tint, as in T. turgidum and other wheats
(see p. 105), being developed to the greatest
extent in dry bright weather, frequently failing
to appear in dull humid seasons ; they are

FIG. 140. — Grains of the spike-
lets on one side of an ear of
Macaroni wheat (T. durum)
(nat. size).

MACARONI WHEAT

211

almost smooth near the base, thus differing from those of T. turgidum,
which are usually scabrid throughout their whole length.

The awns are generally straight and more or less parallel to the sides
of the ear, though in some varieties they are divergent and occasionally
bent in a sinuous manner near the base ; they are longer than those of
any other wheats, varying from n to 23 cm. in length, and are more per-
sistent than those of T. turgidum, except in some Indian forms, which
lose their awns very readily when dead ripe.

The grains are usually white, amber, yellow, or red ; the latter, when
pale and flinty, are extremely difficult to distinguish from white or amber
flinty grains.

The typical form is somewhat narrow, tapering towards both ends,
more or less laterally compressed, with a narrow dorsal ridge, and wanting
in plumpness, the shallow furrow usually having flattish sloping sides ;
the cross section is more or less triangular (Figs. 139, 140).

The endosperm is generally very hard and translucent with a vitreous
or flinty fracture ; a few varieties approximating towards T. dicoccum
or T. turgidum have opaque mealy grains.

The embryo is large with an elongated oval scutellum. Measurements
of grains taken from the middle of the ear of flinty forms gave the following
results :

Length.

Breadth.

Thickness.

mm.

mm.

mm.

Average
Limits
Ratio ....

8-30
7-0-9-7
IOO

3H8
2-8-4-1
41-9

3-61
3-2-4-25
43'5

VARIETIES OF T. durum, Desf.

(i) Ears bearded-

Glumes white, glabrous ; awns white.

a. Grain white ....

b. Grain red .....
Glumes white, glabrous ; awns black.

a. Grain white ....

b. Grain red .....
Glumes white, pubescent ; awns white.

a. Grain white ....

b. Grain red .....
Glumes white, pubescent ; awns black.

a. Grain white ....

b. Grain red .

. var. leucurum, Korn.
var. affine, Korn.

var. leucomelan, Korn.
var. Reichenbachii, Korn.

. var. Valenciae, Korn.
. var.fastuosum, Korn.

var. melanopus, Korn.
var. africanum, Korn.

212

THE WHEAT PLANT

5. Glumes red, glabrous ; awns red.

a. Grain white

b. Grain red ....

6. Glumes red, glabrous ; awns black.

a. Grain white .....

b. Grain red ......

7. Glumes red, pubescent ; awns red.

a. Grain white . . . . .

b. Grain red ......

8. Glumes red, pubescent ; awns black.

a. Grain white .....

b. Grain red ......

9. Glumes blue-black, glabrous.

a. Grain white .....

b. Grain red ......

10. Glumes blue-black, pubescent.

a. Grain white . . . . .

b. Grain red ......

(2) Ears beardless —

1 . Glumes blue-black, glabrous ; grain white .

2. Glumes red, glabrous ; grain white .

var. hordeiforme, Korn.
. var. mur dense, Korn.

var. erythromelan, Korn.
var. alexandrinum, Korn.

var. italicum, Korn.
var. aegyptiacum, Korn.

. var. apulicum, Korn.
. .var. niloticum, Korn.

var. provincial, Korn.
. var. obscurum, Korn.

var. coerulescens, Korn.
var. libycum, Korn.

var. australe, mihi.
var. sub-australe, mihi.

Ear bearded ; glumes white, glabrous ; awns white ; grains white.

T. durum, var. leucurum, Korn. Handb. d. Getr. i. 69 (1885).

One of the most widely cultivated varieties. It is extensively grown in
Italy, Spain, and Portugal, the majority of the forms of these countries having
long ears (8-10 cm.), which are comparatively lax (D = 24-28) and almost square
in section.

It is also found in considerable amount in Asia Minor, but the forms met
with in this region have shorter and denser compressed ears, 5-8 cm. long ;
0 = 34-40 ; oblong in section.

I received lax-eared forms from the Punjab, with ears 9-10 cm. long ; D = 21-
23. Some of them are not far removed from the Indian forms of T. dicoccum.

1. Punjab 3.

Ear, lax, about 10 cm. long, almost square in section, 10-11 mm. across the
face and side ; awns somewhat slender, 12-14 cm. long ; spikelets 21-23 ;
D = 21-23 (Ear types i and 2, Fig. 142).

Empty glume, n mm. long, apex blunt, apical tooth short, blunt, secondary
nerve distinct, and ending a short distance from the base of the apical tooth (3,
Fig. 138).

Grain, flinty, long, and narrow, 9-4 mm. long, 2-6-3 mm. broad, 2-6-3 mm.
thick.

2. Trigo fanfarron. — Received from Spain.
Young shoots, erect ; young leaves glabrous.

Straw, tall, 125 cm. (about 49 inches) long ; upper internode solid.

FIG. 141. — MACARONI WHEAT (T. durum, Desf.).

i . var. leucurum.
(Trigo de espiga larga blanca.)

2. var. leucurum.
(Trigo blanquillo.)

MACARONI WHEAT 213

Ear, lax, large, 9*5-10 cm. long, square, 11-12 mm. across the sides ; spike-
lets 22-24, 3~ to 4-grained ; D = 25 ; awns 10-19 cm. long (Ear type i, Fig. 141).

Empty glume, 10 mm. long ; apical tooth broad, acute, i mm. long (7, Fig. 138).

Grain, flinty, 8-5 mm. long, 3-7 mm. broad, 3-4 mm thick.

Closely similar to this are Trigo de espiga larga blanca, Trigo blanquillo
de la Granja de Jeres, and Trigo bianco Verdeal from Spain, and a large form
from Abyssinia.

3. Saragollo. — Received from Spain and also from Italy ; recorded from
Sicily by Werner.

Young shoots, erect ; young leaves glabrous or slightly rough.

Straw, tall, 124 cm. (about 49 inches) long ; upper internode solid.

Ear, narrow, 9-10 cm. long, square, 10-11 mm. across the sides ; spikelets
23, 2- to 3-grained ; D = 27-29 ; awns 11-13 cm. long (Ear type 2, Fig. 141).

Empty glume, 9-10 mm. long; apical tooth broad and blunt, -5 mm. long
(n, Fig. 138).

Grain, large, flinty, compressed, with prominent dorsal ridge, 9-9-5 mm.
long, 3-6 mm. broad, 4 mm. thick.

4. Trigo Claro. — Received from Spain.
Young shoots, erect ; young leaves glabrous.

Straw, medium to tall, 1 14 cm. (about 45 inches) long ; upper internode solid.

Ear, 7-8 cm. long, 10-11 mm. across the face, 13 mm. across the side ;
spikelets 22, 2- to 3-grained ; D = 28-32 ; awns 17 cm. long (Ear type 2, Fig. 143).

Empty glume, 10 mm. long ; apical tooth narrow, acute, 1-2 mm. long (8,
Fig. 138).

Grain, flinty, 8-6 mm. long, 3-25 broad, 3-55 mm. thick.

Very similar to this is Duro Catanzaro, from Italy with narrower and
slightly shorter ears.

5. Forms of var. leucurum received from Asia Minor are Ak poussan, Arsus,
and Alexandretta wheat, which closely resemble each other.

Ear, moderately lax, 8-10 cm. long, compressed 8-10 mm. across the face,
13-15 mm. across the side ; spikelets about 23 ; awns 12-14 cm. long ; 0 = 25-
30 (Ear type 2, Fig. 144).

Empty glume, about 10 mm. long, with stout, acute, apical tooth, i mm. long.

Grain, flinty, long, and narrow, 8-5 mm. long, 3 mm. broad, 3 mm. thick.

6. A common form also from Asia Minor, under the names Menenem
Boydesi, Tsirali-ak-Sowsam, and Magnesia wheat, has the following characters :

Ear, dense, compressed, 7-8 cm. long, 9-10 mm. across the face, 14-16 mm.
across the side ; spikelets 21-23 ; awns 12-14 cm. long ; 0 = 30-36 (Ear type
2, Fig. 144).

Empty glume, 10 mm. long, 5 mm. wide, apical tooth short, acute (4, 9. Fig.

138)-

Grain, semi-flinty, apex narrowed, dorsal ridge prominent ; 8-8-5 mm-
long, 3-5 mm. broad, 3-5 mm. thick.

7. Short dense-eared forms are also frequent in Asia Minor (Ear type i;
Fig- HS)-

214 THE WHEAT PLANT

Ear, 5-6 cm. long, 8-10 mm. across the face, 13-15 mm. across the side;
awns 7-9 cm. long ; spikelets 21-23 ; D = about 40.

Empty glitme, 10 mm. long, apical tooth acute.

Grain, flinty, dorsal ridge prominent ; 7-5 mm. long, 3-5 mm. broad, 3-3
mm . thick.

8. Derived from an Austrian ear of T. vulgare, grown at Reading, is a dense,
short-eared form. (? a hybrid with T. monococcum grown in an adjacent row.)

Young shoots, erect ; young leaves glabrous.

Straw, tall, 127 cm. (about 50 inches) long; upper internode solid.

Ear, short, very dense with narrow face ; 5-7 cm. long, 8-10 mm. across the
face ; 0=43-47 ; awns 13 cm. long.

Empty glume, 9 mm. long ; apical tooth narrow, acute, 1-5-2 mm. long.

Grain, flinty, much compressed laterally, 1-7 mm. long, 2-75-2-8 mm. broad,
3-5 mm. thick.

Ear bearded ; glumes white, glabrous ; awns white ; grain red.

T. durum, var. affine, Korn. Handb. d. Getr. i. 70 (1885).

A widely distributed variety cultivated in Spain, Portugal, Italy, and North
Africa. Found also in small amounts in European Turkey, Russia (the Don
Provinces), Transcaucasia, Turkestan, and Asia Minor. I have also had a form
of it from Burma.

The majority of the forms have somewhat slender, lax ears about 9 cm. long,
and almost square in section ; a few possess shorter, denser, compressed ears.

1 . The form from Burma is not far removed from forms of T. dicoccum
and has the following characters :

Ear, narrow, lax, about 8 cm. long, almost square in section, 9-10 mm.
across the face and side ; spikelets 19-21 ; 0 = 23-26 ; awns 15 cm. long (Ear
type 2, Fig. 142).

Empty glume, n mm. long, narrow, with acute apical tooth (n, Fig. 138 ;
2, Fig. 142).

Grain, semi-flinty, small, narrow, about 8 mm. long, 3 mm. wide, 3 mm.
thick.

Yiaf boyde from Asia Minor is similar.

2. Trimenia. — An ancient and widely distributed form, cultivated in Spain,
Portugal, Italy, and North Africa. A form received from Italy under the name
Risciola is similar, as is also Trigo Obispado from Spain.

Young shoots, erect ; young leaves glabrous.

Straw, short, 86 cm. (about 34 inches) long ; upper internode hollow or solid.

Ear, lax, somewhat slender, 9-10 cm. long, square, 10-12 mm. across the
side; spikelets 22-24, 2~ to 3-grained; 0=24-26; awns 13-16 cm. long (Ear type
2, Fig. 141).

Empty glume, 9-10 mm. long ; apical tooth narrow, acute, 1-5-2 mm. long
(6, 9, Fig. 138).

MACARONI WHEAT 215

Grain, somewhat short, flinty, dorsal ridge prominent; 7-5-8 mm. long,
3-4 mm. broad, 3-6 mm. thick.

Similar but with longer grain is De Alfaro from Spain.

3. Amarello de barba branca. — Received from Portugal.
Young shoots, semi-erect ; young leaves glabrous.

Straw, slender, of medium height, no cm. (about 44 inches) long ; upper
internode solid.

Ear, 8-10 cm. long, 10-11 mm. across the face, 12-13 mm. across the side ;
spikelets 21-25, 2- to 3 -grained ; D = 28-30 ; awns 15-16 cm. long (Ear type i,
Fig. 141).

Empty glume, 11-12 mm. long ; apical tooth acute, 1-1-5 mm- l°ng (4> 9>
Fig. 138).

Grain, flinty or half mealy, 8-6-9 mm. long, 3-2-3-6 mm. broad, 3-3-3-6 mm.
thick.

4. A short, dense-eared form received from Germany, under the name
Sorrentino ; it appears to be near to Kornicke's var. campylodon (a form of
affine with incurved tooth on the empty glume).

Young shoots, erect ; young leaves glabrous.

Straw, of medium height, 100 cm. (about 40 inches) high ; upper internode
solid.

Ear, 6-7 cm. long, square, 12-15 mm. across the sides; spikelets 22-24, 3-
to 4-grained ; D = 33-35 ; awns 15 cm. long (Ear type i, Fig. 144).

Empty glume, 10 mm. long ; apical tooth short, curved inwards, secondary
tooth distinct (4, Fig. 138).

Grain, flinty, 7-9 mm. long, 3-4 mm. broad, 3-7 mm. thick.

5. Lobeiro. — Received from Portugal and an allied form from Greece.
Young shoots, semi-erect ; young leaves glabrous.

Straw, of medium height, about 96 cm. (about 38 inches) long ; upper inter-
node solid.

Ear, dense, compressed, tapering somewhat towards the apex; 8 cm. long,
10-11 mm. across the face, 13-15 mm. across the side; spikelets 20-22,2- to
3-grained ; 0 = 33-35 ; awns 12-13 cm. long (Ear type i, Fig. 143).

Empty glume, 10-11 mm. long ; apical tooth narrow, acute, 2 mm. long (7,
Fig. 138).

Grain, flinty, large, apex truncate, and a somewhat narrow dorsal ridge ; 8-6
mm. long, 3-65 mm. broad, 3-55 mm. thick.

Ear bearded ; glumes white, glabrous ; awns black ; grain white.

T. durum, var. leucomelan, Korn. Handb. d. Getr. i. 70 (1885).

A common Mediterranean variety, especially frequent in Spain and Italy,
rare in Russia. Most forms have slightly rough young leaves ; they are probably
of hybrid origin and related to T. turgidum or T. vulgare.

The majority possess somewhat short, solid straw ; stout, dense, short ears,
almost square in section, 7-8 cm. long, 12-15 mm. across the face and side.

2i 6 THE WHEAT PLANT

The awns are brown or black, and a dark line is usually present along the
upper part of the keel and edges of the empty glumes. The latter are large,
generally with long, acute, apical teeth.

1. Trigo Macolo. — Received from Spain.
Young shoots, erect ; young leaves slightly rough.

Straw, of medium height, 100 cm. (about 39 inches) long ; upper inter-
node solid.

Ear, 8-5-10 cm. long, square, 12-13 mm. across the side ; spikelets 22-24,
2- to 3-grained ; D =29-32 ; awns 17-18 cm. long (Ear type 2, Fig. 143).

Empty glume, 10-11 mm. long ; upper part of apical tooth usually black ;
apical tooth acute, broad at the base, 1-5-2 mm. long (6, Fig. 138).

Grain, mealy, with truncate apex ; 8 mm. long, 3-85 mm. broad, 3-65 mm.
thick.

Similar are Trigo de espiga azulada and Trigo Alonso from Spain ; in
these the empty glume has a narrow, longer, apical tooth.

2. Trigo Chacon. — Received from Spain.
Young shoots, erect ; young leaves glabrous.

Straw, of medium height, 110-115 cm- (about 40-42 inches) high ; upper
internode solid.

Ear, coarse, stout, short, and dense, 6-8 cm. long, square, 15 mm. across the
sides ; spikelets broad, 20-22, 3- to 4-grained ; D = 30-32 ; awns stout, 13-14 cm.
long (Ear type i, Fig. 143).

Empty glume, 11-12 mm. .long ; apical tooth acute, 2 mm. long (4, Fig. 138).

Grain, large, semi-flinty, 8-8-9 mm- l°ng> 3'8~3'9 mm. broad, 4 mm. thick.

Very similar is Trigo Cannu altu from Spain and Italy.

3. T. raspinegro de espiga peciuefia. — Received from Spain.
Young shoots, semi-erect ; young leaves with few short hairs.

Straw, of medium height, 102 cm. (about 40 inches) long ; upper inter-
node solid.

Ear, short, dense, often convex on one side ; 6-6-5 cm- l°n§> IO mm- across
the face, 17 mm. across the side ; spikelets 23-24, 2- to 3-grained ; 0=44 ;
awns 13-14 cm. long (Ear type i, Fig. 145).

Empty glume, 10 mm. long ; apical tooth narrow, acute, 2 mm. long (6,
Fig. 138).

Grain, flinty, 8-7 mm. long, 3-85 mm. broad, 3-6 mm. thick.

Ear bearded ; glumes white, glabrous ; awns black ; grain red.

T. durum, var. Reichenbachii, Korn. Handb. d. Getr. i. 70 (1885).

First obtained by Kornicke from the Dresden Botanic Garden as " Triticum
meianocus " (? melanocus), and later from the Province of Cadiz in Spain.

A somewhat uncommon variety received from Spain, Greece, and Asia
Minor ; Flaksberger mentions its rare occurrence in Russia (the Don Pro-
vinces and Caucasus) and in Turkestan.

i. Trigo raspinegro. — Received from Spain.

Young shoots, erect ; young leaves slightly rough.

FIG. 142. — MACARONI WHEAT (T. durum, Desf.)>

i . var. Reichenbachii.
(Greece.)

2. var. affine.
(Burma.)

MACARONI WHEAT 217

Straw, very tall, 127 cm. (about 59 inches-) long ; upper internode solid.

Ear, 8-9 cm. long, coarse, square, 11-12 mm. across the sides, or com-
pressed, and 10 mm. across the face, 12-13 mm. across the side ; spikelets 20-24,
2- to 3-grained ; D =25-28 ; awns stout, 17-18 cm. long, sometimes irregularly
bent (Ear type 2, Fig. 143).

Empty glume, 10-12 mm. long, narrow ; apical tooth small, about i mm.
long (4, Fig. 138).

Grain, flinty, long, and narrow, apex truncate ; 8-9 mm. long, 3-3 mm.
broad, 3-85 mm. thick.

Grano forte from Italy resembles this form.

A larger form with square ears, 12-13 mm. across the sides and paler awns,
also from Spain.

2. Trigo marzuolo. — Received from Italy, and similar forms from Greece.

Young shoots, erect ; young leaves glabrous.

Straw, tall, 116 cm. (about 46 inches) long ; upper internode hollow.

Ear, long and narrow, 8-10 cm. long, square, 10-11 mm. across the sides ;
spikelets 23, 2- to 3-grained ; D = 23-26 ; awns 14 cm. long (Ear type i, Fig. 142).

Empty glume, narrow, n mm. long ; apical tooth 1-2 mm. long, acute,
usually with a black margin (9, Fig. 138).

Grain, flinty, with truncate apex and narrow dorsal ridge ; 8 mm. long, 3-45
mm. broad, and 3-6 mm. thick.

I received a smaller form allied to this from Adana, Asia Minor, under the
name Yerli.

Ear bearded ; glumes white, pubescent ; awns white ; grain white.

T. durum, var. Valenciae, Korn. Handb. d. Getr. i. 72 (1885).

Kornicke received this variety from Valencia, Spain, the Government of
Kharkov, Russia, and Chili.

I have received it from Austria, Bulgaria, Asia Minor (Damascus), and the
Punjab, India ; also in samples of Goose wheat from Canada.

Forms of this variety are rare and liable to be confused with examples of the
black-awned var. melanopus, in which the dark pigment does not develop in
some seasons.

Flaksberger says that the forms from Russia should be placed under the
variety melanopus.

i. Trigo Sicilian!. — Received from Italy.

Young shoots, semi-erect ; young leaves glabrous.

Straw, tall, 116 cm. (about 46 inches) high ; upper internode solid.

Ear, 9 cm. long, square, 10-12 mm. across the sides, or compressed, and
10-11 across the face, 12-14 mm. across the side; spikelets 21-23, 2- to 3-
grained ; 0 = 28-30 ; awns 13 cm. long (Ear type i, Fig. 142).

Empty glume, 9-11 mm. long, 4 mm. broad ; apical tooth 1-1-5 mm- long
(12, 13, Fig. 138).

Grain, flinty, apex narrowed ; 8-8-5 mm- l°ng> S'^S'S rnrn. broad. Similar
to this is Beloklassa zagani from Bulgaria.

2i 8 THE WHEAT PLANT

2. Cawnpore 2.

Ear, large, 10 mm. long, compressed 10 mm. across the face, 13-14 mm.
across the side ; spikelets 22-24 '> D =23-26 (Ear type i, Fig. 141).

Empty glume, large, 12 mm. long, 5 mm. broad, apex broad, apical tooth
short and blunt (2, Fig. 138).

Grain, long and narrow, 8 mm. long, 3 mm. wide, 3 mm. thick.

3. Cawnpore 9. — Somewhat similar to the previous form but with smaller,
denser ears.

Ear, about 9 mm. long, 10 mm. square, or 9 mm. across the face, and n mm.
across the side ; spikelets 21-23 ; D = 28 (Ear type 2, Fig. 141).

4. A form of var. Valenciae received from Damascus, Asia Minor, under the
name Yabroudi Bayadi, has the following characters :

Ear, compressed, 8 cm. long, 6-8 mm. across the face, 11-13 mm- across the
side; spikelets 22-27 ; D = 30-36.

Empty glume, 8-5 mm. long, 3 mm. broad, apical tooth acute, short (i, n,
Fig. 138).

Grain, 7-5 mm. long, 3-3-4 mm. broad, 2-5-3 mm. thick.

Ear bearded ; glumes white, pubescent ; awns white ; grain red.

T. durum, var. fastuosum, Korn. Handb. d. Getr. i. 72 (1885).

A rare variety received by Kornicke from the Government of Kharkov,
Russia, and from an Exhibition in Vienna in 1873. Flaksberger states that he
has not seen a single ear of this variety from any part of the Russian Empire.

I have received specimens only from India (Central and United Provinces).
These were narrow lax-eared forms with the following characters :

Young shoots, erect ; young leaves glabrous.

Straw, of medium height, 104 cm. (about 41 inches) long ; upper internode
almost solid.

Ear, small, narrow, 7-9-5 mm. long, square, 10-11 mm. across the sides or
slightly compressed, 8-9 mm. across the face, and 11-12 mm. across the side ;
spikelets 21-23, 3-grained ; D =25-27 ; awns slender, 10-13 cm. long (Ear types
i, 2, Fig. 142).

Empty glume, 9 mm. long, apical tooth short and blunt, -5 mm. long (10,
Fig. 138).

Grain, flinty, apex narrow, 7-5-8-5 mm. long, 3-2 mm. broad, 3-5 mm thick.

Ear bearded ; glumes white, pubescent ; awns black ; grain white.

T. durum, var. melanopus, Korn. Handb. d. Getr. i. 72 (1885).

One of the commonest varieties of T. durum with a wide distribution, being
cultivated in Spain, Portugal, Italy, Roumania, South-East Russia, Turkestan,
Siberia, Asia Minor, and India.

Forms of this variety are found with long and somewhat narrow, square
ears 10 mm. across ; the majority, however, have short, broad ears, 14-15 mm.
square, strong coarse awns, and slightly scabrid young leaves.

MACARONI WHEAT 219

The forms from Asia Minor generally possess short, dense, compressed
ears and stout awns. The ears are usually from 5-5 to 7-5 cm. long, 10-12 mm.
across the face, and 14-15 mm. across the side ; D= usually about 33, but the
density of the shortest ears sometimes reaches 40.

Narrow-eared forms received from Russia and Dobrudja, Roumania, with
pale brown awns, have the following characters :

Young shoots, erect ; young leaves glabrous or slightly hairy.

Straw, slender, tall, 118 cm. (about 47 inches) long ; upper internode hollow.

Ear, slender, 9 cm. long, square, 10-11 mm. across the sides ; spikelets 25-27,
2- to 3-grained ; D = 31-32 ; awns slender, 12-13 cm. long (Ear type i, Fig. 142).

Empty glume, short, and somewhat inflated, 8 mm. long ; apical tooth short,
broad at the base, 1-1-5 mm- l°ng (jS, 19, Fig. 138).

Grain, flinty, small, 7 mm. long, 3-55 mm. broad, 3-2 mm. thick.

1. Punjab 1.

Young shoots, erect ; young leaves glabrous.

Straw, tall, to very tall, 132 cm. (about 53 inches) high ; upper internode hollow.

Ear, 9-10 cm. long, almost square, 11-12 mm. across the sides ; spikelets
22-26, 2- to 3-grained; D =27-28; awns stout, 12-16 cm. long, jet black or
sometimes pale brown (Ear types i, Fig. 141 ; 2, Fig. 143).

Empty glume, 10 mm. long ; apical tooth, short and bluntish, about i mm.
long (i, 2, Fig. 138).

Grain, small, flinty, apex blunt, 7-3-7-5 mm. long, 3-4 mm. broad, 3-1 mm.
thick.

I received a similar form from Bombay, the Shan States, Northern Burma,
and other parts of India ; also from Asia Minor.

2. Durazio mollar. — Received from Portugal.

Young shoots, semi-erect ; young leaves slightly rough.

Str aw, some what slender, tall, 1 16 cm. (46 inches) high ; upper internode solid.

Ear, short, 6-8 cm. long, square, 10-12 mm. across the sides or slightly com-
pressed, 10 mm. across the face, and 11-12 across the side; spikelets 21, 3-
grained ; 0 = 33 ; awns 14-15 cm. long, sometimes pale brown (Ear type 2,

Fig- H3)-

Empty glume, 10 mm. long ; apical tooth narrow, acute, 1-5-2 mm. long
(Form 6, Fig. 138).

Grain, semi-flinty, compressed, with high and narrow dorsal ridge; 8-25 mm.
long, 3-75 mm. broad, 3-6 mm. thick.

Zwaart Baard from South Africa resembles this.

Durazio rijo from Estramadura, Portugal, is also similar with somewhat
longer ears and larger, yellowish-white grain.

3. Trigo Andaluze. — Received from Spain and a similar form from Algeria.
Young shoots, erect ; young leaves glabrous.

Straw, of medium height, 102 cm. (about 40 inches) long; upper internode solid.

Ear, stout and coarse, 8-9 mm. long, square, 13-15 mm. across the sides ;
spikelets 20-22, 3- to 4-grained ; D = 28-32 ; awns strong, black, 15-16 cm. long
(Ear type 2, Fig. 143).

220 THE WHEAT PLANT

Empty glume, 10-11 mm. long ; apical tooth acute, 1-5-2 mm. long (Form 9,
Fig. 138).

Grain, flinty, long, narrow, 8-7-9 mm- l°ng> 4'° mm. broad, 4-0 mm. thick.
Goose from Canada resembles this, but has smaller, narrower ears.

4. Similar forms from Spain are Trigo Macolo, T. negro, with glabrous
leaves, and T. Enano de Larco, Trigo Eecio de Granada, T. Granadino and
others with slightly scabrid young leaves.

Forms with shorter and denser ears (7 cm. long, 15 mm. across the sides,
D = 33-36), and shorter grain, are also met with in Spain under various names.

Ear bearded ; glumes white, pubescent ; awns black ; grain red.

T. durum, var. africanum, Korn. Handb. d. Getr. i. 73 (1885).

Kornicke obtained this variety from Portugal, Egypt, and Asia Minor.

A somewhat rare variety received from Spain, Portugal, Russia, and India.

The Russian and Indian forms have long, somewhat narrow, square ears
(0 = 27-32), those from Spain and Portugal large, coarse, denser ears, showing
affinity with T. turgidum.

1. Alexandre. — Received from Portugal ; cultivated in Santa Cruz, Funchal.
Young shoots, erect ; young leaves glabrous.

Straw, of medium height, 109 cm. (about 43 inches) long ; upper inter-
node solid.

Ear, stout, 6-8 cm. long, square, 12-14 mm. across the sides ; spikelets 22-23,
3 -grained ; D = 31 ; awns 16-18 cm. long (Ear type i, Fig. 143).

Empty glume, 10-11 mm. ; apical tooth very long and narrow, 4-5 mm. long
(6, 7, Fig. 138).

Grain, flinty, large, 8-3-8-7 mm. long, 3-2-3-4 mm. broad, 3-3-4-1 mm. thick.

2. Russian Form.

Young shoots, semi-erect ; young leaves slightly rough.

Straw, somewhat slender, very tall, 140 cm. (about 55 inches) high ; upper
internode hollow.

Ear, 7-5-8 cm. long, square, 10-11 mm. across the sides; spikelets 22,
3-grained ; D = 32 ; awns 15 cm. long (Ear type 2, Fig. 143).

Empty glume ,() mm. long ; apical tooth short, broad, and blunt, i mm. long
(n, Fig. 138).

Grain, flinty, short, with narrow dorsal ridge ; apex truncate ; 7-1 mm. long,
3-6 mm. broad, 3-8 mm. thick.

3. Anafil. — Received from Portugal.

Young shoots, erect ; young leaves slightly rough.

Straw, slender, tall, 130 cm. (about 51 inches) long ; upper internode solid.
, Ear, dense, often broad at the base and tapering towards the apex, 8-9 cm.
long, 12 mm. across the face, 15-18 mm. across the side ; spikelets 27-29, 3-
to 4-grained ; D = 36 ; awns 14-15 cm. long (Ear type 2, Fig. 144).

Empty glume, 10 mm. long ; apical tooth acute, 2-5-3 mm. long (5, Fig. 138).

FIG. 143. — MACARONI WHEAT (T. durum, Desf.).

i. var. leucomelan.
(Alonso o Fanfarron.)

2. var. leucomelan.
(Trigo Macolo.)

MACARONI WHEAT 221

Grain, flinty, with prominent dorsal ridge, 8-5 mm. long, 3-3 mm. broad,
3-6 mm. thick.

4. Punjab 2.

Straw, of medium height, 77-97 cm. (36-38 inches) high.

Ear, long, lax, compressed, 8-10 cm. long, 8-10 mm. across the face, 12 mm.
across the side ; spikelets 22-24 > D =30-34 ; awns 14-15 cm. long (Ear types :
i, Fig. 141 ; 2, Fig. 143).

Empty glume, 8-9 mm. long, 4 mm. broad, with short, blunt, apical tooth
(Form 12, Fig. 138).

Grain, long, narrow, flinty, with prominent, dorsal ridge ; 8-3 mm. long, 3-3
mm. broad, 3-2 mm. thick.

In some seasons this form has pale awns and is then confused with var.
fastnosum.

Ear bearded ; glumes red, glabrous ; awns red ; grain -white.

T. durum, var. hordeiforme, Korn. Handb. d. Getr. i. 71 (1885).

One of the most widely distributed varieties of T. durum, especially common
in Russia, Bulgaria, Turkey, Greece, and Asia Minor. It occurs also in India,
Egypt, the North African Coast, Italy, Spain, and Portugal, but is rare in the
three latter countries.

The Macaroni Wheats usually grown in the United States belong to this
variety.

Four general types of ear are found, namely :

a. An elongated, narrow form about 10 cm. long, square in section, about
10 mm. across the sides ; awns 15-18 cm. long (2, Fig. 141).

b. A similar form with ears 7-8 cm. long ; awns 15-18 cm. long, and stout
(2, Fig. 142).

c. A shorter, denser form, 7-9 cm. long, square in section, 10-12 mm. across
the sides ; awns 13-15 cm. long (Figs. 143, 144).

d. A short, much compressed form, 6-8 cm. long, 8-10 mm. across the face,
and 15-20 mm. across the side ; awns 12-14 cm. long (2, Fig. 144).

1. Most of the " Kathias " of India are forms with narrow ears, stout, very
long awns (Ear type 2, Fig. 142), and narrow, strongly keeled, empty glumes
(n, Fig. 138, and 2, Fig. 142). The upper internodes are almost solid. They
are not far removed from the Indian T. dicoccum.

2. Several spring forms received from Spain and Portugal and showing
affinities with T. vulgare or T. turgidum have the following characters :

Young shoots, erect ; young leaves more or less hairy.

Straw, of medium height, 45-100 cm. (38-40 inches) high ; upper internode
solid or hollow with thick walls.

Ear, long, lax, and narrow, 10-12 cm. long, square, 10-11 mm. across the
sides ; spikelets 24-27, 3-grained ; 0 = 24-26 ; awns 12-14 cm. long (Ear type
2, Fig. 141).

Empty glume, short and somewhat inflated, 8 mm. long ; apical tooth -5-1
mm. long (n, Fig. 138).

222 THE WHEAT PLANT

Grain, semi-flinty, with high dorsal ridge ; 8-8-3 mm- l°ng> 3'3~3'6 mm.
broad, 3-6-3-85 mm. thick.

3. Kubanka.— This is the common form of var. hordeiforme grown in the
Kuban territory, South Russia, and now introduced into most countries where
Macaroni Wheats are cultivated ; most Russian Arnautka forms are similar to
this.

Young shoots, semi-erect or prostrate ; young leaves glabrous.

Straw, very tall, 140 cm. (about 55 inches) long ; upper internode hollow.

Ear, lax, 9-10 cm. long, square 10-11 mm. across the sides ; spikelets 24,
2- to 3 -grained ; 0 = 24-27 ; awns 13-15 cm. long (Ear type i, Fig. 142).

Empty glume, 9 mm. long ; apical tooth acute, 1-1-5 mm- l°ng (Jo, Fig. 138).

Grain, flinty, sometimes opaque, 7-5-8 mm. long, 3-6 mm. broad, 3-4-3-6
mm. thick.

4. Bieloturka ( = White Turkish). — Autumn or spring forms received from
Russia, Greece, Turkey, and Spain ; these closely resemble the forms named
Kubanka, but have shorter and somewhat denser ears.

Young shoots, semi-erect ; young leaves glabrous.

Straw, of medium height, 95-100 cm. (38-40 inches) high ; upper internode
hollow, with thick walls.

Ear, 6-8 cm. long, often tapered towards the apex, square, 10-12 mm. across
the sides ; spikelets 22-25, 3-grained ; 0 = 29-32 ; awns 12-14 cm. long (Ear
type i, Fig. 144).

Empty glume, 8-9 mm. long, apical tooth broad and blunt, i mm. long (u,
Fig. 138).

Grain, flinty or semi-flinty, usually plump with bluntish apex ; 7-6 mm. long,
3-5 mm. broad, 3-3 mm. thick.

Several forms, chiefly received from Russia under the names Bieloturka
and Arnovka, possess rough or slightly pubescent leaves and suggest a relation-
ship with T. turgidum or T. vulgar e.

5. Spring form received from Russia and Greece and from Spain under the
name Bieloturka. Slightly taller than the previous winter form, with denser
ears (0 = 33-36).

6. Forms from Turkey and Asia Minor under the names Sari Bashak, Sari
Sakij, and Kizil Bogdai have the following characters :

Ear, short, dense, compressed, 6-7-5 cm- ^ong> 6-9 mm. across the face, 14-16
mm. across the side ; spikelets 20-25 ; 0 = 33-36 ; awns 12-14 cm. long (Ear
type 2, Fig. 144).

Empty glume, about 10 mm. long, flattened, with acute, apical tooth (8, Fig.

Grain, flinty, long, and narrow, 7-5-8 mm. long, 2-5-3 mm- broad, 2-5-3 mm-
thick.

Ear bearded ; glumes red, glabrous ; awns red ; grain red.
T. durum, var. murciense, Korn. Handb. d. Getr. i. 71 (1885).

MACARONI WHEAT 223

Kornicke records this variety from Spain (Murcia), Portugal, Persia, Altai,
and Egypt.

Flaksberger mentions the occurrence of dense-eared forms in Turkestan and
the Russian provinces of Kuban and Samara.

I have had examples from Portugal, Bulgaria, Greece, Asia Minor, and
Burma.

A widely distributed but somewhat rare variety, the chief forms of which
have long, narrow ears, 8-10 cm. square in section. A few are found in the
eastern Mediterranean region with denser, shorter, and somewhat compressed
ears.

1 . Forms received from Bulgaria, Turkey, Greece, and Jaffa have the follow-
ing characters :

Young shoots, semi-erect ; young leaves slightly rough.

Straw, tall to very tall, 132 cm. (about 53 inches) long ; upper internode
hollow.

Ear, 8-10 cm. long, square, 10 mm. across the sides; spikelets 23-27, 3-
grained ; 0=29-31 ; awns 13-14 cm. long (Ear type 2, Fig. 141).

Empty glume, 9-10 mm. long ; apical tooth narrow, acute, i mm. long (14,
Fig. 138).

Grain, flinty, apex blunt, 7-65 mm. long, 3-15 mm. broad, 3-5 mm. thick.

2. Santa Marta. — Received from Portugal.

Young shoots, erect ; young leaves slightly rough, old leaves glabrous.

Straw, tall, no cm. (about 43 inches) high ; upper internode solid.

Ear, lax, 8-10 cm. long, square, 10-11 mm. across the sides ; spikelets 19-22,
3-grained ; D =24-25 ; awns 14 cm. long (Ear type i, Fig. 142).

Empty glume, 10 mm. long ; apical tooth narrow, acute, about i mm. long
(6, Fig. 138).

Grain, flinty, with truncate apex ; 8-3 mm. long, 3-7 mm. broad, 3-7 mm.
thick.

A form of hordeiforme is also found under this name in Portugal.

3. Received from Greece.

Young shoots, erect ; young leaves glabrous.

Straw, tall, 127 cm. (about 50 inches) high ; upper internode solid.
Ear, short and flat, 7-8 cm. long, 8-9 mm. across face, 12 mm. across side ;
spikelets 22, 2- to 3-grained ; D = 29-32 ; awns slender, 12 cm. long (Ear type 2,

Fig- MS)-

Empty glume, 9 mm. long ; apical tooth short, broad at base, -5-1 mm. long
(10, Fig. 138).

Grain, flinty, with truncate apex, 8-2 mm. long, 3-7 mm. broad, 3-95 thick.

Ear bearded ; glumes red, glabrous ; awns black ; grain white.

T. durum, var. erythromelan, Korn. Handb. d. Getr. i. 71 (1885).
Kornicke received this variety from Murcia, Spain. It is chiefly found in
Spain and Portugal ; very rare in Russia.

224 THE WHEAT PLANT

1. Medea or Trigo raspinegro de Medea. — Received from Spain.
Young shoots semi-erect ; young leaves glabrous.

Straw, of medium height, 106 cm. (about 42 inches) high, upper internode
solid.

Ear, lax and square, 9-10 cm. long, 10-11 mm. across the sides ; spikelets
22, 3 -grained ; 0 = 25-26; awns 17-18 cm. long, their bases jet black (Ear
type i, Fig. 142).

Empty glume, 10 mm. long ; apical tooth acute, i mm. long (6, 9, Fig. 138).

Grain, semi-flinty, 8 mm. long, 3-7 mm. broad, 3-6 thick.

2. Russian Form.

Young shoots, semi-erect ; young leaves glabrous.

Straw, short, 90 cm. (about 35 inches) high ; upper internode hollow.

Ear, lax, 8-9 cm. long, square, 10 mm. across the sides ; spikelets 20-22,

2- to 3-grained ; D =27 ; awns 15 cm. long (Ear type i, Fig. 142).

Empty glume, 9 mm. long ; apical tooth acute, i mm. long (8, Fig. 138).
Grain, flinty, dorsal ridge prominent, 7-7 mm. long, 3-4 mm. broad, 3-2 mm.
thick.

Similar forms from Spain.

3. Trigo pinot and others received from Spain.
Young shoots, erect ; young leaves glabrous.

Straw, tall, 140 cm. (55 inches) high ; upper internode solid.
Ear, dense, 6-5-8 cm. long, square, 10-13 mm. across the side or compressed,
9 mm. across the face, and 11-12 mm. across the side; spikelets 21-23,

3- to 4-grained ; 0 = 32-35 ; awns 15 cm. long (Ear type i, Fig. 144).

Empty glume, 9-10 mm. long ; apical tooth short, acute, i mm. long (5, 7,
Fig. 138).

Grain, flinty, 8 mm. long, 3-8 mm. broad, 3-75 mm. thick.

Trigo Colorado, Trigo berberisco, and others from Spain are similar to this
form.

4. Trigo bascufiano and others from Spain, and Mourisco vermelho from
Portugal.

Young shoots, semi-erect ; young leaves glabrous.

Straw, tall, 130 cm. (about 51 inches) long ; upper internode solid.

Ear, short, dense, and tapering towards the apex ; 5-7 cm. long, 10 mm.
across the face, 13-15 mm. across the sides ; spikelets 20-23, 2- to 3-grained ;
D = 37-40 ; awns 13-15 cm. long (Ear type 2, Fig. 144).

Empty glume, 9-10 mm. long ; apical tooth broad and bluntish, 1-1-5 mm-
long (9, n, Fig. 138).

Grain, flinty, with high dorsal ridge; 8-8-2 mm. long, 3-1-3-4 mm. broad,
3-6-3-8 mm. thick.

5. A form from Alexandretta, Asia Minor, has the following characters :
Ear, short, compressed, 6-7 cm. long, 8-9 mm. across the face, 13-14 mm.

across the side ; spikelets 20 ; D = 35 ; awns stout, jet black, about 14 cm. long
(Ear type 2, Fig. 144).

FIG. 144. — MACARONI WHEAT (T. durum, Desf.).

i. var. affine.
(Sorrentino.)

2. var. alexandrinum.
(Canary Isles.)

MACARONI WHEAT 225

Empty glume, oval, very broad, 9 mm. long, 4 mm. wide (9, 12, Fig. 138).
Grain, flinty, 7-7-5 mm. long, 3 mm. broad, 3 mm. thick.

Ear bearded ; glumes red, glabrous ; awns black ; grain red.

T. durum, var. alexandrinum, Korn. Handb. d. Getr. i. 71 (1885).
Kornicke's type was obtained from Lower Egypt through L. Wittmack. A
comparatively rare variety which I have received only from Spain and the

Canary Isles.

t

1. Trigo bascunano from Spain.

Young shoots, erect ; young leaves glabrous or slightly rough.

Straw, tall, 127 cm. (50 inches) high ; upper internode solid.

Ear, dense, 7-8 cm. long, 10-12 mm. across the face, 13-16 mm. across the
side ; spikelets 22-23, 2~ to 3-grained ; 0 = 33-35 5 awns 15-17 cm. long (Ear
type i, Fig. 144).

Empty glume, 9-10 mm. long ; apical tooth slightly curved, acute, i mm.
long (9, Fig. 138).

Grain, semi-flinty, 8-25 mm. long, 3-65 mm. broad, 3-8 mm. thick.

Allied to this are short-eared forms from the Canary Isles and a long-eared
form (ears 9-10 cm. long and 0 = 28-30).

2. Spanish Form.

Young shoots, erect ; young leaves glabrous or slightly rough.

Straw, of medium height, 90 cm. (about 35 inches) high ; upper internode
hollow.

Ear, short, dense, 6-8 cm. long, square, 10-12 mm. across the sides or slightly
compressed; spikelets 20-24, 3- to 4-grained ; 0 = 30; awns divergent, 12-
13 cm. long (Ear type i, Fig. 144).

Empty glume, 9-10 mm. long, keel and outer edge often black ; apical tooth
acute, 1-5-2 mm. long (9, Fig. 138).

Grain, flinty, 8 mm. long, 3-3-5 mm. broad, 3-3-5 mm. thick.

Ear bearded ; glumes red, pubescent ; awns red ; grain white.

T. durum, var. italicum, Korn. Handb. d. Getr. i. 73 (1885).

Kornicke obtained this variety from Italy, Spain (Murcia and Valencia),
Greece, and Egypt.

Forms of var. apulicum in some seasons do not develop the black colour in
the awns and may often be placed under this variety. All the forms I have seen
of this variety exhibit characters which suggest affinity with T. turgidum or T.
vulgar e.

Trigo Siciliani. — Received from Italy.

This form is not typical of T. durum ; the characters of leaves, ears, straw7,
and grain suggest an affinity with T. turgidum.

Young shoots, semi-erect ; young leaves pubescent.

Straw, of medium height, 90 cm. (about 36 inches) high ; upper internode solid.

Ear, large, 9-11 cm. long, square, 11-13 mm- across the sides; spikelets

226 THE WHEAT PLANT

25-26, 3- to 4-grained ; D =23-26 ; awns stout, 15 cm. long, somewhat divergent
(Ear type i, Fig. 141).

Empty glume, 10 mm. long ; apical tooth 3 mm. long ; secondary notch
distinct (4, Fig. 138).

Grain, very large, flinty, 10 mm. long, 3-9 mm. broad, 4 mm. thick.

A similar but smaller form, with ears 6-8 cm. long (D =30-35), was received
from Italy under this name.

Ear bearded ; glumes red, pubescent ; awns red ; grain red.

T. durum, var. aegyptiacum, Korn. Handb. d. Getr. i. 73 (1885).
Kornicke obtained this variety from Egypt, Apulia, and the Government of
Kharkov, Russia.

Vermelho fino. — A form showing affinities with T. turgidum or T. vulgar e
received from Portugal.

Young shoots, semi-erect ; young leaves with a few hairs.

Straw, of medium height, 90 cm. (about 36 inches) high; upper internode
hollow with thick walls.

Ear, 7-9 cm. long, square, 10-14 mm. across the sides; spikelets 19-22, 3-
grained ; 0=26-27 ; awns 15-17 cm. long, somewhat divergent (Ear type i,

Fig- 143)-

Empty glume, 11-12 mm. long ; apical tooth short, blunt, -5-1 mm. long (18,

*9> Fig- I38)-

Grain, very long, pale red, flinty, 10 mm. long, 3-5 mm. broad, 3-5 mm.

thick.

Ear bearded ; glumes red, pubescent ; awns black ; grain white.

T. durum, var. apulicum, Korn. Handb. d. Getr. i. 73 (1885).

Kornicke obtained forms of this variety with lax and dense ears from Apulia
(Italy), Andalusia (Spain), and a dense-eared form from Lower Egypt.

Flaksberger states that a lax-eared spring form and a dense-eared winter
form of this variety are cultivated in the Transcaucasus. I received a form with
long, lax, narrow ears from the United Provinces, India, and one with short
dense ears from Spain.

Cawnpore 18. — Indian form.

Ear, long, lax, and narrow, almost square in section, 8-9 cm. long, 9-10 mm.
across the sides ; spikelets 22-24 ; D = 28-32 ; awns 11-13 cm. long (Ear types
1,2, Fig. 142).

Empty glume, short, 8 mm. long ; apical tooth stout (8, 10, Fig. 138).

Grain, flinty, small, 7-7 mm. long, 3-1 mm. broad, 3 mm. thick.

Ear bearded ; glumes red, pubescent ; awns black ; grain red.

T. durum, var. niloticum, Korn. Handb. d. Getr. i. 73 (1885).

Kornicke obtained dense-eared specimens of this variety from Upper Egypt
and Apulia, Italy.

Flaksberger records isolated ears from Russia (the Don Provinces), and I
have had it from Asia Minor. Some of the forms I have seen of this variety

FIG. 145. — MACARONI WHEAT (T. durum, Desf.).

i. var. leucomelan.
(Trigo raspinegro de espiga pequena.)

2. var. niloticum.
(A dense eared Spanish form.)

MACARONI WHEAT 227

show affinities with T. turgidum or T. vulgare. The ears are chiefly of two types :
(i) very long, lax, and square in section, and (2) short, dense, compressed ears.

1 . A form from Spain, showing affinities with T. vulgare.
Young shoots, semi-erect ; young -leaves slightly pubescent.

Straw, of medium height, 95-100 cm. (about 36-38 inches) high ; upper
internode hollow.

Ear, 8-10 cm. long, square, 10-12 mm. across the sides ; spikelets 22-25,
3-grained ; D = 25-27 ; awns 13-14 cm. long (Ear type i, Fig. 142).

Empty glume, 8-10 mm. long ; apical tooth acute, 1-5-2 mm. long (5, Fig.

138).

Grain, pale red, flinty, 7-5-8-5 mm. long, 3-3 mm. broad, 3-4 mm. thick.

2. Spanish Form.

Young shoots, semi-erect ; young leaves glabrous.

Straw, of medium height, 95-100 cm. (38-40 inches) high ; upper internode
solid.

Ear, short and dense, 6 cm. long, compressed, 13-14 mm. across the face,
17-20 mm. across the sides ; spikelets 21, 3- to 4-grained ; D = 38 ; awns 15-16
cm. long (Ear type 2, Fig. 145).

Empty glume, 10-11 mm. long; apical tooth narrow, 1-5-2 mm. long (4,
Fig. 138).

Grain, flinty, long, and pointed at the apex, 9-4 mm. long, 3-3 mm. broad,
3-8 mm. thick.

Ear bearded ; glumes blue-black, glabrous ; grain white.

T. durum, var. provinciale, Korn. Handb. d. Getr. i. 71 (1885).

In some seasons the glumes are a dark reddish tint.

A comparatively rare variety received from Spain and Morocco, and recorded
by Flaksberger as of casual occurrence in South Russia.

Spanish Form.

Young shoots, erect ; young leaves glabrous.

Straw, tall, 135 cm. (about 53 inches) high ; upper internode hollow.

Ear, somewhat lax, 9 cm. long, square, 12-14 mm. across the sides ; spikelets
23, 3- to 4-grained ; 0 = 27 ; awns 15-17 cm. long (Ear type 2, Fig. 141).

Empty glume, 10-11 mm. long ; apical tooth narrow, acute, -5-1 mm. long (4,
Fig. 138).

Grain, flinty, with high dorsal ridge and blunt apex ; 8-25 mm. long, 3-65
mm. broad, 3-8 mm. thick.

Ear bearded ; glumes blue-black, glabrous ; grain red.

T. durum, var. obscurum, Korn. Handb. d. Getr. i. 72 (1885).
A rare variety received by Kornicke from Spain.

Ear bearded ; glumes blue-black, pubescent ; grain white.

T. durum, var. coerulescens, Korn. Handb. d. Getr. i. 73 (1885).
The hairs on the glumes are usually few and inconspicuous. In some seasons

r

228 THE WHEAT PLANT

the empty glumes of most forms are dark reddish tint with a black stripe along
the outer margins. Kornicke obtained samples of this variety from Spain.
Italy, Upper Egypt, and Chili.

Flaksberger records its occurrence in S. and S.E. Russia, Transcaucasia,
Siberia, and Turkestan, and Asia Minor, Turkey (Adrianople).

I have had specimens from Russia.

1. Received from Semipalatinsk, Russia, and in commercial " Hard Tagan-
rog."

Young shoots, erect ; young leaves glabrous.

Straw, tall, slender, 132 cm. (about 52 inches) long ; upper internode hollow.

Ear, lax and square, 8-9 cm. long, 10-12 mm. across the sides ; spikelets
20-23, 3- to 4-grained ; D =26-28 ; awns 14-15 cm. long (Ear type i, Fig. 142).

Empty glumes, 10-11 mm. long ; apical tooth narrow, acute, 1-1-5 mm. long
(8, n, Fig. 138).

Grain, flinty, with narrow apex, 8-5 mm. long, 3-5 mm. broad, 3-5 mm. thick.

2. Kara Bashak from Adrianople and Arab boydd from Asia Minor are
forms of coerulescens with compressed ears, 6-7 cm. long (Ear type 2, Fig. 144).

Ear bearded ; glumes blue-black, pubescent ; grain red.

T. durum, var. libycum, Korn. Handb. d. Getr. i. 73 (1885).

The glumes are only slightly pubescent, and the hairs easily overlooked as
in the previous variety.

Kornicke's type was collected by P. Ascherson in the oasis Merdisheh in the
Libyan Desert (1876) ; the native name is "Tuedi."

Flaksberger records the rare occurrence of var. libycum in the Samara Pro-
vince, Russia.

I have received forms of it from Morocco, Spain, Portugal, and Bulgaria.

1. Morocco Form.

Young shoots, erect ; young leaves glabrous.

Straw, of medium height, 106 cm. (about 42 inches) long ; upper internode
solid.

Ear, 7-8 cm. long, square, 11-13 mm- across the sides ; spikelets 17-20, 3- to
4-grained ; D = 28-30 ; awns stout and very long, some of them reaching a length
of 23 cm. (Ear type 2, Fig. 138).

Empty glume, 11-13 mm- l°ng > apical tooth acute, curved, 1-2 mm. long
(4, Fig. 138).

Grain, flinty, 8-2-8-6 mm. long, 3-3-4 mm. broad, 3-3-4 mm. thick.

2. Madonna. — Deceived from Haage and Schmidt, Erfurt, Germany.
Young shoots, semi-erect ; young leaves glabrous.

Straw, very tall, 150 cm. (about 60 inches) high ; upper internode hollow
with thick pithy walls.

Ear, dense, 7-8 cm. long, square, 11-13 mm- across the sides, or compressed,
10-12 mm. across the face, and 14-15 mm. across the side ; spikelets 26, 3-
to 4-grained ; D = 33-34 ; awns 15-17 cm. long (Ear type 2, Fig. 143).

FIG. 146. — MACARONI WHEAT (T. durum, Desf.).

i. var. affine.
(Asia Minor.)

2. var. australe.
(Huguenot.)

MACARONI WHEAT 229

Empty glume, 9 mm. long; apical tooth 2-3 mm. long, the tip sometimes
curved outwards (5, Fig. 138).

Grain, flinty, small, with high dorsal ridge, 7-1 mm. long, 3-15 mm. broad,
3-4 mm. thick.

Javardo from Portugal is similar, but in some seasons the empty glumes are
reddish or yellowish with black keels and edges.

3. Trigo raspinegro espiga negra. — Received from Spain.

Young shoots, erect ; young leaves glabrous.

Straw, of medium height to tall, 135 cm. (about 53 inches) high ; upper inter-
node solid or hollow with thick walls.

Ear, slightly tapering towards the apex, 8-9 cm. long, almost square, 11-13
mm. across the sides ; spikelets 19-20 ; 0 = 30-34 ; awns 18 cm. long, slightly
divergent (Ear type i, Fig. 143).

Empty glume, black, with a bluish-white surface, 9 mm. long ; apical tooth
1-1-5 mm- long with broad base (4, 6, Fig. 138).

Grain, flinty, compressed, with high dorsal ridge, apex truncate, 8-2 mm.
long, 3-65 mm. broad, 3-9 mm. thick.

A closely similar form to this from Bulgaria has ears 7-9 cm. long, almost
square, 10-12 mm. across the sides, or compressed, 10 mm. across the face, and
13 mm. across the sides.

Ear beardless ; glumes blue-black, glabrous ; grain white.

T. durum, var. australe, mihi.

Huguenot or Bald Medeah (2, Fig. 146). — Received from Australia, where
it is grown as a hay crop and for green feed in the coastal regions.

At Reading it is very early, coming into ear before the end of May.

Guthrie states that it originated from a single plant with two beardless heads
in a crop of Medeah (var. erythromelan, p. 224). The selection was made in
1897 by J. Correll, Arthur River, Western Australia. G. L. Sutton notes that
the original plant is supposed to have been a natural cross between Medeah and
Purple straw ( T. vulgare, var. lutescens).

Young shoots, erect.

Straw, stiff, erect, tall, 127 cm. (about 50 inches) high ; upper internode solid
and often wavy below the ear.

Ear, dense, 6-5-7 cm. long, tapering towards apex and bases, 9-10 mm.
across the face, 15 mm. across the side ; spikelets 22-24, 3~grained ; D = 38.

Empty glume, 9-10 mm. long; apical tooth small, very short, -5 mm. long;
secondary tooth of empty glume distinct (14, Fig. 138).

Grain, flinty, with high dorsal ridge and bluntish apex ; 7-8 mm. long, 3-5
mm. broad, 3-5 mm. thick.

Ear beardless ; glumes red, glabrous ; grain white.
T. durum, var. sub-australe, mihi.

Received from Australia in a sample of Bald Medeah (the previous variety),
with which form it agrees in all respects except in the colour of the glumes.

CHAPTER XVI

POLISH WHEAT

Triticum polonicum, L. Sp. PI. ed. ii. 127 (1762).
T. levissimum, Haller. Stirp. ind. Helv. 209, No. 1423 (1768).
T. glaucum, Moench. Method. 174 (1794).

Gigachilon polonicum, Seidl. Bercht. und Seidl, Oek. tech. Flora Bohmens, i.
425 (1836).

Deina polonica, Alef. Landw. Fl. 336 (1866).

POLISH wheat is the most modern of all the races or sub-races of wheat,
there being no evidence of its existence before the first half of the seven-
teenth century.

Bauhin and Cherler (Historia Plantarum universalis, ii. 410, 1651)
mention a " Triticum speciosum grano longo " obtained from the Botanic
Garden at Stuttgart, which probably refers to Polish wheat, but the name
T. polonicum is first met with in Hermann's list of plants grown in the
Botanic Garden at Leyden during 1681-1686 (P. Hermann, Hortus
Academicus Lugduno-Batavus, 609 (1687)).

The second reference to this wheat occurs in 1690 in Bobart's catalogue
published as an appendix in Ray's Synopsis Methodica Stirpium Britan-
nicarum (p. 236), where it is described as " Triticum Poloniae. Triticum
majus gluma foliacea," with the note, " This I received first out of Wor-
cestershire, where it is sown in the fields."

Plukenet in 1692 gives an excellent figure of an ear of the wheat in
his Phytographia (Plate CCXXXL, Fig. 6) under the title " Triticum
Poloniam H. Oxon ab ingeniosissimo Hortulano D. Harrison accepimus,"
and an ear similar to the one figured exists in Plukenet's Herbarium in
the British Museum (Herb. Sloane, vol. xcviii. fol. 124).

Later in his Almagestum, 1696 (p. 378), Plukenet refers to the wheat
as Triticum Polonicum, the name ultimately adopted by Linnaeus.

Morison (Plantarum Historia Oxoniensis, iii. 175 (1699)) describes it
as " Triticum majus longiore grano glumis foliaceis incluso Poloniae
dictum," and two specimens, var. levissimum and var. villosum, are found
in the Morisonian Herbarium at Oxford.

230

POLISH WHEAT 231

From its first appearance it seems to have been grown in Botanic
Gardens throughout Europe, and Haller in 1768 (Stirp, ind. Helv. ii. 209)
says it was cultivated in Thuringia and in small amount in Switzerland.

Although the name " Polish " is associated with it in almost all the
countries in which it is found, the origin of the name and the early con-
nection of the wheat with Poland is obscure. Seringe (Cer. eur. 182)
states that in 1816 and 1817 sixty vessels laden with this wheat arrived in
France from Dantzig, and concludes that it was cultivated on a large scale
in some of the northern countries. I have, however, discovered no
satisfactory evidence of its growth in Poland before 1870 (Rostafinski,
Fl. Poloniae prodromus, 26 (1873)).

Polish wheat is a delicate spring cereal, and requires a hot climate and
fertile soil for satisfactory growth. In damp seasons at Reading it is
almost completely sterile and in the best years there gives only poor yields
of grain ; the anthers frequently remain very small and shrivel, the fila-
ments not elongating as in the other wheats. It is nowhere prolific and its
cultivation is of little importance, since it possesses no qualities which
are not found in greater degree in T. durum.

Small isolated areas are grown in several countries bordering the
Mediterranean, more especially in Algeria, Spain, and Italy, where its
grain is employed in the manufacture of macaroni. Schimper found it
in Abyssinia with the native name " Fellasito," and I have had samples
of it from Idaho, U.S. A., and from Argentina under the name " Trigo
Chile de fideos." Flaksberger informs me that it is not found as a field
crop in European Russia or Siberia, but is cultivated on a small scale in
the Heyrabad district in Turkestan, where it is considered to be of Siberian
origin.

To Polish wheat various names have been given, such as Siberian,
Astrakhan, Mogadur, Surinam, and Chinese wheats. In outline the long
narrow grain somewhat resembles a grain of rye, and with a specimen in
the Herbarium at Kew is the record of 4 acres grown in Essex in 1858 and
described by Dickson & Sons as " Giant Rye."

Kornicke states that a sample was sent to the Vienna Exhibition
(? date) as Montana Rye from the United States, and that two samples
obtained first and second prize as Rye at the Denver Exhibition, U.S.A.,
of 1882.

GENERAL CHARACTERS OF T. polonicum, L.

The typical form of T. polonicum is strikingly different from all other
wheats in possessing large ears with long, narrow, empty glumes of a
glaucous hue, which extend beyond the rest of the spikelet.

It is one of the tallest wrheats, its culms usually reaching a height of
140-160 cm. (about 55-60 inches). These generally have 5 internodes

232

THE WHEAT PLANT

above ground, averaging about 5-5 cm., 10-5 cm., 15 cm., 28 cm., and 54
cm., increasing from below upwards.

The upper internodes are generally solid, the
lower ones more or less hollow. The tillering
power is small, each plant rarely producing more
than three or four straws.

I

i

FIG. 147. — Empty glumes FIG. 148. — .Grains of
of Polish wheat (T. Polish wheat (T.

polonicum) (nat. size). polonicum), front,

back, and side
views (nat. size).

The shoots of the young plants are of
upright habit, their leaves bluish-green,
with comparatively smooth surfaces like
those of T. durum.

The culm leaves are almost glabrous,
the upper ones 2 cm. or more broad with
small auricles, which are generally free
from hairs.

The ears have from 19 to 23 spikelets,
three or four of the lower ones being
abortive ; in some varieties they are lax,
14-16 cm. long, somewhat square in sec-
tion, with a density of 15-18 ; in others
they are 7-9 cm. long, oblong in section,
and compact, the density being about 30.

The rachis is smooth and flattened,
the margins fringed with hairs, which
are longest near its nodes ; a frontal tuft
of hairs 2-2-5 mm- l°ng is als° present

FIG. 149. — Grains of the spikelets
of one side of an ear of Polish
wheat (T. polonicum) (nat.
size).

POLISH WHEAT 233

at the base of each spikelet. The spikelets are large, flattened, 3-4 cm.
long, consisting of four or five flowers, of which two (or three) only
produce grain, the others having anthers devoid of pollen and imperfectly
developed ovaries.

The empty glumes are glabrous or pubescent, the hairs in the latter
case usually very short and easily overlooked ; they are as long as, or
slightly longer than, the rest of the spikelet, 2-4 cm. long, lanceolate,
and narrow, each measuring 4-5 mm. across its outer, wider, and convex
half, and 2-5-3 mm- across the inner concave half, which is closely ad-
pressed to the former.

They are fringed along the keel and possess 8-10 nerves, 5 or 6 on the
outer broad half, 2 or 3 on the inner half, and a central nerve, which
terminates on the short primary tooth -5-3 mm. long ; the secondary
tooth is very short or altogether absent (Fig. 147).

The flowering glumes of the two lowest flowers are from 2 to 3 cm. long
and 8-10 mm. broad, the upper ones being much shorter ; they are boat-
shaped, rounded on the back with 15-17 nerves, their membranous edges
fringed with short hairs.

Those of the two lowest flowering glumes of the spikelet are bearded,
each awn being 7-12 cm. long, frequently with a kink or bend in the basal
portion ; the awns of the upper flowering glumes are either very short,
measuring not more than about 5 mm. long, or are entirely missing.

The paleae are only i'2-i'5 cm. long, even in the lower flowers, whose
flowering glumes are about twice this length ; they are of the usual
bicarinate lanceolate form, with a slightly divided tip, and possess 4
nerves.

The thin rachilla is convex, with fine hairs on one side and flat and
more or less glabrous on the other.

The grains, which are the largest of all wheats, are yellowish- white,
or pale red with flinty endosperm ; they are long and narrow, measuring
11-12 mm. long and 4 mm. broad (Figs. 148, 149).

VARIETIES OF T. polonicum, L.

1. Glumes white, glabrous ; awns short.

a. Ears cylindrical ...... var. incertum, Korn.

b. Ears quadrate ..... var. submuticum, Link.

c. Ears flattened, compact.

i. Short ears ..... var. compactum, Link,

ii. Long ears ..... var. elongatum, Korn.

2. Glumes white, glabrous ; awns long.

a. Ear quadrate.

i. Grain white ..... var. levissimum, Korn.

ii. Grain red ..... var. chrysospermum, Korn.

234 THE WHEAT PLANT

b. Ear flattened,
i. Awns white.

A. Ear broad at base.

(1) Spikelets narrow . . . var. abessinicum, Korn.

(2) Spikelets wide . . . var. attenuatum, Korn.

B. Ear broad at apex . . . var. intermedium, Korn.
ii. Awns black ..... var. nigrobarbatum, Korn.

3. Glumes white, pubescent ; awns short.

a. Ears flattened ....... var. vestitum, Korn.

4. Glumes white, pubescent ; awns long.

a. Ear cylindrical.

i. Grain white ..... var. speltiforme, Korn.

ii. Grain red . . . . . var. novissimum, Korn.

b. Ears quadrate ....... var. villosum, Korn.

c. Ears flattened.

i. Grain white ..... var. Martinari, Korn.

ii. Grain red ...... var. Halleri, Korn.

5. Glumes pale red, glabrous ; awns long.

a. Ears cylindrical ...... var. rufescens, Korn.

6. Glumes pale red, pubescent ; awns long.

a. Ears cylindrical ..... var. rubrovelutinum, Korn.

b. Ears quadrate.

i. Awns pale red ...... var. Seringei, Korn.

ii. Awns black ...... var. Vilmorini, Korn.

7. Glumes black, pubescent ; awns short.

a. Ears cylindrical ...... var. anomalum, Korn.

8. Glumes pale violet, pubescent.

a. Ears quadrate ...... var. violaceum, Korn.

9. Glumes blue-black, pubescent.

a. Ears quadrate ...... var. nigrescent, Korn.

A large number of varieties of T. polonicum have been described or figured ;
the majority of them are, however, not now in cultivation and do not appear to
have been grown except in Botanic Gardens.

I have not been able to procure viable grain of any forms except those
belonging to the varieties levissimum, villosum, and Martinari, and a single
specimen of aristinigrum found among a sample of villosum ; the three first
mentioned appear to be the only varieties found at present in field culture.

Seringe divided T. polonicum into four groups, viz. :

1. T. polonicum quadratum, long, lax, square-eared forms.

2. T. polonicum oblongum, forms with long, lax, somewhat cylindrical ears
with affinities near to T. durum.

3. T. polonicum compactum, forms with ovate dense ears.

4. T. polonicum deformatum, forms with deformed or branched ears with
short, poorly-developed grains.

POLISH WHEAT 235

In each group Seringe mentions several " variations," but these are now
entirely lost. Early specimens I have seen in various herbaria, and several of
the figures given by Seringe belonging to Groups i and 2 have narrow regular
ears, and empty glumes intermediate in length between the typical long-
glumed T. polonicum and T. durum ; I have little doubt that these are of hybrid
origin.

It is probable that T. polonicum originated as a mutation of T. durum, or as a
race resulting from the hybridisation of the latter with some other wheat, and
may have been specially unstable and subject to further mutation or natural
hybridisation in Seringe's day.

Branching of the ear is uncommon in T. polonicum, but occurs occasionally ;
Seringe's deformatum shows this character, and Bayle-Barelle (Mon. agr. Cer.
38, t. 2, Fig. 8) figures a proliferous ear under the title " T. polonicum hybridum
horti ticinensis " ; Metzger also describes and figures a compound ear of this
species.

Glumes white, glabrous ; ear cylindrical ; awns short.

T. polonicum, var. incertum, Korn. Handb. d. Getr. i. 100 (1885).
T. polonicum oblongum, var. J., Ser. Cer. eur. 147 (185) (1841).
The ears are long and cylindrical, the spikelets wide apart and adpressed to
the rachis.

Glumes white, glabrous ; ear quadrate ; awns short.

T. polonicum, var. submuticum, Link. Hort. Berol. i. 28 (1827).

T. polonicum, D., Metzger. Eur. Cer. 25, t. 5, B., C. (1824).

T. polonicum aristis brevibus, Krause. Getr. Heft iv. 5, t. 2, B. (1836).

Of this variety Metzger gives the following characters :

Straw, tall, 110-120 cm. (about 44-48 inches) high, solid, striated, yellowish-
white ; leaves 15-17 cm. long, 1-5 cm. wide.

Ears, 12-15 cm. (4-75-6 inches) long, lax, 4-sided, compressed, tapering
towards the apex ; spikelets 20-24, 25-30 mm. long, 18-20 mm. wide, 2- to 3-
grained, with short awns 1-5-5 cm. l°ng-

Empty glume, 2-5 cm. long, with six prominent nerves and two teeth, one
longer than the other ; flowering glumes as long as the empty glume with a short
awn and many nerves ; palea half as long as the flowering glume.

Grain, long, pointed, smooth, pale greyish- white, flinty.

Glumes white, glabrous ; ear flattened, short, compact ; awns short.

T. polonicum, var. compactum, Link. Hort. Berol. i. 28 (1827).
T. polonicum, E., Metzger. Eur. Cer. 25, t. V. Fig. C. (1824).
T. polonicum compactum, Krause. Getr. Heft iv. 7, Taf. 2, C., D. (1*836).
T. polonicum compactum, var. M., Ser. Cer. eur. 148 (186) (1841).
Metzger states that this variety is grown in Granada, and notes that in wet
years it produces very little grain.

The following description is from Metzger :

236 THE WHEAT PLANT

Straw, tall, 120 cm. (about 48 inches) high, erect, striated, and solid ; leaves
15-20 cm. long, 1-5 cm. wide.

Ear, usually very dense, but sometimes laxer, 7-5-10 cm. long, quadrate,
generally tapered towards the apex, but occasionally broader upwards ; spike-
lets 14-16, 2-5 cm. long, 1-25 cm. wide, 2- to 3 -grained ; awns i or 2, 1-5-5 cm-
long.

Empty glume, 2-5 cm. long, glabrous, with 5 nerves on the broadest half ;
apical teeth two, the outer larger than the inner one.

Grain, 1-25 cm. long, greyish-white, pointed, flinty, the surface wrinkled.

Glumes white, glabrous : ear long, flattened, compact ; awns short.

T. polonicum, var. elongatum, Korn. Handb. d. Getr. i. 102 (1885)
T.polonicum compactum, var. L., Ser. Cer. eur. 148 (186) (1841).
A variety with long compact ears mentioned by Seringe ; the ears are often
curved.

Glumes glabrous, white ; grain white ; ear quadrate, lax ; awns long.

T. polonicum, var. levissimum, Korn. Handb. d. Getr. i. 100 (1885).

T. levissimum, Haller. Hist. Stirp. indig. Helvetiae, ii. 209, No. 1423 (1768).

Deina polonica alba, Alef. Landw. Fl. p. 336 (1866).

This and its pubescent counterpart var. villosum are two of the most widely
distributed varieties of Polish wheat.

Young plants, erect ; young leaves glabrous.

Straw, tall, 110-120 cm. (about 44-48 inches) high, erect, solid, striated;
leaves 15-20 cm. long, 1-5-2 cm. wide.

Ear, 12-5-17-5 cm. (5-7 inches) long, very lax, slightly tapered at the apex ;
spikelets 14-18, 2- to 3-seeded, 2-5-3-5 cm- l°ng-

Empty glume, 25-30 mm. long, 3 mm. wide, with 5-6 prominent nerves on
the outer half ; keel ciliated, the hairs very fine ; apex with two teeth.

Flowering glume, as long as the empty glume, with fine brittle awns half as
long as the ear.

Grain, 1-25 cm. long or longer, somewhat pointed, flinty, the surface
wrinkled.

Glumes glabrous, white ; grain red ; ear quadrate, lax ; awns long.

T. polonicum, var. chrysospermum, Korn. Handb. d. Getr. i. 101
(1885).

Kornicke's type has white smooth ears 11-5 cm. long ; the empty glumes
2-5-3 cm. long ; awns of moderate length, and red grain 9 mm. long. He
states that at first the glumes were slightly pubescent, but in later seasons became
glabrous.

Glumes white, glabrous ; ear dense, flattened, broad at base ; spikelets narrow ;

awns long, white.

T. polonicum, var. abessinicum, Korn. Handb. d. Getr. i. 103 (1885).

POLISH WHEAT 237

Collected by Schimper in Abyssinia ; the native name is " Fellasito."

Kornicke states that the ears are 8 cm. long, slightly narrowed towards the
apex, very dense, strongly compressed, narrow across the face, but 2 cm. wide
across the 2-rowed side ; awns 15 cm. long. The glabrous pale yellow, empty
glumes are 2-5 cm. long, with a terminal awn-like point which in some instances
reaches a length of 18 mm.

Grain, white, somewhat short and broad, 7 mm. long (unripe specimens).

Steudel's T. abyssinicum (Synops. Gram. 342, No. 19) appears to be a similar
dense-eared form of T.polonicum grown in Abyssinia.

Glumes white, glabrous ; ear dense, flattened, broad at base ; spikelets wide ;

awns long, white.

T. polonicum, var. attenuatum, Korn. Handb. d. Getr. i. 104 (1885).
T.polonicum, var. longiaristatum, b., crassispicatum, Korn. Syst. Uebers. 15

(1873)-

Kornicke's type, which was obtained from a Botanic Garden, possesses ears
8 cm. long or longer and white grain.

Glumes white, glabrous ; ear broadest at apex ; awns long, white.

T. polonicum, var. intermedium, Korn. Handb. d. Getr. i. 104 (1885).

T. polonicum compactum, var. K., Ser. Cer. eur. 148 (186), t. 8 (16) Fig 4
(1841).

A " club-eared " form of T. polonicum mentioned and figured by Seringe.
The ears are lax at the base and dense at the blunt apex.

Glumes white, glabrous ; ear flattened ; awns long, black.

T. polonicum, var. nigrobarbatum, Korn. Handb. d. Getr. i. 109 (1885).
T. polonicum nigrobarbatum, Desv. From. 149 (1833).
T.polonicum compactum, var. O., Ser. Cer. eur. 148 (186) (1841).
A compact-eared variety mentioned by Desvaux and Seringe.

Glumes white, pubescent ; ear dense, flattened ; awns short.

T. polonicum, var. vestitum, Korn. Handb. d. Getr. i. 103 (1885).
T.polonicum compactum, var. N., Ser. Cer. eur. 148 (186) (1841).
A dense compact-eared variety mentioned by Seringe ; similar to var.
compactum, Link, but with pubescent glumes.

Glumes white, pubescent ; ear lax, cylindrical ; grain white or pale yellow.

T. polonicum, var. speltiforme, Korn. Archiv f. Biontologie, ii. 4.0-?
(1908).

A form sent to Kornicke by H. de Vilmorin and apparently of hybrid
origin (T. polonicum levissimum (or villosuni) x T. turgidum, var. lusitanicum,
Petianelle blanche). The ears are very narrow, lax, with awns of variable
length (0-18 cm.) ; empty glumes as long as the spikelets, the grain compara-
tively short like that of T. durum.

238 THE WHEAT PLANT

Glumes white, pubescent ; ear cylindrical ; awns long ; grain red.

T. polonicum, var. novissimum, Korn. Archiv. f. Biontologie, ii. 404
(1908).

A narrow-eared form apparently of hybrid origin (T. polonicum x T. durum ? ).

Glumes white, pubescent ; ear quadrate ; awns long.

T. polonicum, var. villosum, Korn. Handb. d. Getr. i. 101 (1885).

T. polonicum villosum, Desv. From. 146(1833).

T. polonicum, Lagasca. Gen. et sp. pi. 6 (1814).

T. polonicum, C., Metzger. Eur. Cer. 25 (1824).

T. polonicum quadratum, var. B., Ser. Cer. eur. 146 (184) (1841).

T. polonicum velutinum, Krause. Getr. Heft iv. 5, t. 2, A. (1836).

Deina polonica velutinum, Alef. Landw. Fl. 336 (1866).

This and its glabrous counterpart var. levissimum are two of the most widely
known varieties of T. polonicum.

Young shoots, erect or semi-erect.

Straw, tall, 125-130 cm. (about 50-52 inches) high ; upper internode almost
solid.

Ear, lax, 15-17 cm. long, 10 mm. across the face, 12-15 mm. across the 2-
rowed side ; spikelets 15-22, 30-35 mm. long, 3-4 abortive at the base of the
ear, the rest 3- to 4-flowered, the lower 2 or 3 flowers of each fertile ; D = 15-18.

Edges of the rachis fringed with hairs 1-2 mm. long.

Empty glume, thin, narrow, lanceolate, those of the lower flowers 30-35 mm.
long, keeled, outer half with five prominent longitudinal ciliate nerves, the
hairs very fine ; apex with short acute primary tooth and rudimentary secondary
tooth close to the primary tooth (i, Fig. 147).

Flowering glumes of the two lower flowers with awns 9-11 cm. long, those of
the upper flowers with a short point about i mm. long, palea of the lower flowers
about half the length of the flowering glume.

Grain, flinty, long and narrow, pointed at the base, blunt at the apex ; 10-12
mm. long, 3-3-1 mm. broad, 2-9 mm. thick (i, Fig. 148).

Glumes white, pubescent ; ear dense ; awns long ; grain white.

T. polonicum, var. Martinari, Korn. Handb. d. Getr. i. 104 (1885).

Kornicke's type has dense ears, 8 cm. long, attenuated towards the apex
the empty glumes with awn-like points 6 mm. long. It originated from a
sample exhibited as " Grano S. Martinaro " at the Italian Exhibition in Paris,
1878.

Trigo Chile de fideos (i, Fig. 150). — A form of var. Martinari received from
Argentina and Spain.

Young shoots, semi-erect.

Straw, slender, of medium height, 110-115 mm. (42-44 inches) high ; upper
node solid.

Ear, dense, 8-9 cm. long, outline more or less ovate, tapering towards the
•apex ; compressed, 10-15 mm. across the face, about 20 mm. across the 2-rowed

FIG. 150.— POLISH WHEAT (T. polonicum, L.).

i. var. Martinari. 2. var. villosum.

(Tngo Chile de fideos.)

POLISH WHEAT 239

side ; awns of the two lowest flowering glumes of each spikelet 10-13 cm. long,
those of the third flower about i cm. long ; spikelets 21-23, each 20-25 mm-
long, 10-14 mm. broad ; 0 = 28-30.

Empty glume, keeled to the base, 25-27 mm. long, outer half narrow, lanceo-
late with 5 or 6 nerves ; primary apical tooth 1-3 mm. long or longer ; secondary
tooth about i mm. long (2, Fig. 147).

Grain, flinty, pointed at the base, apex blunt; 9-5-11 mm. long, 3-3 mm.
broad, 3-2 mm. thick (2, Fig. 148).

Glumes white, pubescent ; ear dense ; awns long ; grain red.

T. polonicum, var. Halleri, Korn. Handb. d. Getr. i. 104 (1885).

A variety with dense ears 5-7 cm. long, 2 cm. or more wide, the shorter ears
more or less oval in outline. Kornicke's type was obtained from a Botanic
Garden.

Kornicke describes a var . pseudohalleri with a somewhat lax, square, glabrous
ear, remarking that it " differs from var. Halleri in its red grain " ! (Archiv f.
Biontologie, ii. 404 (1908).

Glumes pale red, glabrous ; awns long ; ears cylindrical.

T. polonicum, var. rufescens, Korn. Handb. d. Getr. i. 100 (1885).

Kornicke's type has narrow pale red ears, 9-12 cm. long, moderately dense
with long awns and reddish grain.

T. polonicum oblongum, var. G., Ser. (Cer. eur. 147 (185) (1841)), is similar,
but with very lax ears.

Glumes pale red, pubescent ; awns long, red ; ear cylindrical, lax.

T. polonicum, var. rubrovelutinum, Korn. Handb. d. Getr. i. 100 (1885).

T. polonicum oblongum, var. H., Ser. Cer. eur. 147 (185) (1841).

A variety with very lax, pale red, pubescent ears mentioned by Seringe.

Glumes red, pubescent ; awns long, pale red ; ear quadrate, lax.

T. polonicum Seringei, Korn. Handb. d. Getr. i. 101 (1885).
T. polonicum quadratum, var. C., Ser. Cer. eur. 146 (184) (1841).
A variety mentioned by Seringe.

Glumes pale red, pubescent ; awns long, black ; ear quadrate, lax.

T. polonicum, var. Vilmorini, Korn. Handb. d. Getr. i. 101 (1885).

T. polonicum quadratum, var. D., Ser. Cer. eur. 146 (184) (1841).

A variety mentioned by Seringe ( = T. polonicum, Vilmorin, No. 74, 1636).

Glumes black, pubescent ; ear cylindrical ; awns short.

T. polonicum, var. anomalum, Korn. Handb. d. Getr. i. 100 (1885).
T. polonicum oblongum, var. I., Ser. Cer. eur. 147 (185), PI. IX. (Fig. 2)
(1841).

240 THE WHEAT PLANT

A variety with lax, slightly-compressed ears, which are almost beardless,
possessing only a few short awns at the apex.

Glumes pale violet, pubescent ; ear quadrate, lax.

T. polonicum, var. violaceum, Korn. Handb. d. Getr. i. 101 (1885).
T. polonicum quadratum, var. E., Ser. Cer. eur. 146 (184) (1841).
A variety mentioned by Seringe, cultivated in the Botanic Garden at Lyon,
1836.

Glumes bluish-black, pubescent ; ear quadrate, lax.

T. polonicum, var. nigrescens, Korn. Handb. d. Getr. i. 101 (1885).
T. polonicum quadratum, var. F., Ser. Cer. eur. 146 (184) (1841).
A variety mentioned by Seringe with regularly arranged spikelets and blue-
black glumes and awns ( = T. polonicum, Vilmorin, No. 5 (1836)).

CHAPTER XVII

RIVET OR CONE WHEAT

T. turgidum, L. Sp. PI. 86 (1753).

T. vulgar e turgidum, Alef. Landw. Fl. 325 (1866).

T. sativum turgidum, Hackel. Nat. Pfl. ii. 2, 85 (1887).

THE existence of Rivet wheat in prehistoric times is very doubtful. Heer
obtained from the pile-dwelling deposits of Robenhausen (Switzerland)
grains and a portion of a dense ear which he attributed to this race, and
large grains described as those of T. turgidum have been recorded also
from deposits of Neolithic and Bronze Ages in Italy (Castione, Parma) ;
owing, however, to the difficulty of distinguishing grains of some of the
forms of T. turgidum, T. vulgar e, and T. durum, even in recent material
these conclusions must be considered problematical.

On similar grounds the statement of A. de Candolle, that he recognised
grains of this wheat among seeds taken from the sarcophagi of ancient
Egyptian tombs, should be accepted with reserve.

There are no descriptions of wheats among ancient Greek or Latin
authors which conclusively point to this race ; it is possible, however,
that the " triticum ramosum " of Pliny refers to one or other of the varieties
of T. turgidum with branched ears, of which the simple-eared forms
would, no doubt, exist.

The clear separation of T. turgidum from T. vulgare and T. durum
does not appear to have been recorded until the first half of the sixteenth
century, when Fuchs (1542) and other " herbalists " accurately describe
and figure it under the name " Welsch " wheat, i.e. wheat from the
Welschland (France and Italy).

Dodoens also speaks of it as Roman wheat or Triticum romanorum,
and distinguishes it from the allied Triticum typhinum (T. durum),
' Typhew " wheat, which he obtained from the Canary Isles and Spain.

Several varieties of this race of wheat were cultivated somewhat
widely in England in the sixteenth, seventeenth, and eighteenth centuries,
at which period they were designated Rivet, Cone, or Pollard wheats.
During the latter half of the nineteenth century they were less commonly

241 R

242 THE WHEAT PLANT

grown ; at the present time only one or two forms of Rivet wheat are
found on farms in this country, and these only on comparatively small
areas, chiefly in the southern counties. Their high yield will, however,
always attract attention, and there is some likelihood of their being in-
creasingly grown in the future on this account.

In Germany this race is termed " English Wheat " ; under the name
Ble* Poulard it is met with in France, but the home of Rivet wheat is in
the countries bordering the Mediterranean. It is especially frequent in
Spain and Portugal and Italy, but several forms are grown in lesser degree
in Bulgaria, Greece, and Turkey. It is also cultivated extensively in the
Provinces of Tiflis and Baku in Transcaucasia, and in smaller amount in
Turkestan and Siberia ; Howard records its occurrence in Baluchistan,
but I have not seen specimens.

Small areas are occasionally devoted to this wheat in Australia, Canada,
the United States, South Africa, Chili, and Algeria.

I have no specimens nor records of its cultivation from India, China,
Japan, or Abyssinia.

Although some of the red and blue-black velvet-chaffed varieties can
be cultivated as winter wheats in the south of England, France, and
Germany without much risk in ordinary seasons, the majority of the Rivet
wheats are delicate and easily damaged, or destroyed altogether by hard
frosts or continued rain.

They are practically immune to rust fungi. I have never seen a field
crop nor yet a small plot or row of any variety of T. turgidum attacked on
ordinary cultivated soils in this country, but in 1908 I observed an exten-
sive invasion of yellow rust (Puccinia glumarum) on Blue Cone wheat
(T. turgidum, var. iodurum) sown late in spring on a small plot of highly
manured ground, which had been trenched 3 feet deep.

Their tall straw is strong, the crop rarely lodges, and the stiff awned
ears are not readily damaged by sparrows and other birds.

The varieties grown in England have very narrow blue-green leaves,
which lie close to the ground during winter ; in February, when ordinary
varieties of T. vulgare usually exhibit a vigorous growth of leafy shoots
2 or 3 inches high, Rivet wheats have a disappointing appearance, and the
farmer who is not familiar with their habit is inclined to plough them up
at this season. They should, however, be left, for their progress later
invariably compensates for the apparent lack of vigour in the early stages
of their development.

The productive power of most varieties of T. turgidum is greater than
that of any other race of wheat when the soil is suitable and the climate
allows of a long growing period for the crop.

Their high grain-yielding capacity is correlated with the long vegetative
period of the plants, the usual possession of six or seven green leaves on

RIVET OR CONE WHEAT 243

each straw, as well as with the great length of their stems and leaves, and
consequent abundance of green assimilating tissue. Moreover, the
number of spikelets possessed by each ear is usually greater among the
varieties of T. turgidum than among the varieties of T. vulgar e.

To ensure the greatest success with these wheats in England they
should be sown not later than the middle of October on warm, well-drained
soils ; when sown in spring they ripen very late, and the grain at harvest
is often poorly filled.

A few varieties possess flinty grains, but the majority of Rivet wheats
have a soft, opaque, starchy endosperm, from which is obtained " weak "
flour, suited to the requirements of the biscuit-maker, but of less value to
the baker on account of the poor physical quality and reduced amount
of its gluten. The dough is somewhat greyish and " short " ; loaves of
bread made from Rivet flour are more or less dense and non-porous in
texture, and of small volume when compared with loaves made from the
same weight of dough prepared from the " strong " flour of certain
varieties of T. vulgar e.

The bran is thick and abundant.

Rivet wheats are chiefly utilised by the miller for mixing with the strong
Canadian and Russian wheats.

GENERAL CHARACTERS OF T. turgidum, L.

T. turgidum is the tallest of the wheats ; its straw, which is somewhat
slender but strong, with a dull striate surface, averages about 150 cm.
(69 inches), the length in different varieties measuring from 120 to 180 cm.
(48 to 72 inches) or more ; the upper internode is curved and in many
cases solid or filled with pith ; even in those varieties with all the internodes
hollow the wall of the straw is often thick and pithy and the central
cavity comparatively small. I have found none with walls as thin as
those of T. vulgar e.

Well-developed culms of many kinds possess six or seven internodes
above ground ; the average lengths of the successive internodes from
below upwards were found in a few common varieties to measure 5, 10-5,
14-5, 19, 29, and 60 cm. respectively.

In some varieties the young shoots have the prostrate " winter " habit ;
others have the semi-erect or erect " spring " habit.

The leaves of the young plants are short, narrow, and usually dark
bluish-green, those of the culm long and broad.

In almost all varieties both sides are invested with soft white hairs
from 1 20 to 250 p long, as in T. dicoccum, the leaves to the touch resembling
velvet and appearing clothed with a silvery covering when viewed along
the surface (Fig. 151). Some forms of var. dinurum have young leaves

244 THE WHEAT PLANT

which appear almost smooth, the hairs upon them measuring not more
than 32-64 p. long, the older leaves of such varieties being somewhat
scabrid.

The auricles are usually fringed with a few long hairs.

The ears are large, pendulous, heavy, and

|<4/j^j^ a nearly always bearded, the sides usually

parallel, though a few taper a little towards
the tip. The ears are square or oblong in

section, the form in this respect depending
FIG. 151. — Diagrammatic trans- 11 j •. r ^\- j Di-

verse sections of young leaves uPon the laxness or density of the ear and the

of (a) T. turgidum and T. number and size of the grains which develop
pyramidale. (6) T. turgidum . , -11^ T-> ->_i i i 1 j

(var. dinurum). *n each spikelet. Ears with closely packed

spikelets containing four or five grains are

almost square ; those in which the spikelets are densely packed and
contain only two or three grains are oblong in section and broadest across
the side. A few forms with lax ears and well-filled spikelets are broad
across, the face and narrow across the side like most of the varieties of
T. vulgar e.

The ears of different forms are from 7 cm. to 11-5 cm. in length, their
" density " varying from 21 to over 40, the individual rachis internodes
measuring from 2-5 mm. to 4-66 mm.

The number of spikelets possessed by an average ear varies from 19 1033.

The rachis is tough, except in a few forms, which are brittle like those
of T. dicoccum ', it is smooth, but copiously fringed along its edges with
white hairs, and at the apex of each of its internodes at the base of the
spikelets is a tuft of similar hairs from i to 2 mm. in length.

The spikelets are from 8 to 15 mm. broad, 10 to 13 mm. long, and about
4 mm. thick. They are usually 5- to y-flowered, and in some varieties many
of them produce three to five well-developed grains.

The empty glumes are white, yellow, red, or dark bluish, the black
or dark brown pigment in the latter case being modified by a glaucous
covering.

As in T. durum and other wheats, the power of forming black pigment
in the glumes and awns appears to be absent from some varieties ; in those
forms which possess it the colour is not always produced, its appearance
being dependent upon climatic conditions. In damp cold seasons, and
in very dry seasons, when the plant is prevented from developing its ears
fully, the colour is more or less completely kept in abeyance, but in bright
warm seasons it appears again, such variation occurring in plants of
" pedigree " lines.

In some forms the glumes are glabrous," in others they are pubescent.
They are comparatively short ; those of the lateral spikelets are 8-n mm.
long, more inflated than those of T. durum, unsymmetrical, and 5- to 7-

RIVET OR CONE WHEAT 245

nerved, the portion from the midrib to the outer edge being 4-5 mm. across,
the inner membranous part being half as wide. The strong apical tooth in
some varieties is acutely pointed, i-i'5 mm. long, and curved inwards, in
others short and blunt. A well-marked keel runs from the base to the

HIM

I 23456789

FIG. 152. — Empty glumes of Rivet wheat (T. turgidum) ( x 2).

tip, and a more or less prominent lateral nerve is present on the outer
broad half of the glume, running from the base to a point close to the
apical tooth, where in some forms it terminates in a short secondary
tooth (Fig. 152).

oooocce »»

FIG. 153. — Grains of Rivet wheat (T. turgidum), front, back, and side views (nat. size).

The empty glumes of the terminal spikelet are ovate and more or less
symmetrical (Fig. 78).

The flowering glumes are thin, fragile, and pale ; even in the black-
eared varieties the dark tint is not developed in the part covered by
the empty glume. They are oval, inflated, and boat-shaped, without a

246

THE WHEAT PLANT

keel, but possessing 9-15 fine nerves, which converge at the tip into the

terminal awn.

The awns are stout, yellowish- white, red,
or black, from 8 to 16 cm. in length, triangular
in section, the angles from the base to the
apex being set with scabrid forward-pointing
projections.

In a few varieties the awns diverge slightly
outwards ; usually, however, they are parallel
to each other and to the long axis of the ear.
Several kinds shed their awns when the grain
is ripe.

As a rule only the two lower flowering
glumes of the spikelet bear long awns ; those
of the third and higher flowers are much
shorter or altogether missing.

The palea is membranous and of the
normal bi-nerved form.

The typical grain is white, yellow, or
red, large, broad, and plump, though some-
times flattened on one side by pressure of
the empty glume, blunt or truncate at the
" brush " end, with a high dorsal arch or
hump behind the embryo. The furrow is
shallow and the ventral flanks rounded in
well-grown grains (Fig. 153).

In a few forms the caryopses are narrower
towards the apex, though rarely so much as
in T. durum and T. dicoccum. In a small
number the dorsal arch is reduced, the grain
then approximating to that of T. vulgare.

In some seasons, and especially when the
crop is sown late, the grains at harvest ex-
hibit a somewhat rough surface, and are
poorly filled ; this result is most evident
among varieties possessing the largest grains.

The embryo is comparatively small and
the endosperm usually opaque and starchy,
although in a few forms it is flinty.
Measurements of grains taken from the middle of the ear of thirty-
eight varieties gave the following results :

FIG. 154. — Grains of the spike-
lets of one side of an ear of
Rivet wheat (T. turgidum)
(Blue Cone) (nat. size).

RIVET OR CONE WHEAT

247

Length.

Breadth.

Thickness.

mm.

mm.

mm.

Average
Limits
Ratio ....

TAB

6-7-8-37

IOO

3-88
3-26-4-43
52-0

3.70
3-23-4-1
49-6

The long growing period, number of culm leaves, velvety leaf-surface,
the height, surface, and pithy interior of its straw, the density of the ear,
as well as the frequent occurrence of compound ears, indicate a close
relationship of T. turgidum with the tall European forms of T. dicoccum.

A few forms suggest affinity with T. durum, while others appear to be
allied to T. vulgare ; these are probably hybrids.

I. Ears bearded —

VARIETIES OF T. turgidum, L.

Glumes white, glabrous ; awns white,
i. Ear simple, grain white
ii. Ear simple, grain red
iii. Ear branched, grain red .
Glumes white, glabrous ; awns black,
i. Ear simple, grain white
ii. Ear simple, grain red
iii. Ear branched, grain red .
Glumes white, pubescent ; awns white.
i. Ear simple, grain white
ii. Ear simple, grain red
iii. Ear branched, grain red .
iv. -Ear branched, grain red-violet .
Glumes white, pubescent ; awns black.

i. Ear simple, grain white
Glumes red, glabrous ; awns red.
i. Ear simple, grain white
ii. Ear branched, grain white
iii. Ear simple, grain red
iv. Ear branched, grain red .
Glumes red, glabrous ; awns black,
i. Ear simple, grain white
ii. Ear branched, grain white
iii. Ear simple, grain red
Glumes red, pubescent ; awns red.
i. Ear simple, grain white
ii. Ear branched, grain white
iii. Ear simple, grain red
iv. Ear branched, grain red .

var. lusitanicum, Korn.

var. gentile, Korn.

var. columbinum, Korn.

var. melanatherum, Korn.

var. nigrobarbatum, Korn.

var. pavoninum, Korn.

var. megalopolitanum, Korn.

var. buccale, Korn.

var. centigranum, Korn.

var. modigenitum. Korn.

. var. Salomonis, Korn.

var. Dreischianum, Korn.

\a.r.pseudocervinum, Korn.

. var. speciosum, Korn.

. var. cervinum, Korn.

var. speciosissimum, Korn.

var. Plinianum, Korn.

. var. Mertensii, Korn.

var. pseudomirabile, mihi.

var. mirabile, Korn.

var. dinurum, Korn.

var. Linnaeanum, Korn.

248 THE WHEAT PLANT

8. Glumes red, pubescent ; awns black.

i. Ear simple, grain red . . . var. rubroatrum, Korn.

9. Glumes blue-black, glabrous ; awns black.

i. Ear simple, grain white .... var. Herrerae, Korn.
10. Glumes blue-black, pubescent.

i. Ear simple, grain red .... var. iodurum, Korn.

ii. Ear branched, grain red .... var. coeleste, Korn.

II. Ears beardless —

i. Glumes blue-black, pubescent.

i. Ear simple, grain red . . . var. subiodurum, Korn.

Ear bearded, simple; glumes white, glabrous; awns white; grain white.

T. turgidum, var. lusitanicum, Korn. Handb. d. Getr. i. 59 (1885).

A widely distributed variety cultivated in France, Spain, and Greece, and
according to Kornicke in Portugal, Italy, and Chili ; Flaksberger mentions the
occurrence of this variety in small amounts in Russia, Turkestan, and Siberia.

1. Trigo Santa Salamanca. — A spring form received from Spain.
Young shoots, semi-erect ; young leaves pubescent.

Straw, very tall, 140 cm. (about 55 inches) high ; upper internode solid.

Ear, lax, 10-11 cm. long, square, 10-11 mm. across the sides, or somewhat
compressed, 9-11 mm. across the face, and 11-14 mm- across the side ; edges of
the rachis with long hairs ; spikelets 23-26, 2- to 3-grained ; D =25-27 ; awns
rough, 16-20 cm. long (Ear type 2, Fig. 156).

Empty glume, flattish, 8 mm. long with acute apical tooth (2, Fig. 152).

Grain, very large, flinty, narrowed at the truncate apex, dorsal hump pro-
minent ; 8-9 mm. long, 4 mm. broad, 3-75 mm. thick.

2. A winter form received from Greece and Italy.

Young shoots, prostrate or semi-erect ; young leaves pubescent.

Straw, hollow, with thick walls.

Ear, 7-10 cm. long, square, 10-12 mm. across the side, or somewhat com-
pressed, 9-10 mm. across the face, and 10-12 mm. across the side ; spikelets
22-24, 3-grained ; D = 30-33 ; awns 9 cm. long (Ear type 2, Fig. 157).

Empty glume, 8 mm. long ; apex narrowed, sometimes with a black line along
the margin ; apical tooth short, acute (i, Fig. 152).

Grain, large, mealy ; apex truncate ; dorsal hump prominent ; 8'2-8'5 mm.
long, 4-4*4 mm. broad.

3. Pdtianelle blanche or Bid Hybride Galland. — A winter or early spring
form received from France.

Vilmorin states this form is cultivated in Central and Southern Italy, from
which country it was introduced into France ; it is a strong-growing form
adapted to alluvial and the stiffer loamy soils in a warm climate.

Young shoots, semi-erect ; young leaves pubescent.

Straw, very tall, coarse, striate, 152. cm. (about 60 inches) high; upper
internode hollow with thick walls.

FIG. 155.— RIVET WHEAT (T. turgidum, L,).

var. gentile.
(Pisana blanca.)

RIVET OR CONE WHEAT 249

Ear, 9-11 cm. long, much compressed, 9-10 mm. across the face, 14-15 mm.
across the side; spikelets 27-32, 2- to 3 -grained ; 0 = 33-36; awns 11-12 cm.
long, deciduous (Ear type 2, Fig. 156).

Empty glume, 6-5-8 mm. long, apical tooth short, blunt (6, Fig. 152).

Grain, more or less flinty, large, often dark brown at the radicle end, dorsal
hump prominent ; 8-5-8-9 mm. long, 4 mm. broad, 3-7 mm. thick.

4. A smaller form resembling Petianelle blanche, received from Greece.
Ears, 7-8 cm. long, compressed, 9-10 mm. across the face, 12-14 mm. across
the side. Empty glume, with more acute tooth, but grain as in Petianelle blanche.

Ear bearded, simple ; glumes white, glabrous ; awns white ; grain red.

T. turgidum, var. gentile, Korn. Handb. d. Getr. i. 59 (1885).
A widely cultivated variety chiefly grown in Spain, Italy, Turkey, and to a
small extent in France and the Midlands and eastern parts of England.

1 . Pisana blanca. — A distinct winter form received from Spain ; the flat-
faced and narrow-sided ear suggests affinity with T. vulgar e.

Young shoots, prostrate ; young leaves pubescent.

Straw, very tall, 152 cm. (about 60 inches) high ; upper internode solid.

Ear, lax, 9-11 cm. long, 13-15 mm. across the face, 9-10 mm. across the side ;
spikelets 19-24, widely separated and somewhat irregularly arranged on the
rachis, 3- to 4-grained ; 0 = 20-23 ; awns 10 cm. long, slender, divergent, and
deciduous (Ear type, Fig. 155).

Empty glume, 7-8 mm. long ; apical tooth blunt (6, Fig. 152).

Grain, mealy, somewhat narrowed at the apex, dorsal hump prominent ;
7-2-7-8 mm. long, 3-3-3-8 mm. broad, 3-3 mm. thick.

2. Poulard Blanc Taganrog. — A winter or spring form received from
France and Germany ; the name suggests that this form is of Russian origin,
but Flaksberger states that var. gentile is not found in Russia. Vilmorin states
that it is grown in Central France and gives a good yield if sown not later than
the early part of November ; it is adapted to calcareous soils and those of
moderate quality.

Young shoots, semi-erect ; young leaves pubescent.

Straw, tall, 140 cm. (about 56 inches) high ; upper internode solid or hollow
with thick walls.

Ear, 9-11 cm. long, apex tapered, square, 10-12 mm. across the face and side ;
spikelets 28-32, 3-grained ; 0 = 28-32 ; awns slender, 10-11 cm. long, decidu-
ous (Ear type i, Fig. 157).

Empty glume, short (7 mm. long), apex narrow, apical tooth acute, 1-1-5 mm-
long (i, Fig. 152).

Grain, visible between the small short glumes, mealy, plump ; apex narrowed,
truncate ; dorsal hump prominent ; 7-7-5 mm. long, 3-5-3-8 mm. broad, 3-5-3-8
mm. thick.

3. Biancone. — Spring form received from Italy.

Young shoots, erect ; young leaves pubescent ; hairs short.

250 THE WHEAT PLANT

Straw, of medium height, 125 cm. (about 50 inches) high ; upper internode
hollow.

Ear, 8-5-10 cm. long, more or less compressed, 9-10 mm. across the face,
12-13 mm. across the side ; spikelets 25-28, 3-grained ; 0 = 30 ; awns 12-13
cm. long, deciduous (Ear type i, Fig. 157).

Empty glume, 7-8 mm. long, apex narrow, apical tooth short and blunt (3, 6,
Fig. 152).

Grain, mealy, large, pale reddish-white, dorsal hump prominent ; 8-8-5 nun.
long, 4-4-3 mm. broad, 4 mm. thick.

4. Ak Bashak ( = White Ear). — A winter form received from European
Turkey.

Young shoots, prostrate ; young leaves pubescent.

Straw, of medium height, 125 cm. (about 50 inches) high ; upper internode
solid, or hollow with very thick walls.

Ear, 9-10 cm. long, compressed, 10 mm. across the face, 14-15 mm. across
the side; spikelets 22-25, 2~ to 3-grained; 0=25-29; awns 12-14 cm. long
(Ear type i, Fig. 157).

Empty glume, 9 mm. long, apical tooth short and blunt (6, 7, Fig. 152).

Grain, semi-flinty, large, reddish-white ; apex somewhat narrowed ; dorsal
ridge not very prominent ; 8-2 mm. long, 4 mm. broad, 3-5-3-7 mm. thick.

5. Poulard a six rangs. — A winter form received from France.
Young shoots, semi-erect ; young leaves pubescent.

Straw, tall, 127 cm. (about 50 inches) high ; upper internode hollow with thick
walls.

Ear, 6-8 cm. long, very dense, compressed, 11-12 mm. across the face, 14-15
mm. across the side ; spikelets 28-31, 3- to 4-grained ; D = 38-45 ; awns slender,
10 cm. long, sometimes deciduous (Ear type 2, Fig. 158).

Empty glume, 7 mm. long, apical tooth acute, i mm. long (2, Fig. 152).

Grain, mealy, plump, yellowish-red, dorsal ridge prominent ; 7-7-5 mm.
long, 3-5-4 mm. broad, 3-5-4 mm. thick.

Ear bearded, branched ; glumes white, glabrous ; awns white ; grain red.

T. turgidum, var. columbinum, Korn. Handb. d. Getr. i. 63 (1885).
The form of var. gentile with compound ears.

Ear bearded, simple ; glumes white, glabrous ; awns black ; grain white.

T. turgidum, var. melanatherum, Korn. und Wern. Handb. d. Getr. ii.
396 (1885).

T. turgidum, var. nemausense, Wittmack ex Korn. Handb. d. Getr. i. 59
(1885).

The empty glumes usually have a narrow black line along the outer margin,
or are more or less blotched all over with black pigment. This variety has a
wide range of cultivation, being grown in considerable amount in Italy, Spain,
Greece, and the Baku Province of Transcaucasia, and to a lesser extent in
France.

FIG. 156. — RIVET WHEAT (T. turgidum, L.).

i . var. iodurum.
(Blue Cone.)

2. var. Mertensii.
Greece.

RIVET OR CONE WHEAT 251

Kornicke received samples as " Giant wheat " from an exhibition in Denver,
Colorado, U.S.A.

A dense-eared form of wheat from Baluchistan has been referred to var.
melanatherum by Howard.

The Ble G-aragnon du Languedoc, cultivated in the south and north-west
of France, is a form of this variety or of var. nigrobarbatum .

1. Received from Italy and Greece.

Young shoots, semi-erect ; young leaves pubescent.

Straw, tall, 130 cm. (about 51 inches) high ; upper internode hollow.

Ear, long and lax, 9-10-5 cm. long, square, 12-13 mm. across the sides ;
spikelets 22-24, 3~grained ; 0=25-28 ; awns 14-15 cm. long (Ear type i, Fig.
158).

Empty glume, 9 mm. long, with a black line along the outer margin ; apical
tooth short and blunt (i, 8, Fig. 152).

Grain, mealy, large, dorsal ridge prominent ; 8 mm. long, 4 mm. broad, 3-9
mm. thick.

2. Carosellone del Molise. — A well-marked form received from Italy ;
similar forms also from Spain and Greece.

A form from Spain under this name has red grains (see below, var. nigrobar-
batum, Desv.).

Young shoots, erect or semi-erect ; young leaves pubescent.

Straw, tall, 132 cm. (about 52 inches) high ; upper internode hollow with
thick walls.

Ear, 6-5-8 cm. long, dense, compressed, 10-12 mm. across the face, 15 mm.
across the 2-rowed side ; spikelets 23-25, 2- to 3-grained ; D =38-44 ; awns 9-10
cm. long, black at the base, the upper half pale (Ear type i, Fig. 158).

Empty glume, 7 mm. long, apical tooth acute, in some cases blunter, 1-2 mm.
long (Forms i, 4, Fig. 152). Sometimes the glume is entirely white ; usually,
however, there, is a narrow black line along the outer margin, and in some
seasons the dark pigment spreads over the rest of the glume ; these colour
variations are found in plants raised from a single ear.

Grain, mealy, apex truncate, dorsal hump prominent, radicle end sometimes
dark brown ; 7-8 mm. long, 3-5-4 mm. broad, 3-5-4 mm. thick.

Ear bearded, simple ; glumes white, glabrous ; awns black ; grain red.

T. turgidum, var. nigrobarbatum, Korn. Handb. d. Getr. i. 60 (1885).

Kornicke obtained this variety from Spain, and Werner mentions its cultiva-
tion in the south of France.

Flaksberger states that a winter form of it is found in the Tiflis and Baku
Provinces of Transcaucasia.

I have had it from Greece, Spain, and the United States.

i. Phillipino, received from the United States of America, and forms sent
from Spain and Greece, belong to this variety.

Young shoots, semi-erect ; young leaves pubescent.

252 THE WHEAT PLANT

Straw, tall, 132 cm. (about 52 inches) high ; upper internode solid or hollow
with thick walls.

Ear, 8-9 cm. long, usually square, 12-15 mm. across the sides; small ears
much compressed, 9-10 mm. across the face, 12-14 mm. across the side ; spike-
lets 22-25, 3" to 4-grained ; 0 = 28-32 ; awns 12-13 cm. long (Ear type 2, Fig.

157).

Empty glume, 8-9 mm. long, with a black line along the outer margin, or, in
some seasons, more or less uniformly black ; apical tooth 2 mm. long, acute,
broad at the base (i, Fig. 152).

Grain, more or less flinty, dorsal ridge prominent ; 7-6 mm. long, 4 mm.
broad, 3-5 mm. thick.

2. A form from Spain under the name Carosellone del Molise, with short
dense grain similar to those of Carosellone del Molise (var. melanatherum) from
Italy.

Ear bearded, branched ; glumes white, glabrous ; grain red ; awns black.

T. turgidum, var. pavoninum, Korn. Handb. d. Getr. i. 63 (1885).
The form of var. nigrobarbatum with compound ears.

Ear bearded, simple ; glumes white, pubescent ; awns white ; grain white.

T. turgidum, var. megalopolitanum, Korn. Handb. d. Getr. i. 60 (1885).
A rare variety found by Kornicke in a seedsman's sample of " Greek wheat
from Megalopolis," and also in samples from Italy.

Ear bearded, simple ; glumes white, pubescent ; awns white ; grain red.

T. turgidum, var. buccale, Korn. Handb. d. Getr. i. 60 (1885).

Kornicke states that var. buccale is cultivated in Germany, England, France,
Spain, and Italy ; I have also received it from Portugal.

The Cone Rivet or Anti-fly wheat formerly grown in England was a form of
this variety.

1. Pombinho. — Received from Portugal.

Young shoots, semi-erect ; young leaves pubescent.

Straw, tall, 132 cm. (about 52 inches) high ; upper internode hollow with
thick walls.

Ear, 7-9 cm. long, square, 12-14 mm. across the sides, or slightly compressed,
10 mm. across the face and 12-13 mm. across the side ; spikelets 21-25, 3- to 4-
grained ; D = 28-30 ; awns stout, 12-15 cm. long (Ear type 2, Fig. 157).

Empty glume, 8 mm. long, apex narrow, apical tooth acute, i mm. long
(Form 6, Fig. 152).

Grain, flinty or semi-flinty, dark red, truncate ; dorsal ridge prominent ; 7-3
mm. long, 3-6 mm. broad, 3-6 mm. thick.

2. Sicilio, with very villose glumes and short starchy grains, also from
Portugal, closely resembles (i).

FIG. 157.— RIVET WHEAT (T. turgidum, L.).

i. var. nigrobarbatum.
(Phillipino.)

2. var. Mertemii.
(Moula Oglau.)

RIVET OR CONE WHEAT 253

Ear bearded, branched ; glumes white, pubescent ; awns white ; grain red.

T. turgidum, var. centigranum, Korn. Handb. d. Getr. i. 63 (1885).
The form of var. buccale with compound ears, the result of a cross and not
constant (Kornicke).

Ear bearded, branched ; glumes white, pubescent ; awns white ;
grain reddish-violet.

T. turgidum, var. modigenitum, Korn. Arch. f. Biontologie, ii. 403
(1908).

A form originating from grains of T. durum, var. Schimperi, apparently the
product of a cross with a white, villose, branched ear of T. turgidum (Kornicke).

Ear bearded, simple ; glumes white, pubescent ; awns black ; grain white.

T. turgidum, var. Salomonis, Korn. Handb. d. Getr. i. 61 (1885).

A comparatively rare variety, the type of which was obtained by Kornicke
from a sample of " Grano tenere bianco," sent by G. Salomone from Catania,
Sicily, to the Vienna Exhibition in 1873.

A short dense-eared wheat from Baluchistan has been referred to this variety
by Howard.

Ear bearded, simple ; glumes red, glabrous ; awns red ; grain white.

T. turgidum Dreischianum, Korn. Handb. d. Getr. i. 60 (1885).

Kornicke's type was sent to him under the name " Frumento bianco " by
Dr. Dreisch from the Italian Exhibition held in Paris in 1878. This variety is
chiefly confined to the Balkan area, most examples being received from Greece,
Bulgaria, and Turkey.

A winter form from Greece, Bulgaria, and Turkey.

Young shoots, prostrate or semi-erect ; young leaves pubescent.

Straw, tall, 130 cm. (about 52 inches) high, hollow with thick walls.

Ear, 9-11 cm. long, lax, narrow, square, 10-11 mm. across the sides ; spike-
lets 22-24, somewhat elongated, 2- to 3-grained ; 0 = 22-24 '•> awns about 13-14
cm. long, parallel to the sides of the ear (Ear type 2, Fig. 156).

Empty glume, 9 mm. long, apex narrow, apical tooth very short and blunt (3,
6, Fig. 152).

Grain, mealy, large, apex truncate, dorsal ridge prominent ; 8-5 mm. long,
4-2-4-4 mm. broad, 3-65 mm. thick.

Ear bearded, branched ; glumes red, glabrous ; awns red ; grain white.

T. turgidum, var. pseudocervinum, Korn. Handb. d. Getr. i. 63 (1885).

This is one of the most frequently cultivated varieties with compound ears,
and may be considered as var. Dreischianum with a branched rachis.

I have received examples from Turkey under the name Keupely, from
Idaho and other parts of the United States as Alaska, Seven-headed, and
Miracle wheat, and from Greece.

254 THE WHEAT PLANT

Young shoots, semi-erect ; young leaves pubescent.

Straw, tall, 125-130 cm. (about 50 inches) high, hollow with thick walls. The
plants tiller very little, each usually sending up not more than two straws.

Ears, 8-9 cm. long, varying in width according to the amount of branching.

In a typical ear (i, Fig. 160), which is generally less compound than var.
mirabile (p. 256), the upper third is normal, bearing single spikelets at each
notch of the rachis. At each notch of the middle third of the ear two spikelets
arise, side by side, arranged at right angles to the spikelets above ; the notches
of the lower part of the ear bear short secondary ears, 2-2-5 cm- l°ng> usually
composed of 3-7 spikelets. A well-grown ear possesses 60-70 spikelets, yielding
90-100 small grains.

Empty glume, pale red, 6 mm. long ; apical tooth very short and blunt ; awns
of the flowering glume 2-5-7 cm. long (9, Fig. 152).

Grain, small, semi-flinty, laterally compressed, with narrow apex ; 6-7 mm.
long, 3-5 mm. broad, 3-55 mm. thick.

A form of pseudocervinum is recorded by Kornicke and Werner from Castile
Spain, under the name Trigo Rubio ; they state that it is sometimes cultivated
in Southern Europe and Northern Africa.

Ear bearded, simple ; glumes red, glabrous ; awns red ; grain red.

T. turgidum, var. speciosum, Kdrn. Handb. d. Getr. i. 60 (1885).

A fairly common variety cultivated in Italy and France. Flaksberger
records its occurrence from the Tiflis and Baku Provinces of Transcaucasia,
and notes that pure lines of the typical red-awned variety sometimes give black-
awned forms (var. Mertensii).

1 . A winter form from Spain.

Young shoots, prostrate or semi-erect ; young leaves pubescent ; hairs short.

Straw, tall, 122 cm. (about 48 inches) high ; upper internode solid.

Ear, lax, 9-10 cm. long, compressed, the face broader than the side as in
T. vulgare, 12-15 mm. across the face, 10-11 mm. across the 2-rowed side;
spikelets 21-23, broad, 3- to 4-grained ; 0 = 23-25 ; awns 15 cm. long.

Empty glume, 9 mm. long, narrow, apical tooth acute.

Grain, flinty, narrow, dorsal ridge prominent ; 7-8-7-9 mm. long, 3-7-3-9 mm.
broad, 3-8-4 mm. thick.

2. Mazzochio. — A form grown extensively in Italy (Tuscany).
Young shoots, semi-erect ; young leaves pubescent.

Straw, very tall, somewhat slender, 157 cm. (about 62 inches) high; upper
internode solid.

Ear, 8-9 cm. long, square, 11-13 mm. across the sides, or slightly compressed,
10 mm. across the face and 12-14 mm. across the side ; spikelets 22-25, 2" to 3"
grained ; D = about 30 ; awns stout, 12-15 cm. long, deciduous (Ear types i, 2,

Fig- 157).

Empty glume, 6-7 mm. long, apical tooth very short, blunt (3, 6, Fig. 152).

Grain, mealy, large, yellowish-red, dorsal ridge prominent ; 8-2 mm. long,
4-5 mm. broad, 4 mm. thick.

FIG. 158. — RIVET WHEAT (T. turgidum, L.).

i . var. melanatherum.
(Carosellone del Molise.)

2. var. gentile.
(Poulard a six rangs.)

RIVET OR CONE WHEAT 255

Poulard rouge de Gatinais, cultivated in the Central Departments of
France, is a form of speciosum nearly allied to this.

3. Buca vera. — A winter form received from Haage and Schmidt, Erfurt,
Germany.

Young shoots, prostrate ; young leaves pubescent.

Straw, tall, 130 cm. (about 51 inches) high ; upper internode hollow with
very thick walls.

Ear, 9-10 cm. long, square, 12 mm. across the sides ; spikelets short and
wide, 22-25, 3~ to 4-grained ; D = about 30; awns slender, 10-11 cm. long,
deciduous (Ear type i, Fig. 156).

Empty glume, 7 mm. long, often with a dark line along the outer margin ;
apical tooth acute, curved, with a broad base, 1-5 mm. long (Form 4, Fig. 152).

Grain, meah , dorsal ridge not prominent ; 7-6 mm. long, 3-75 mm. broad,
3 -4 mm. thick.

Ear bearded, branched ; glumes red, glabrous ; awns red ; grain red.

T. turgidum, var. cervinum, Korn. Handb. d. Getr. i. 63 (1885).
A rare form derived from a cross.

Ear bearded, simple ; glumes red, glabrous ; awns black ; grain white.

T. turgidum, var. speciosissimum, Korn. Handb. d. Getr. i. 60 (1885).

Kornicke obtained his type from a mixed sample of wheat designated
" Frumento veneto " at the Italian Exhibition held in Paris in 1878. According
to Flaksberger, this variety is cultivated in considerable amount in Transcaucasia.
I received samples from Turkey.

A winter form received from Dr. Regel, Petrograd, from the district of
Tiflis, Transcaucasia, and in a sample from Turkey under the name Moula
Oglau (var. Mertensii).

Young shoots, prostrate ; young leaves pubescent.

Straw, very tall, 150 cm. (about 60 inches) high ; upper internode hollow with
thick walls.

Ear, 8-10 cm. long, compressed, 9-11 mm. across the face, 13-14 mm.
across the side ; spikelets 23-27, 2- tO3-grained ; 0 = 29-31 ; rachilla with con-
spicuous fulvous hairs ; awns 10-13 cm- l°ng> Jet black, parallel to the sides of
the ears (Ear type i, Fig. 157).

Empty glume, 7-5-8 mm. long, apical tooth short, blunt (6, 7, Fig. 152).

Grain, mealy, short, apex truncate, dorsal ridge prominent ; about 7 mm.
long, 4 mm. broad, 3-8 mm. thick.

Ear bearded, branched ; glumes red, glabrous ; awns black ; grain white.

T. turgidum, var. Plinianum, Korn. Handb. d. Getr. i. 63 (1885).
The form of var. speciosissimum with compound ears. It is recorded by
Flaksberger from Turkestan.

256 THE WHEAT PLANT

Ear bearded, simple ; glumes red, glabrous ; awns black ; grain red.

T. turgidum, var. Mertensii, Korn. Handb. d. Getr. i. 60 (1885).

Kornicke's type in the Berlin Herbarium has a dense square ear 7 cm.
long and awns 15 cm. long. It was collected by G. Mertens in 1818 at " Vetrego
nella campagna di Gottardo " as " Frumento faro o vicentin."

A comparatively uncommon variety chiefly found in the Balkan area. I
have received samples from Turkey and Greece, and Flaksberger records the
occurrence of dense-eared winter forms of this variety in Transcaucasia.

Moula Oglau. — A winter form received from Turkey and a similar form
from Greece.

Young shoots, prostrate ; young leaves pubescent.

Straw, tall, 140 cm. (about 55 inches) high ; upper internode hollow with
thick walls.

Ear, 8-10 cm. long, compressed, 10 mm. across the face, 13 mm. across the
side ; spikelets 22-26, 2- to 3-grained ; D= 28-30 ; awns 9-10 cm. long, upper
part red, lower part black (Ear type i, Fig. 157).

Empty glume, 8 mm. long, apical tooth blunt (3, 6, Fig. 152).

Grain, mealy, dorsal ridge not very prominent ; 7-8 mm. long, 4 mm. broad,
3-8 mm. thick.

Ear bearded, simple ; glume red, pubescent ; awn red ; grain white.

T. turgidum, var. pseudomirabile, mihi.
A rare variety received only from Spain.

Trigo Salmeron. — A form with long, narrow, very lax ears, which when
young are a yellow-green tint.

Young shoots, erect or semi-erect ; young leaves with short hairs.

Straw, stout, tall, 120-135 cm. (48-53 inches) long ; upper internode solid.

Ear, long, lax, and narrow, 10-12 cm. long, square, 10-11 mm. across the
sides ; spikelets 22-27, 3-grained ; 0 = 22-24 5 awns 12-14 mm. long, scabrid
to the base (Ear type 2, Fig. 159).

Empty glume, short, somewhat inflated, 8-9 mm. long ; apical tooth short
and curved (18, Fig. 138).

Grain, white, opaque, with prominent dorsal hump ; 8-2-8-9 mm. long, 3-8
mm. broad, 3-8 mm. thick.

Ear bearded, branched ; glumes red, pubescent ; awns red ; grain white.

T. turgidum, var. mirabile, Korn. Handb. d. Getr. i. 63 (1885).

This is the most commonly grown variety having compound ears, being
met with in the British Isles, France, Germany, Italy, Spain, Portugal, and the
North African coast. In most instances it is cultivated on a small scale more as
a curiosity than a profitable farm crop.

Miracle or Mummy Wheat (2, Fig. 160). — The commonest branched form
of T. turgidum. Its massive ear suggests a high yield, but in field culture the
return per acre from it is low except on rich soil in a warm climate.

FIG. 159. — RIVET WHEAT (T. turgidum, L.).

i. var. dinurum. 2. var. rubro-atrum.

(Red Rivet.)

RIVET OR CONE WHEAT 257

Young shoots, semi-erect ; young leaves almost smooth, the hairs on the ridge
being reduced in this form to short projections from the epidermis.

Straw, tall, 125-130 cm. (about 50 inches) high ; upper internode solid.

Ear, compressed, 7-9 cm. long, and 4-5 cm. across the widest part. The
branching of the ear is hereditary, but the extent of its manifestation is influenced
by soil conditions and season and by the space allotted to the plants.

In the simpler, least compound ears the upper part is normal, with a single
spikelet at each notch of the rachis, a few of the lower notches producing a pair
of spikelets placed side by side and arranged at right angles to those of the
normally placed single spikelets above. In large well-developed ears the upper
third is usually normal ; below this, one or two notches bear pairs of spikelets ;
at the remaining lower notches secondary ears are produced, their spikelets
being arranged at right angles to those on the apical portion of the main axis.
The secondary ears vary in number, length, and development ; ten or twelve
are often found on well-grown ears, the largest 3-4 cm. long with 10-14 small
crowded spikelets, whose grains often protrude from between the short glumes ;
the basal secondary ears are often rudimentary, possessing 4-10 diminutive
spikelets, usually barren or nearly so.

A large well-developed ear frequently has a total of 115-120 spikelets and
140-150 grains, one of average size 85-90 spikelets and 80-100 grains ; the wheat
is nevertheless not prolific in field culture, the tillering power being low, each
plant rarely producing more than two straws.

Empty glume, 7 mm. long, reddish, with a glaucous covering and a dark line
often along the outer margin (Forms 4, 6, Fig. 152). The awns of the flowering
glumes are short and slender, 4-9 cm. long, sometimes black at the base.

Grain, mealy, small, with prominent dorsal ridge and blunt apex ; 6-5 mm.
long, 3-8 mm. broad, 3-8 mm. thick.

Ear bearded, simple ; glumes red, pubescent ; awns red ; grain red.

T. turgidum, var. dinurum, Korn. Handb. d. Getr. i. 63 (1885).

This is perhaps the most widely cultivated variety of T. turgidum, some forms
being comparatively hardy and adapted to a great range of climate. It is grown
in England, France, Germany, Italy, Spain, Portugal, Bulgaria, Greece, and
Turkey ; apparently not in Russia. The young leaves of the majority of the
representatives of this variety are almost glabrous, the hairs being very short,
from 32 to 64 // long, those of typical turgidum being 120-250 p. long.

i. Trigo focense. — Received from Spain. (I have had forms of T. vulgare,
var. erythrospermum, from Spain under the name Trigo focense.)

Young shoots, semi-erect ; young leaves almost glabrous.

Straw, very tall, 150 cm. (about 60 inches) high ; upper internode hollow
with thick walls.

Ear, lax, 10-12 cm. long, vulgare~\ike, 12-15 mm. across the face, 10-11 mm.
across the 2-ranked side ; spikelets 26-30, wide, 3- to 4-grained ; 0 = 26-27 ;
awns 12-13 cm. long, deciduous (Ear type i, Fig. 156).

Empty glume, 8 mm. long, apical tooth acute, 1-5 mm. long (4, Fig. 152).

s

258 THE WHEAT PLANT

Grain, flinty, large, dorsal ridge prominent, apex truncate ; 7-5 mm. long,
3-7 mm. broad, 3-7 mm. thick.

2. Nonette de Lausanne. — Received from France ; cultivated in Italy under
the name Andriolo rosso pelosa.

A winter form grown formerly in England under the name Giant St.
Helena wheat. It is a prolific wheat, fairly resistant to frost and grows satis-
factorily on comparatively poor strong soils.

Young shoots, semi-erect ; young leaves almost glabrous.

Straw, very tall, stout, 152 cm. (about 60 inches) high ; upper internode hollow.

Ear, of medium density, pendent when ripe, 8-9-5 cm. long, square, 12-14
mm. across the sides; spikelets 27-31, 3- to 4-grained ; 0 = 29-32; awns 9-10
cm. long (Ear type i, Fig. 156).

Empty glume, 8 mm. long, with acute apical tooth, 1-5-2 mm. long, and dis-
tinct lateral tooth (Forms i, 4, Fig. 152).

Grain, more or less flinty, long and somewhat narrow with indistinct dorsal
ridge ; 8-3 mm. long, 3-9 mm. broad, and 3-6 mm. thick. Closely similar forms
were received from Bulgaria and Greece and from the United States under the
name Ratel, and as Gigante Milanese from Italy, France, and Spain.

3. A form received from Turkey under the name Hudavendigar resembles
the preceding, but has narrower, square ears 9-11 mm. across the sides, divergent
awns, and smaller semi-flinty grains narrowed at the apex, with a prominent
dorsal ridge ; 6-8-7 mm. long, 3-5-3-7 mm. broad, 3-5-3-8 mm. thick.

4. Bed Eivet ; Clock Wheat.

Young shoots, semi-erect ; young leaves almost glabrous.

Straw, tall, stout, 130-132 cm. (about 52 inches) high ; upper internode
hollow with thick walls.

Ears, dense, 7-5-9 cm. long, square, 12-14 mm. across the sides ; spikelets
28-36, 3- to 4-grained ; D = 38-40 ; awns divergent, 8-9 cm. long, pale red (Ear
type i, Fig. 159).

Empty glume, 6-7 mm. long, apex narrow ; apical tooth narrow, acute, 1-5
mm. long (4, Fig. 152).

Grain, starchy, laterally compressed, dorsal ridge prominent ; 7 mm. long,
3-45 mm. wide, 3-65 mm. thick.

The term Rivet is obscure ; its first recorded use is by Tusser in his Fine
hundred pointes of good Husbandrie (ed. 1580 ; not in the first ed. 1573).

White wheat or else red, red riuet or whight,
far passeth all other for land that is light,

White pollard or red, that so richly is set :
for land that is heauie is best ye can get.

So Turkey or Purkey wheat, many do loue :
because it is flourie as others aboue.

This wheat is often erroneously termed " Rivett's wheat," but the word
rivet is correctly used in the plural — rivets — in a generic sense for this and other
varieties of turgidum or samples of their grain. Red Rivet wheat also had
various other names such as Pendule or Pendulum wheat (doubtless on account

FIG. 160. — RIVET WHEAT (T. turgidum, L.).

i. var. pseudocervinum.
(Alaska or Seven-headed.)

2. var. mirabile.
(Miracle or Mummy.)

RIVET OR CONE WHEAT 259

of its drooping ears), Red Cone or Eed Pollard wheat, though Tusser implies
that Red Pollard was different from Red Rivet ; White Rivet is also mentioned
by Worlidge (1681).

Neither this nor the previously mentioned forms of var. dinurum are much
grown in England at the present day, their place being taken by forms of var.
iodurum, to which the generic term Rivet is frequently applied.

A sample of wheat received from Germany under the name Payne's De-
fiance consisted of Red Rivet and Blue Cone (see p. 260).

Ear bearded, branched ; glumes red, pubescent ; awns red ; grain red.

T. turgidum, var. Linnaeanum, Korn. Handb. d. Getr. i. 63 (1885).
The rare form of var. dinurum with compound ears.

Ear bearded, simple ; glumes red, pubescent ; awns black ; grain red.

T. turgidum, var. rubroatrum, Korn. Handb. d. Getr. i. 61 (1885).

A rare variety found by Kornicke among other wheats from Italy. I
received a narrow lax-eared form of this variety from Spain ; some of its
characters suggest a relationship to T. vulgar e.

Young shoots, erect or semi-erect ; young leaves slightly hairy.

Straw, of medium height, 90-100 cm. (38-40 inches) high ; upper internode
solid.

Ear, long, lax, and narrow, 10-11 cm. long, square, 10-11 mm. across the
sides ; spikelets 21-23, 3-grained ; D = 22 ; awns 12 cm. long, scabrid to the
base (Ear type 2, Fig. 159).

Empty glume, short, somewhat inflated, 8 mm. long ; apical tooth short
(Forms 2, 5, Fig. 152).

Grain, pale red, flinty, short, apex truncate, dorsal ridge prominent ; 7-6-7-9
mm. long, 3-6-3-9 mm. thick.

Ear bearded, simple ; glumes blue-black, glabrous ; awns black ; grain white.

T. turgidum, var. Herrerae, Korn. Handb. d. Getr. i. 60 (1885).

A rare variety the type of which was obtained by Kornicke from Valencia,
Spain.

According to Vasilieff, it is grown in the Tiflis Province of Transcaucasia.

The Ble Garagnon noir, which Heuze says is cultivated in Algeria and
occasionally in the olive regions of France, appears to be a form of var. Herrerae.

Ear bearded, simple ; glumes blue-black, pubescent ; grain red.

T. turgidum, var. iodurum, Korn. Handb. d. Getr. i. 61 (1885).

In some forms of this variety the colour of the glume is a dense blue-black ;
in others, especially those with dense ears, it is a pale bluish-grey.

The ground tint in most cases is reddish, and when this tint preponderates,
as it often does in damp seasons, the forms are difficult to separate from those
of var. dinurum. The empty glumes of the pale forms of the var. iodurum,

a6o THE WHEAT PLANT

however, usually have a dark outer margin. Like var. dinurum this is one of the
most extensively cultivated varieties of T. turgidum, being adapted to a compara-
tively wide range of climate. It is grown in England, France, Germany, Italy,
Spain, and Portugal, and occasionally in Australia and the United States of
America. Flaksberger says it is not found in Russia.

1. Pisana francesca. — A distinct winter form received from Spain and
Portugal ; the flat-faced, narrow-sided ear suggests a relationship to T. vulgar e.

Young shoots, prostrate ; young leaves pubescent.

Straw, tall to very tall, 130-150 cm. (about 50-60 inches) high, hollow with
thick walls.

Ear, 8-9 cm. long, 13-15 mm. across the face, 10-12 mm. across the side ;
spikelets broad, 22-24, 3" *° 4-grained ; D = 24-27 ; awns divergent, 9-10 cm.
long, deciduous (Ear type, Fig. 155).

Empty glume, short, inflated, blue-grey on a red ground, 6 mm. long ; apical
tooth curved, acute, i mm. long (4, Fig. 152).

Grain, mealy, pale yellowish-red, large, with prominent dorsal ridge ; 7-5 mm.
long, 4 mm. broad, 3-9 mm. thick.

Pole-rivet, formerly cultivated in England, appears to have been allied to
this form.

2. Pe'tianelle noire de Nice. — A spring form received from France.
Young shoots, erect ; young leaves pubescent.

Straw, very tall, 140 cm. (about 56 inches) high, striate ; upper internode solid.

Ear, lax, 11-15 cm- l°ng> square, narrow, 10-12 mm. across the sides ;
spikelets 26-30, 3- to 4-grained ; 0=22-27 '•> awns 12-14 cm. long, deciduous
when ripe (Ear types i, 2, Fig. 156).

Empty glume, 7 mm. long, blue-black with a glaucous surface ; apical tooth
acute, broad at the base, about i mm. long (Forms i, 4, Fig. 152).

Exposed parts of the flowering glume blue-black, the parts covered by the
empty glumes pale yellowish-red.

Grain, more or less flinty, large, not always plump ; 7-5 mm. long, 3-5 mm.
broad, 3-8 mm. thick.

A distinct and striking form only suited to a warm climate.

Forte Nero from Italy and an unnamed form from Spain closely resemble
Petianelle noire de Nice, and "Andriolo nero " and Grano nero are synonym-
ous Italian forms.

Very similar also is Reynold's Discovery received from Australia, with
hollow straw and large flinty grain ; 8-2 mm. long, 3-9 mm. broad, 3-9 mm. thick.

3. Blue Cone. — A winter form, a select line of which, introduced by the
author, is grown in the southern half of England.

A similar form under the name Poulard d'Australie or Poulard bleu is
grown in the north of France.

Young shoots, prostrate ; young leaves pubescent ; blades narrow.

Strazu, very tall, 150 cm. (about 58 inches) high, striate ; the upper internode
hollow with thick walls.

Ear, pendulous when ripe, 9-10 cm. long, tapering a little towards the apex,

RIVET OR CONE WHEAT 261

square, 12-14 mm. across the sides ; spikelets 26-30, well filled, 3- to 4-grained ;
0 = 30-32 ; awns 10-11 cm. long, deciduous (Ear type i, Fig. 156).

Empty glume, 8 mm. long, blue-black on a reddish ground ; in damp seasons
the ear is pale ashy-brown or " mouse coloured " ; apical tooth 1-1-5 mm- l°ng
(4, Fig. 152).

Grain, starchy, yellowish-red, plump, apex truncate, dorsal ridge prominent ;
7-5-8 mm. long, 3-7 mm. broad, 3-65 mm. thick.

The term " Cone " applied to this and similar forms of T. turgidum probably
refers to the slightly tapering conical ear, which results when the lower spikelets
develop more and plumper grains than the upper.

Long Cone and White Cone wheats are mentioned by Plot in his Natural
History of Oxfordshire in 1677, along with a Eed Cone form called Pendule
wheat.

Grey Cone or Grey Pollard, also called Dugdale and Duckbill wheat,
grown frequently in the seventeenth and eighteenth centuries in England,
was a form with shorter ears and reddish-grey glumes. The specimen labelled
" Grey Pollard " in Petiver's Herbarium (British Museum) is about 7 cm. long,
one of " Cone wheat " in the same collection being 8-5 cm. long.

A sample of Payne's Defiance from Germany consisted of Blue Cone and
Red Rivet (var. dinurum).

Ear bearded, branched ; glumes blue-black, pubescent ; grain red.

T. turgidum, var. coeleste. Handb. d. Getr. i. 64 (1885).

From the cross T. dicoccum, var. cladurum x T. turgidum, var. iodurum,
Kornicke obtained this variety and a similar beardless unbranched variety (var.
coelestoides], neither of which was constant.

Ear beardless, simple ; glumes blue-black, pubescent ; grain red.

T. turgidum, var. subiodurum, Korn. Arch.f. Biontologie, ii. 401 (1908).

The product of a cross obtained by Gustav Bestehorn. Kornicke states

that it is constant, and differs only from var. iodurum in possessing beardless ears.

CHAPTER XVIII

EGYPTIAN CONE WHEAT

Triticum pyramidale, mihi.

A SMALL race which I have seen only from Egypt. It has some of the
characters of T. turgidum, but, unlike this, has short straw, characteristic
yellow-green culm leaves, pointed grain, and is among the earliest wheats,
coming into ear at Reading at the end of May or the first few days in June.
The Dwarf wheats collected by W. Schimper in Abyssinia and referred
to T. compactum by Kornicke, doubtless belong to this race.

GENERAL CHARACTERS OF T. pyramidale, mihi.

The young plants have erect shoots ; the leaf-surface is clothed with
soft hairs of equal length, like those of T. dicoccum and T. turgidum.

The straw is very short, rarely more than 80-100 cm. (32-40 inches)
high, solid or hollow with thick walls.

The culm leaves are generally a yellow-green tint. The ears, which

usually taper towards the apex, are
short and very dense, rarely meas-
uring more than 5 or 6 cm., and
having a density usually over 40 ;
spikelets 20-23.

The edges of the rachis are
fringed with conspicuous white
hairs, and there is a frontal tuft at
the base of each spikelet.

3 The spikelets are about 12 mm.

FIG. 161. — Empty glumes of Egyptian Cone long and 12 mm. wide, and often

ripen 3 or 4 grains in each.

The empty glumes are strongly keeled from the tip to the base ; the
apical tooth in some forms is long and acute, in others short and blunt
(Fig. 161).

The flowering glume bears a long, somewhat slender awn from 9 to 15
cm. in length, which is scabrid to the base.

262

EGYPTIAN CONE WHEAT

263

The grains are a fine white tint, generally mealy, laterally compressed,
narrowed at the apex, with a prominent dorsal ridge or hump ; they are
7*5-8-5 mm. long, 3-3 mm. broad, and 3'5-3'6 mm. thick (Fig. 162).

VARIETIES OF T. pyramidale, mihi.

var. recognitum, mihi.
var. compressum, mihi.

1. Ear bearded ; glumes white, glabrous ; awns white

2. Ear bearded ; glumes white, pubescent ; awns white

3 . Ear bearded ; glumes white, pubescent ; awns black .

var. pseudo-compressum, mihi.

4. Ear bearded ; glumes pale red, pubescent ; awns

red .... var. copticum, mihi.

5. Ear bearded ; glumes pale red, pubescent ; awns

black . . . var. pseudo-copticum, mihi.

Ear bearded ; glumes white, glabrous ; awns white ;
grain white.

T. pyramidale, var. recognitum, mihi.

White Saidi (i, Fig. 163). — An early wheat received
from Upper Egypt.

This is a very distinct form, referred by Seringe
to T. durum complanatum and described by Delisle
(Descr. de I'Egypte, ii. 177) as T. sativum pyramidale.
Kornicke's T. compactum, var. recognitum, is probably
this form.

In its habit and the morphological characters of its
leaves, it is quite unlike T. durum, T. sativum, or T.
compactum.

Young shoots, erect ; young leaves pubescent, the
hairs slightly shorter in this than in most varieties of
T. dicoccum.

Straw, short, 80-85 cm- (about 33 inches) high ;
upper internode hollow with thick pithy walls.

Ear, short and very dense, 4-5-5 cm. long, 10-11
mm. across the face, 15-18 mm. across the side ;
spikelets 20-21, densely crowded on the rachis, many
with three grains in each; 0=45-50. Awns about
ii cm. long, scabrid to the base. The ear is flattish on one side and convex on
the other. It tapers from the base towards the apex, which in the larger ears is
often curved to one side.

Empty glume, 8 mm. long ; apical tooth broad and blunt, i mm. long
(3, Fig. 161).

Grain, mealy, dorsal hump very prominent ; 7-5 mm. long, 3-3 mm. broad,
3-65 mm. thick.

FIG. 162. — Grains of the
spikelets of one side of
an ear of Egyptian Cone
wheat (T. pyramidale)
(nat. size).

264 THE WHEAT PLANT

Ear bearded ; glumes white, pubescent ; awns white ; grain white.

T. pyramidale, var. compressum, mihi.

This variety I obtained from Egypt mixed with the succeeding variety, from
which it differs only in the colour of the awn.

Ear bearded ; glumes white, pubescent ; awns black ; grain white.

T. pyramidale, var. pseudo-compressum, mihi.

A form of this variety I received from Egypt under the names Tunis wheat
and Fayum wheat.

It is an early, short-strawed form, with the characteristic dense ear of the
race.

Kornicke's T. compactum, var. compressum, appears to be the same.

Young shoots, erect ; young leaves pubescent.

Straw, short, rigid, 56-80 cm. (22-32 inches) long ; upper internode solid
hollow with thick walls.

Ear, short and dense, 4-6 cm. long, oblong in section, 10-11 mm. across the
face, 15-18 mm. across the 2-rowed side; spikelets 21-23, 3 -grained ; D =

4°-43-

Empty glume, 9-10 mm. long, apex narrow, apical tooth acute (2, Fig. 161);
awn of flowering glume 10-13 mm. long.

Grain, white, flinty or semi-flinty, narrow at the apex, laterally compressed,
with a prominent dorsal ridge ; 8-8-4 mm- l°ng> 3*° mm. broad, 3-5-3-6 mm.
thick.

Ear bearded ; glumes pale red, pubescent ; awns red ; grain white.
T. pyramidale, var. copticum, mihi.

Ein el Bent (Eye of the Girl) (2, Fig. 163). — A distinct form received from
Egypt, and probably the same as Kornicke's T. compactum, var. copticum, from
Abyssinia.

Young shoots, erect ; young leaves pubescent.

Straw, short to medium height, 76-105 cm. (about 30-42 inches) high,
glaucous ; upper internode hollow with thick walls.

Ear, yellow-green, short and compressed, dense, 5-8 cm. long, 12 mm.
across the face, 20 mm. across the two-ranked side ; spikelets 20-25 ; D = about
36 ; awns 14-17 cm. long, scabrid to the base.

The ears usually taper from the base to the apex and are often curved towards
the flat side near the tip.

Empty glume, 10 mm. long, pale red, pubescent, with prominent lateral nerve ;
keel tooth broad and bluntish (i, Fig. 161).

Grain, white, semi-flinty, laterally compressed, with prominent dorsal hump ;
6-5-7-5 mm. long, 2-8-3-2 mm. broad, 3-3-3-6 mm. thick.

Ear bearded ; glumes pale red, pubescent ; awns black ; grain white.

T. pyramidale, var. pseudo-copticum, mihi.

Received with the preceding variety, from which it differs only in the posses-
sion of black awns.

FIG. 163. — EGYPTIAN CONE WHEAT (T. pyramtdale, mihi).

i. var. recognitum.
(White Saidi.)

2. var. copticum.
(Ein el Bent.)

CHAPTER XIX

COMMON BREAD WHEAT

Triticum vulgare, Host. Icon, et descr. Gram. Aust. iii. 18 (1805).
T. aestivum, L. (bearded). \ e P7 Q Qf- /T_r.\
r. *«•»•«;, L. (beardless). I *' P/' ®5' 8& ('753)'
7\ sativum, Lamk. Enry. Me*A. ii. 554 (1786).

(T. aestivum, L.
= - T. hybernum, L.

{T. turgidum, L.

T. vulgare, Vill. ( = T. aestivum, L.). #w*. P7. </. Dauph. ii. 153 (1787).
T. sativum, Pers. 5jyw. P/. i. 109 (1805) :

a. aestivum. (3. hybernum. \ j. durum.

ALTHOUGH bread can be made from Macaroni (T. durum), Rivet (T.
turgidum), and other wheats, it is from Triticum vulgare that the world's
bread is chiefly produced. The peculiar physical quality of the gluten of its
grain specially fits it for the manufacture of spongy, digestible bread,
and in this respect it surpasses all other races. In addition, on account
of its great adaptability to a variety of climatic conditions, it is the most
widely distributed of all. More of this race is grown than of any other,
some forms of it being found in every country wherever wheat of any kind
is cultivated.

Bread wheat is one of the most ancient of cereals, examples of ears of
grain in considerable abundance having been discovered in various parts
of Europe on sites occupied by man in Neolithic times and the Bronze and
Iron Ages.

In the earliest periods the grains were distinctly smaller than the
forms now cultivated, but all grades up to the large plump grains near
to those of T. turgidum are met with in the later deposits.

The most primitive of prehistoric wheats of this race appears to be a
variety hitherto found only in the Neolithic store-chambers at Lengyel
(Hungary) and described by Deininger under the name T. sativum Scythi-
cum. In this form the apex of the grain tapers to a narrow point, the
whole being pear-shaped, with little or no sign of a furrow on the ventral

265

266 THE WHEAT PLANT

side and more or less circular in transverse section. The grains measure
3-5-4-5 mm. long, 2-3 mm. broad, and 1-8-2-6 mm. thick.

Deininger infers that the spikelets of this wheat probably bore only
one grain, and concludes that this form is the prototype of the Bread
wheats and a form found more widely distributed throughout Europe
in the Neolithic period named T. vulgar e antiquo/nan by Heer. The ears
of the latter are beardless, short, and dense (4-5 cm. long and 10 mm.
wide), with three or four grains in each spikelet ; the empty glumes are
strongly keeled throughout their length with an inwardly curved apical
tooth. The grain is especially small and blunt at the apex, and markedly
convex on the dorsal side ; the average dimensions are : length 5 mm.,
breadth 3-5 mm. Kornicke and Buschan place this wheat under T.
compactum, a view in which I concur, the examples I have seen being most
like some Chinese compactums of the present day.

It is not until the Bronze and Iron Ages that grains similar in size and
form to those now cultivated became common in prehistoric deposits.

In Britain small-grained forms of T. vulgare, possibly mixed with those
of T. compactum, were cultivated in Roman times.

I have seen no examples which can be attributed with certainty to
this race from ancient Egypt, but the wheats described under the name
TTvpos by the Greeks and triticum by Roman authors were wheats with
grain loosely invested by the glumes, and doubtless included T. vulgare
as well as- T. durum.

GENERAL CHARACTERS OF T. vulgare

The young shoots are erect, semi-erect, or prostrate.

A number of forms from China, Japan, Persia, and India have yellowish-
green leaves ; the majority, however, have more
or less glaucous blades and ears.

On the surface of the young leaves hairs
are always present. A characteristic single line
of long hairs is found along the summit of all
the longitudinal ridges or upon those near the

leaf-margins ; in some cases this is the only
FIG. 164. — Diagrammatic ,. r , . , , . 11-^-

transverse sections of young line of hairs present, others have in addition

leaves of T. vulgare, T. a number of shorter hairs covering the flanks

compactum, 1 . sphaerococ-

cum, T. Spelta. of the ridges (rig. 164).

The culms vary in height from 75 cm.

(about 30 inches), or less in some Asiatic forms, to 140-150 cm. (55 or
60 inches). They possess 5 or 6 internodes, which are usually hollow,
with comparatively thin walls, but in a few forms {e.g. New Zealand
Tuscan) they are solid. The ears are generally grouped into bearded

COMMON BREAD WHEAT 267

and beardless forms ; a few Japanese and Australian forms are quite
awnless, but most of the so-called beardless wheats have short awns on the
flowering glumes of the upper spikelets of the ear, and in a few forms fine
short awns of uniform length are found on all the flowering glumes.

In most varieties the ears are flattened and broadest across the face ;
sometimes the width across the 2-ranked side and the face is the same,
but I have met with none in which the width of the 2-ranked side is
uniformly greater than that of the face, as in many forms of T. durum and
T. turgidum.

The length of the ear varies from 6 to 18 cm. (about 2-5-7 inches), with
an average of 20 spikelets per ear, the density varying from 14 or 15 to
36 or more per 10 cm. of rachis.

The rachis is tough and smooth, and fringed with short hairs along its
margins, with a few hairs immediately below the base of each spikelet.

The spikelets are 5- to Q-flowered ; near the middle of the ear they not
infrequently ripen 4 or 5 grains, but those near the base of the ear are
often barren or contain i or 2 grains only ; generally in the apical spikelets
only i or 2 grains are developed.

The spikelets are 10-15 mm. long, 9-18 mm. across the face, and 4-5
mm. thick. The empty glumes are glabrous, pubescent, or villose,
creamy-white, canary-yellow, or various shades of red, brown, or blue-
black ; those of the lateral spikelets are 6-n mm. long, unsymmetrical,
3-5 mm. from the midrib to the outer edge, terminating in a tooth, which
in the beardless forms is short and usually blunter than that of T. durum,
and in the bearded form often prolonged into a slender awn from i to 3 cm.
or more in length. The midrib is commonly prominent only in the upper
half, but in some cases there is a well-developed keel running from the
apex to the .base of the glume ; the various forms of the empty glume
and their apical teeth are illustrated in Figs. 165 and 166.

The flowering glumes are thin and pale, rounded on the back with
7-11 nerves. In bearded wheats the awns are from 5 to 10 cm. long, trique-
trous, scabrid, and persistent, and in some Chinese forms very fine, in
others, especially Persian and Indian sorts, stout and brittle. They are
generally of the same tint as the glume, namely, reddish- or yellowish- white,
black or dark brown awns being of exceptional rarity in T. vulgare, though
frequent in other races. The awns are usually straight or only slightly
curved, but in some Asiatic wheats they are tortuose or bent into the form
of a hook or loop (i, Fig. 179). A peculiar form is illustrated in Figs.
179 and 217.

The grains are white, yellow, pale orange, or varying shades of red or
brown, the colour being most readily determined in those with mealy,
opaque endosperm.

They are generally plump, somewhat broad and bluntish at the apex,

268

THE WHEAT PLANT

456

12 13 14 15 16 17 18 19 20 21 22

FIG. 165. — Empty glumes of bearded forms of Bread wheat (T. vulgare) ( x 2).

13 14 15 16 17 18 19 20 21 22 23 24 25

FIG. 166.— Empty glumes of beardless forms of Bread wheat (T. vulgare).

COMMON BREAD WHEAT

269

where there is a " brush " of hairs ; the furrow is usually shallow, and the
cheeks on each side of it are convex and plump (Fig. 167).

The endosperm is flinty, semi-flinty, or completely mealy.

Measurements of grains taken from the middle of the ear of sixty forms
gave the following results :

Length.

Breadth.

Thickness.

mm.

mm.

mm.

Average
Limits
Ratio ....

6-78

5-2-8-5

IOO

3'63
2-9-4-2

53'5

3^5
2-6-3-9

47'9

In number of its varieties and forms Bread wheat is the richest of the
races of wheat. More than 1300 have been collected and studied for many

DO 0 0 1 0 0 1U) * • t * 1 1 ft I M

> III *)ii IM IIHftt

FIG. 167. — Grains of Bread wheats (T. vulgare) ; front, back, and side views (nat. size).

years at Reading. These exhibit an extraordinary diversity of morpho-
logical characters, as well as differences in habit of growth and period of
ripening, and an almost unbroken series of transitional forms are found
connecting the most widely different varieties.

Long lax ears pass insensibly into the short dense Squareheads, and
in the form of the empty glume and the length of the awns of the flowering
glumes similar gradation is observed. Even between the Spring wheats
with erect young shoots and the prostrate Winter kinds forms are common
with young shoots which spread outwards at intermediate angles.

There is little doubt that T. vulgare is a vast collection of mutants
and hybrids, which I regard as having originated from the crossing of

270

THE WHEAT PLANT

T. dicoccoides or T. dicoccum with one or two species of Aegilops (see Chapter
XXIIL).

The natural classification and description of such a series presents
much the same problems and a similar degree of complexity as the classi-
fication of the human race, and nothing less than a minute description of
individuals would adequately represent it.

The artificial classification into varieties presents
no difficulty, and a few common forms of each
variety are described later.

Some broadly outlined natural groups are readily
recognised when the living plants are studied,
although the lines of demarcation are in some cases
obliterated by the normal fluctuating variations
of the different forms composing them. Among
such groups of apparently genetically related forms
are the following :

GROUP I. — The endemic forms of T. vulgar e
from India and also several from Persia and
Turkestan. These are all early, rapid-growing
Ifc^? {& forms, which come into ear at Reading about the

end of May or first week in June when sown in
autumn or early spring ; under favourable climatic
conditions many of them are able to produce ripe
grain in one hundred days or less from the time of
sowing.

The young shoots are erect, straw slender, in-
clined to curve towards the ground, in cool seasons
short, 76-90 cm. (30-36 inches) high, in hotter
years taller, leaves yellowish-green in the early, and
more or less glaucous in the later forms.

The ears are often quadrate, short, or of
medium length, from 6 to 10 cm. long, usually
lax and rigid, with a somewhat brittle rachis. The
density of " pedigree " lines in this group is more
variable than in any other, especially when com-
parison is made between plants sown in autumn and those of the
same line sown in spring.

The spikelets, 12-20, are sometimes arranged irregularly on the rachis,
especially in the case of lax ears ; in the denser ears they overlap each
other.

The empty glumes are rigid, scabrid, and not infrequently keeled to
near the base ; in the bearded forms the awns of the flowering glumes are
short (4-6 cm. long), scabrid, brittle, and divergent.

FIG. 168. — Grains of the
spikelets of one side of
an ear of Bread wheat
(T. vulgare) (nat. size).

FIG. 169.— BREAD WHEAT (T. vulgare, Host).

var. erythrospermum.
i. (Rosso gentile aristata.) 2. (Wagia, Bombay.)

COMMON BREAD WHEAT 271

The grains, usually 3 or 4 per spikelet, are flinty or semi-flinty, very
soft, mealy endosperm being uncommon ; they are held somewhat firmly
in the glumes.

All forms of this group are very susceptible to attacks of Yellow Rust,
which in England often damages the plants so much that few or no grains
attain their full development.

GROUP II. Endemic Persian and Central Asiatic Wheats. — The young
shoots are erect or semi-erect ; the straw of medium height, somewhat
thick and soft ; the leaves yellow-green or glaucous.

They are chiefly early or mid-season forms, with lax quadrate ears
and characteristic swollen spikelets.

The ears are stiff, 9-12 cm. long ; the awns of the bearded forms stout,
brittle, and scabrid ; spikelets 16-20, elongated, often irregularly arranged
on the rachis ; the grain large and generally soft (Figs. 174, 179).

GROUP III. — Early or mid-season forms, which show affinities with
bearded T. Spelta ; they are endemic in Persia, Bokhara, and Turkestan,
and I have met with examples among wheats from Portugal, Spain, and
Argentina (2, Fig. 173).

The young shoots are erect, the straw stiff, of medium height, leaves
glaucous or yellow-green.

The ears are 9-12 cm. long, narrow, quadrate, 8-10 mm. across the
face and side, lax, with narrow elongated spikelets, the empty glumes
rigid and frequently keeled to the base, lateral nerve prominent.

The awns of the flowering glumes 6-10 mm. long, scabrid.

The grain is long, narrow, semi-flinty, and firmly invested by the
glumes.

The rachis breaks readily, but is not so brittle as in the typical bearded
forms of T. Spelta.

GROUP IV. Endemic Japanese and Chinese Wheats. — Very early
wheats, which come into ear at Reading about the end of May or first week
of June. The young shoots are erect ; the straw somewhat short, 95-
115 cm. (38-46 inches) high, soft, hollow, and of large diameter; the
leaves in some forms yellow-green, in others more or less glaucous. The
ears are of medium length (9-12 cm. long), well filled from the base to the
apex, the spikelets 22-26, 5- to 7-flowered, frequently ripening 4 grains in
each : in several forms some of the ears of a plant are clubbed while the
rest are uniformly dense throughout (Fig. 189). The empty glumes
are frequently keeled to the base, the awns of the flowering glumes of
bearded forms slender and comparatively short (4-6 cm. long). The
glumes are thin and very easily separated from the rachis, and the grain
often visible between them and readily shed.

The grains are dark red, of fair milling quality.

272 THE WHEAT PLANT

Although the number in each ear is large, the spikelets even at the
base and apex being usually fertile, the total yield in weight is low as the
individual grains are small.

GROUP V. — In this group is included a large series of Spring wheats,
bearded forms of which are most prevalent. They are very extensively
cultivated in Spain, Portugal, Italy, the United States, Canada, and
Argentina, and are early forms, coming into ear at the end of May or first
ten days of June at Reading.

The young shoots are erect or semi-erect, and the straw usually of
medium height ; in some forms the leaves are yellowish-green, in others
glaucous.

The ears are lax and compressed, the face considerably wider than the
2-rowed side, the usual density 17-21, spikelets 2- to 3-grained, glumes
somewhat thin, the grain often visible between the glumes, generally
flinty and of good milling quality.

GROUP VI. — The wheats of this group are winter forms, late, coming
into ear at Reading during the second or third week in June. The young
shoots are prostrate, the young leaves narrow, and in many Russian forms
strikingly pubescent, the straw somewhat tall and slender. The ears
are similar to those of the previous group, lax and compressed, 9-13 cm.
long, usual density 17-21, but some forms have a density of 23-27 ; the
spikelets are broad, 2- to 3-grained, with thin glumes, which frequently
do not cover the grain.

The grain is generally flinty and of good milling quality.

The bearded forms of these wheats are very widely distributed in
Eastern Europe, and are grown also in the United States and Canada, and
occasionally in Holland and France and other countries of Western
Europe.

GROUP VII. The Squarehead Group (see p. 296). — In this are included
a number of Winter wheats grown chiefly in Western Europe, where
intensive cultivation is practised, and there is need of the highest yields
per acre in order to secure a remunerative return from the land.

The young shoots are prostrate or semi-prostrate, the straw short
and stiff, ears dense, of the Squarehead type (2, Fig. 189 ; i, 2, Fig. 190).

They have a long growing period and are very prolific. The grain is
usually soft and mealy and the flour of moderate baking quality.

The majority are beardless forms.

FIG. 170. — BREAD WHEAT (T. vulgare, Host).

i. var. erythrospermum.
(Rieti.)

2. var. erythrospermum.
(Preston.)

COMMON BREAD WHEAT 273

VARIETIES OF T. vulgar e
I. Ears bearded —

1. Glumes white, glabrous.

a. Grain white ..... var. graecum, Korn '

b. Grain red.

i. Awns white . . . var. erythrospermum, Korn. *-~

ii. Awns black . . . var. nigroaristatum, Flaksb .

2. Glumes white, pubescent.
a. Grain white.

i. Awns white .... var. meridionale, Korn.

ii. Awns black . . . . var. pseudomeridionale, Flaksb.
b. Grain red.

i. Awns white .... var. Hostianum, Korn.

ii. Awns black . . . var. pseudo-Hostianum, Flaksb.

3. Glumes red, glabrous.

a. Grain white ..... var. erythroleucon, Korn.

b. Grain red.

i. Awns red ..... var.ferrugineum, Korn.
ii. Awns black .... var. Sardoum, Korn.

4. Glumes red, pubescent.

a. Grain white ..... var. turcicum, Korn.

b. Grain red . . . . . . var. barbarossa, Korn.

5. Glumes grey-blue, glabrous.

- ' b. Grain red ... . . . var. caesium, Korn.

6. Glumes blue, pubescent.

b. Grain red . . . . . var. coeruleovelutinum, Korn.

7. Glumes black, pubescent.

b. Grain red . . . . . . \ar.fuligmosum, Korn.

II. Ears beardless —

8. Glumes white, glabrous.

a. Grain white ..... var. albidum, Korn. -

b. Grain red ...... var. lutescens, Korn.

9. Glumes white, pubescent.

a. Grain white .... var. leucospermum, Korn.

b. Grain red ...... var. velutinum, Korn.

10. Glumes red, glabrous.

a. Grain white ..... var. alborubrum, Korn.

b. Grain red ...... var. milturum, Korn.

11. Glumes red, pubescent.

a. Grain white . ... . . var. Delfii, Korn.

b. Grain red . ..... var. pyrothrix, Korn.

12. Glumes blackish-yellow, glabrous . . . var. triste, Flaksb.

13. Glumes bluish, pubescent.

b. Grain red ...... var. cyanothrix, Korn.

14. Glumes black, pubescent ..... var. nigrum, Korn.

T

274 THE WHEAT PLANT

BEARDED VARIETIES

Of some 1300 examples of T. vulgar e collected from all parts of the world
about one-half are bearded.

Among them Spring and Winter wheats are found in almost equal proportion,
some of the former being exceptionally rapid in their growth.

In all countries with hot summers, except Australia, bearded varieties pre-
vail, and although the baking quality of the flour of these wheats is usually good,
the yield per acre is generally very low.

In Western Europe, where intensive cultivation is practised, such wheats are
rarely grown. Occasionally bearded spring forms are sown when adverse
climatic conditions have prevented the sowing of the higher-yielding, beardless,
Winter wheats, but even in these circumstances a better financial return is
generally obtainable from oats, so that on most farms the latter crop, or barley
where the soil is suitable, takes the place of wheat when sowing is necessarily
delayed until March or April.

There is much less diversity of morphological and physiological characters
among the bearded than among the beardless forms of T. vulgare. More than
90 per cent possess lax, compressed ears, the density of which average 18-20
and rarely exceed 23-25. The awns are usually divergent, short in comparison
with those of T. durum or T. turgidum, varying in length from 5-10 cm. The
empty glumes have fine awns, 1-4 cm. long, or acute, apical teeth, which are never
so blunt as those of many beardless varieties and rarely so stout as those of T.
durum (cf. Figs. 138, 165, 166).

The varieties including the greatest number of forms are erythrospermum and
ferrugineum.

Ear bearded ; glumes white, glabrous ; grain white.

T. vulgare, var. graecum, Korn. Syst. Uebers. n (1873).

Kornicke received his type from Greece ; he obtained examples also from
Central Asia, Persia, and India.

The variety is described by Metzger (Eur. Cer. p. i A), who refers to it as an
uncommon wheat found among other varieties in Germany, France, Spain,
Italy, and England.

Although one of the less commonly grown varieties of T. vulgare, it is widely
distributed, especially in the warm wheat-growing regions of Asia.

Of fifty examples collected from different parts of the world, the majority
came from Persia, India, and Central Asia (Bokhara and the province of Semiret-
chensk, Turkestan), single forms only being obtained from Bulgaria, Austria,
France, Spain, Egypt, the Transvaal, New Zealand, United States, and Canada.

i. An early form received from the United Provinces, the Punjab, and
Bombay, and from Egypt under the name Hindi.

Young shoots, erect.

Straw, short and slender, 80-102 cm. (32-40 inches) high ; leaves more or less
glaucous.

Ear, 7-9 cm. long, almost square in section, 10 mm. across the face and side,
with stout awns 4-6 cm. long ; spikelets 16 to 19, usually overlapping, some-

FIG. 171. — BREAD WHEAT (T. vulgare, Host).

i. var. ferrugineum.
(Rouge prolifique.)

2. var. erythrospermum.
(Roumania.)

COMMON BREAD WHEAT 275

times irregularly arranged on the rachis, 3-grained ; 0 = 21-25 (Ear type i,
Fig. 176).

Empty glume, 10 mm. long ; apical tooth 2-3 mm. long (9, 14, Fig. 165).

Grain, short, mealy, or semi-flinty, 6-1-6-5 mm. long, 3-2-3-5 mm. broad,
3-2-3-4 mm. thick.

Roermaker and Ecksteen, with softer glumes, and pale yellowish-green
leaves, obtained from the Transvaal, are allied forms, probably derived originally
from India.

It is almost impossible to grow any of these wheats in England on account
of their susceptibility to attacks of Yellow Rust (Puccinia glumarum).

2. An early form received from Ispahan, Persia.
Young shoots, erect.

Straw, of medium height, 100 cm. (about 40 inches) high ; leaves glaucous.

Ear, lax, 9-10 cm. long, somewhat quadrate, 10 mm. across the face and side,
with plump spikelets and short, stiff awns 3-6 cm. long ; spikelets 15-18, plump,
2- to 3-grained ; D = 16-19 (^ar tvPe 2» Fig. 174).

Empty glume, 9 mm. long, keeled to the base, lateral nerves distinct, apex
broad ; apical tooth short, curved, 1-2 mm. long.

Grain, large, semi-flinty, plump ; 7-4 mm. long, 3-55 mm. broad, 3-35 mm.
thick.

3. Trigo Candeal. — A mid-season spelt-like form obtained from several
districts in Spain and from Italy.

Young shoots, erect.

Straw, tall, 120 cm. (about 48 inches) high ; upper internode with thick, pithy
walls or solid ; leaves glaucous.

Ear, lax, rigid, 10-12 cm. long, somewhat quadrate, 2-11 mm. across the
face and side ; awns slender, 6-9 cm. long ; spikelets 18-22, 3-grained ; D = IJ-
21 (Ear type 2, Fig. 173, lax).

Empty glume, 9 mm. long, keeled to the base ; apex broad, lateral nerve
prominent ; apical tooth acute, 3-5 mm. long (6, 20, Fig. 165).

Grain, large, semi-flinty, 7-3-8 mm. long, 3-7-4 mm. broad, 3-4 mm. thick.

4. Agh Bogda (White Wheat). — A mid-season spelt-like form received from
Tabriz, Persia.

Young shoots, erect or semi-erect.

Straw, of medium height, 90-110 cm. (36-44 inches) high ; leaves glaucous.

Ear, lax, 9-11-5 cm. long, flattish, 13 mm. across the face, 8 mm. across the
two-rowed side, with somewhat slender awns 6-9 cm. long ; spikelets 18-22,
3-grained ; D = 18-21 (Ear type i, Fig. 171).

Empty glume, 8-9 mm. long, keeled to the base, apex broad, apical tooth or
awn 2-5 mm. long (6, 8, Fig. 165).

Grain, semi-flinty or mealy, 6-5 mm. long, 3-2-3-7 mm. broad, 3-3-2 mm.
thick.

Allied spelt-like forms under the names Kizil Goon (golden colour) and
Ablakh (spotted), with broader glumes and larger grain (7-1 mm. long, 3-85 mm.
broad, 3-5 mm. thick), were obtained from Tabriz and also from Turkestan.

276 THE WHEAT PLANT

5. Winter forms from Russia, Bulgaria, Austria, and France.
Young shoots, prostrate or semi-prostrate.

Straw, tall, 110-120 cm. (44-48 inches) high.

Ear, 10-11 cm. long, awns slender, 6-7 cm. long; spikelets 18-21, 2- to
3-grained ; D = 18-21 (Ear type 2, Fig. 170).

Empty glume, 9-10 mm. long, keeled to the base, apex broad ; apical tooth
acute, 1-3 mm. long (6, 8, Fig. 165).

Grain, flinty or semi-flinty, 6-8-7 mm- l°ng> 3'9 mm. broad, 3-3 mm. thick.

6. Shirreff's White Bearded. — A winter form introduced by P. Shirreff
about 1860 ; it is a hardy wheat, productive and late, coming into ear at Reading
in the second or third week of June.

Young shoots, prostrate.

Straw, very tall, slender, 130-140 cm. (about 52-56 inches) high.

Ear, 10-12 cm. long, almost square in section, 10-11 mm. across the face
and side, with divergent awns 8-10 cm. long ; spikelets 22-25, 3-grained ;
0 = 23-26 (Ear type i, Fig. 178, with awns).

Empty glume, 7 mm. long, apex narrow ; apical tooth acute, 2 mm. long.

Grain, semi-flinty, small, plump ; 6-2-7 mm. long, 3-7-4 mm. broad, 3-2-3-5
mm. thick (9, Fig. 165).

Ears bearded ; glumes white, glaucous ; awns white ; grain red.

T. vulgare, var. erythrospermum, Korn. Syst. Uebers. n (1873).

This is one of the most widely distributed varieties of T. vulgare, forms of
it being found in almost all countries where wheat is cultivated. It is grown
more especially in regions having cold winters and hot summers.

Of the 250 examples collected from various parts of the world the majority
came from Russia, Roumania, Hungary, Spain, India, and the United States,
but a few were received also from Asia Minor, Persia, China, Japan, Manchuria,
Morocco, South Africa, Argentina, Australia, Portugal, Austria, Germany,
Holland, France, and England.

The greatest number are Spring and Winter wheats with lax, compressed ears.
With the exception of one or two prolific, bearded, Squarehead wheats they give
poor yields of grain, although the baking quality of their flour is generally
excellent.

i. An early form received from the Punjab, India, and common among
commercial samples from Karachi and Calcutta ; in ear at Reading about the
end of May.

Young shoots, erect.

Straw, short and slender, 80-96 cm. (32-38 inches) high ; leaves yellowish-
green.

Ear, 7-8 cm. long, somewhat square in section, with slender, divergent awns
5-6 cm. long ; spikelets 17-19, 3-grained ; D = 20-23 (Ear type i, Fig. 176).

Empty glume, 9 mm. long, apex broad ; apical tooth 2 mm. long (9, 21, Fig.
165).

Grain, flinty or semi-flinty, 6-5-6-7 mm. long, 3-5 mm. broad, 3-1 mm. thick.

FIG. 172. — BREAD WHEAT (T. vulgare, Host).

var. ferrugineum.
(Japanese forms.)

COMMON BREAD WHEAT 277

2. From the Punjab and the United Provinces was received a common form
similar to (i) in most morphological characters, but possessing more erect ears
and blue-green foliage.

3. Early forms received from various parts of Persia ; some forms of Chul
belong to these.

Young shoots, erect.

Straw, stout, of medium height, 95-110 cm. (38-44 inches) high.

Ear, lax, 11-13 cm- l°ng> awns 3-5 cm. long ; spikelets inflated and elongated,
18-20, 2- to 3-grained ; 0 = 17-20 (Ear types i, Fig. 173 ; i, 2, Fig. 174).

Empty glume, 10 mm. long, broad, keeled to the base ; apical tooth acute,
1-3 mm. long (6, 7, 10, Fig. 165).

Grain, large, flinty, or semi-flinty, 8-2 mm. long, 3 mm. broad, 3-4 mm. thick.

4. An early, somewhat spelt-like form, with rough, rigid ears and narrow
grains ; received from Yezd, Ispahan, Tabriz, and from Bokhara, and Semiret-
chensk. Similar forms occur among Portuguese, Spanish, and Argentine
wheats.

Young shoots, erect.

Straw, tall, 112-125 cm. (about 44-50 inches) high ; leaves glaucous.

Ear, narrow, 10-13 cm. long, almost square in section, 8-10 mm. across the
face and side, with hard glumes and stiff awns 6-8 cm. long ; spikelets 19-22,
2- to 3-grained ; 0 = 16-19 (Ear type 2, Fig. 173).

Empty glume, 10 mm. long, keeled to the base, apex broad ; apical tooth 3
mm. long (6, 10, Fig. 165).

Grain, semi-flinty, long, apex narrow, 7-4-8-4 mm. long, 3-3-6 mm. broad,
3-3-6 mm. thick.

5. Rieti. — An early form ; received from France, Italy, and Spain under
various names ; it comes into ear at Reading about June 9.

Most forms of Rieti have a pale red chaff.
Young shoots, erect.

Straw, slender, tall, 116 cm. (46 inches) high ; leaves yellow-green.
Ear, very lax, 12-13 cm. long ; awns 8-9 cm. long, spreading ; spikelets
21-22 ; D = 16-18 (Ear type i, Fig. 170).

Empty glume, 9 mm. long, apex narrow, its awn 1-5-2 cm. long (2, Fig. 165).
Grain, long, semi-flinty, 7-65 mm. long, 3-6 mm. broad, 3-15 mm. thick.

6. Soshu Shirokawa. — A very early form from Japan, having characteristic
ears with well-filled spikelets, which shed their glumes and grain very readily.

Young shoots, erect.

Straw, stout, erect, of medium height, 100-110 cm. (about 40-44 inches)
high ; leaves glaucous.

Ear, 10-11 cm. long with broad, well-filled spikelets, thin glumes often not
completely enclosing the grain, and fine, spreading awns 5-6 cm. long ; spikelets
20-22, 3- to 4-grained ; D = 20-22 (Ear type i, Fig. 172).

Empty glume, 7-8 mm. long, keeled to the base ; apical tooth or awn 2-6
mm. long (5, 17, Fig. 165).

278 THE WHEAT PLANT

Grain, short, dark red, flinty or semi-flinty, with blunt apex, 6 mm. long,
3-5-3-6 mm. broad, 3-1 mm. thick.

Kintana and other forms resembling (i), but with yellowish-green leaves,
were also received from Japan.

Allied to these early Japanese wheats, with non-glaucous, yellow-green leaves
and small, dark red, semi-flinty grain, are two forms obtained from Chungking,
China :

(a) With lax ears, 10-11 cm. long ; 0 = 18-19.

(b) With denser, square ears, 9 cm. long ; D = 25-27.

7. An early or mid-season form received from Italy and various districts in
Spain.

Young shoots, erect.

Straw, fine, tall, about 120 cm. (48 inches) high ; leaves glaucous.

Ear, very lax, 11-14 cm- l°ng > awns slender, 6-10 cm. long; spikelets
20-23, large> 3 -grained ; D = i6-i8 (Ear type i, Fig. 169).

Empty glume, 8-9 mm. long, keeled to the base ; apical tooth or awn 4-6 mm.
long (14, Fig. 165).

Grain, long, narrow, semi-flinty, 7-1 mm. long, 3-3-3-5 mm. broad, 3 mm.
thick.

8. A widely distributed form of Spring wheat, received chiefly from Spain,
Italy, Portugal, and Argentina ; the grain is of good quality, the ears shorter,
squarer, and slightly denser, and spikelets more overlapping than the pre-
ceding.

Young shoots, erect.

Straw, tall, 115-140 cm. (about 55 inches) high ; young leaves glaucous.

Ear, lax, 9-11 cm. long, somewhat square in section, 9-11 mm. across the
face and side ; awns divergent, 8-9 cm. long (Ear types i, 2, Fig. 170) ; spike-
lets 18-21, 3-grained ; D = 18-21.

Empty glume, 9-10 mm. long, keeled to the base ; apical tooth or awn 4-6
mm. long (3, 14, Fig. 165).

Grain, flinty, 7 mm. long, 3-4 mm. broad, 3 mm. thick.

9. A very widely distributed winter form with grain of fine quality ; culti-
vated extensively in European Russia, Austria, Hungary, and Roumania, the
United States, and Canada; also received from France and Holland. Turkey
Red, Malakov, Banat, and Lancaster are examples.

Young shoots, prostrate.

Straw, tall, 120-130 cm. (48-52 inches) high.

Ear, lax, 9-13 cm. long, compressed, 12-14 mm. across the face, 8-10 mm.
across the 2-ranked side ; awns divergent, 6-8 cm. long ; spikelets 23-26 broad,
3-grained ; D = 18-22 (Ear types 2, Fig. 170 ; 2, Fig. 171).

Empty glume, 9-10 mm. long ; apical tooth 2-5 mm. long (8, 21, Fig. 165).

Grain, flinty, somewhat short, apex blunt ; 6-8-7 mm- l°ng> 3'5 mm. broad,
3 mm. thick.

10. Badger. — A prolific Winter wheat selected by the author from an Austrian
form.

FIG. 173. — BREAD WHEAT (T. vulgar e, Host).

i. var. graecum.
(Chul.)

2. var. erythrospermum,
(Persia.)

COMMON BREAD WHEAT 279

Allied to it, but with less dense ears, are Beal and Strube's Hybrid from
Germany.

Young shoots, prostrate.

Straw, stiff, of medium height, 96-115 cm. (38-44 inches) high; leaves
glaucous.

Ear, dense, 8-9 cm. long, 12-14 mm. across the face, 10 mm. across the 2-
ranked side ; awns slender, 5-6 cm. long ; spikelets 23 to 26, 3-grained ; D =
30-32 (Ear type 2, Fig. 176).

Empty glume, 8 mm. long, inflated ; apical awn or tooth curved at the base,
2-3 mm. long (15, Fig. 165).

Grain, semi-flinty, plump, 6-9-7 mm. long, 3-9 mm. broad, 3 mm. thick.

Ear bearded ; glumes white, glabrous ; awns black ; grain red.
T. vulgare nigroaristatum, Flaksb. Bull. App. Bot. Petrograd, viii. 195

According to Flaksberger, this variety occurs sporadically among other
wheats in Russia.

Ear bearded ; glumes white, pubescent ; grain white.

T. vulgare, var. meridionale, Korn. Handb. d. Getr. i. 47 (1885).

Kornicke's type was obtained from Greece, but the variety is endemic in
Asia.

Of the 22 examples grown at Reading, 2 were from India, i from Rhodesia,
i from the Province of Syr Daria, Turkestan, the rest (18) came from Persia
(Bushire, Tabriz, Ispahan, Arabistan).

All the forms of this variety are early wheats ; those from India with short
straw and ears, the Persian forms having straw of medium height and lax ears
9-11 cm. long.

i . A very early form received from Cawnpore.

Young shoots, erect.

Straw, of medium height, 89-102 cm. (35-40 inches) high ; leaves blue-green.

Ear, 7-8 cm. long, with rigid, spreading awns 5-6 cm. long ; spikelets 15-18
overlapping, often irregularly arranged on the rachis ; D = 20-23 (Ear type i ,
Fig. 176).

Empty glume, 8 mm. long, keeled to the base, apex broad ; apical tooth
stout, acute, 2 mm. long.

Grain, small, flinty, 6-25 mm. long, 3-2 mm. broad, 3-1 mm. thick.

Allied forms with yellow-green leaves were obtained from Arabistan and
Rhodesia, the latter probably derived originally from India.

2. Agh Bogda (White Wheat). — A mid-season form received from Tabriz,
Persia.

Young shoots, semi-erect.

Straw, medium to tall, 104 cm. (40-46 inches) high ; leaves glaucous.

Ear, lax, 9-11 cm. long, somewhat square in section; with stout awns 7-9
cm. long; spikelets 17-19, inflated, often 4-grained ; D = 16-20 (Ear type i,
Fig. 179, with awns).

28o • THE WHEAT PLANT

Empty glume, 10 mm. long, keeled to the base ; apical awn acute, 10-20 mm.
long, with a narrow, dark brownish-purple line along the inner margin (3,6,
Fig. 165).

Grain, large, more or less mealy, plump ; 7-5-7-8 mm. long, 3-8 mm. broad,
3-45 mm. thick.

Similar forms are Sari Bogda (yellow wheat), Kizil Goon (golden colour),
and Ablakh (spotted), from the same district ; others also from Ispahan,
Persia, and one from Syr Daria, Turkestan.

Ear bearded ; glumes white, pubescent ; awns black ; grain white.

T. vulgare, var. pseudo-meridionale, Flaksb. Bull. App. Bot. Petrograd,
viii. 163 (1915).

A very rare variety, hitherto found only in Asia.

Flaksberger states that this variety frequently occurs among other wheats
in Persia and Turkestan.

Howard's var. meridionale, United Provinces Class 6, appears to belong to
this also (Wheat in India, p. 193).

Ear bearded ; glumes white, pubescent ; awns white ; grain red.

T. vulgare, var. Hostianum, Korn. Handb. d. Getr. i. 47 (1885).

A rare variety known to Kornicke only from Botanic Gardens. It occurs
among commercial White Karachi wheat from India, and I have received ten or
twelve forms of it from the United Provinces, India, Tabriz, and Seistan in
Persia, the province of Semiretchensk, Turkestan, and a single form from
Professor Eriksson, Stockholm.

1. An early form found in commercial white Karachi wheat and received
also from Cawnpore, India. At Reading it comes into ear about the end of
May or first week of June.

Young shoots, erect or semi-erect.

Straw, short, slender, erect, 76 cm. (30 inches) high ; leaves blue-green.

Ear, dense, 7-5-8 cm. long, quadrate, 10-11 mm. across the face and side,
with somewhat slender awns 6-7-5 cm- l°ng 5 spikelets 16-19 5 0=24-28 (Ear
type i, Fig. 176).

Empty glume, 9 mm. long, keeled to the base, with apical tooth or awn 4-10
mm. long (15, 16, Fig. 166).

Grain, flinty, dark red, small, plump ; apex blunt, 6-2-6-8 mm. long, 3-6-4
mm. broad, 3-3-25 mm. thick.

2. Ablakh (Spotted). — An early wheat received from Tabriz, Persia.
Young shoots, erect or semi-erect.

Straw, stout, of medium height 100-110 cm. (40-44 inches) high ; leaves
blue-green.

Ear, lax, 10-11 cm. long, with stout awns, 7-8 cm. long ; spikelets 15-16,
plump, inflated, 3- to 4-grained ; D = 16-19 (Ear type 2, Fig. 187, with awns).

Empty glume, 10 mm. long, keeled to the base ; apical tooth 2-4 mm. long,
with a narrow, dark brownish-purple line along the inner margin.

FIG. 174. — BREAD WHEAT (T. vulgar e, Host).

i. var. pseudoturcicum. 2. var. erythroleucon.

(Kadiz Oglau.) (Kalkori.)

COMMON BREAD WHEAT 281

Grain, large, dark red, flinty, 7-5 mm. long, 3-5-4 mm. broad, 3-5 mm. thick.
A similar form with more hairy leaves from Seistan, Persia, under the name
Nishkin.

3. An early form received from the province of Semiretchensk, Turkestan.
Young shoots, erect.

Straw, of medium height, 100-110 cm. (40-44 inches) high.

Ear, 8-10 cm. long, with spreading awns 5-6-5 cm. long ; spikelets 17-19 ;
0 = 20-25 (Ear type 2, Fig. 170).

Empty glume, 8 mm. long, keeled to the base ; apical tooth 2 mm. long,
lateral nerve distinct ; no dark margin to the glume (8, 14, Fig. 165).

Grain, dark red, 6-5 mm. long, 3-4 mm. broad, 3-3 mm. thick.

4. A very late winter form, received from Prof. Eriksson, Stockholm ; it
comes into ear at Reading about the third week in June.

Young shoots, prostrate.

Straw, slender, erect, of medium height, 109 cm. (43 inches) ; leaves blue-
green.

Ear, 9-10.5 cm. long with slender, spreading, somewhat wavy awns 5-6 cm.
long ; spikelets 22, short, 3-grained ; D = 22-24 (Ear type 2, Fig. 177, with awns).

Empty glume, 7 mm. long, keeled to the base ; apical tooth slender, 2-3-5 mm-
long (Form 21, Fig. 165).

Grain, more or less visible between the glumes, semi-flinty, 6-45-6-6 mm.
long, 3-45-4 mm. broad, 3-4 mm. thick.

Ear bearded ; glumes while, pubescent ; awns black ; grain red.

T. vulgare, var. pseudo-Hostianum, Flaksb. Bull. App. Bot. Petrograd,
viii. 165, 196(1915).

A very rare variety occurring only in Central Asia.

Flaksberger states that examples are occasionally found in Turkestan and
Persia, mixed with other wheats.

I obtained examples among Ablakh wheat from Tabriz, Persia, exactly
similar to the Ablakh form described under var. Hostianum (p. 280), but with
black awns.

Punjab 9. — Received from India. A distinct form with dense, square
ears and villose glumes, the hairs conspicuous and spreading.

Straw, tall, hollow, 118-120 cm. (46-47 inches) high.

Ear, 6-5-7-5 cm. long, dense, square, 10-11 mm. across the sides ; spikelets
18-20; 0 = 30 (Ear types 2, Fig. 175 ; 2, Fig. 176 but narrower) ; awns rigid,
stout, and short, 5-5-6-5 cm. long.

Empty glume, 8-9 mm. long, with prominent apex and awn 5-6 mm. long
(Form 20, Fig. 165).

Grain, dark red, flinty, 6-8 mm. long, 3-4 mm. broad, 3-1 mm. thick.

Ear bearded ; glumes red, glabrous ; grain white.

T. vulgare, var. erythroleucon, Korn. Handb. d. Getr. i. 47 (1885).
A rare variety endemic in Central Asia, Persia, and India.

282 THE WHEAT PLANT

Kornicke's examples were derived from Turkestan, Persia, and India. Of
the 22 examples grown at Reading 12 were obtained from India (the Punjab
and United Provinces), 3 from Persia, 3 from Syr Daria and Semiretchensk,
Turkestan, 2 from the Transvaal, and i each from France and Spain.

1. An early form received from the Punjab and the United Provinces.
Several closely similar forms with bluish-green leaves and semi-flinty grain
were received from Karachi, Cawnpore, and Calcutta.

Young shoots, erect.

Straw, short, slender, 90-125 cm. (36-45 inches) high ; leaves pale-yellowish
green.

Ear, lax, 6-5-8-5 cm. long ; awns short, spreading, 6-7 cm. long ; spikelets
1 6, irregularly arranged on the rachis and usually overlapping each other in the
denser ears ; D = 18-23 (Ear types 2, Fig. 173 ; i, Fig. 176).

Empty glume, rigid, 10 mm. long, keeled to the base ; apical tooth acute, 2-3
mm. long (8, 14, Fig. 165).

Grain, flinty, dorsal ridge prominent, apex truncate, 7-2 mm. long, 3-9 mm.
broad, 3-5 mm. thick.

An allied form with yellow-green leaves and one under the name Siebritz
with somewhat glaucous foliage were received from the Transvaal.

2. An early form received from Tabriz, Persia, Syr Daria, and Semiret-
chensk, Turkestan.

Young shoots, erect.

Straw, of medium height, 100-1 10 cm. (40-44 inches) high ; leaves blue-green.

Ear, glaucous, 8-12 cm. long, narrow, quadrate, 9-10 mm. across the face
and side, with slender awns 6-8 cm. long; spikelets 16-18, 2-grained ; D =
17-21 (Ear type 2, Fig. 173).

Empty glume, 9 mm. long, keeled to the base ; apical tooth or awn 3 mm.
long (2, 6, Fig. 165).

Grain, flinty, long, and somewhat narrow, 8 mm. long, 3-6 mm. broad, 3-1
mm. thick.

3. Perle de Nuisement. — A very late winter form received from France ;
it comes into ear June 16-28 at Reading.

Young shoots, prostrate.

Straw, slender, tall, 127 cm. (50 inches) high ; leaves yellow-green.

Ear, 9-12 cm. long, with broad, well-filled spikelets and spreading awns 8-9
cm. long ; spikelets 20-24, often 4-grained ; D = 20-22 (Ear type i, Fig. 171).

Empty glume, 9-10 mm. long, keeled to the base, apex narrow ; apical tooth
acute, 3 mm. long (15, Fig. 165).

Grain, flinty, 6-5 mm. long, 3'5-3'7 mm. broad, 3-25 mm. thick.

A closely similar form with blue-green leaves was received from Spain under
the name Del Frailte.

Ear bearded ; glumes red, glabrous ; awns red ; grain red.

T. vulgare, var. ferrugineum, Korn. Handb. d. Getr. i. 47 (1885).
Like the corresponding white-chaffed var. erythrospermum this is widely

FIG. 175. — BREAD WHEAT (T. vulgar e, Host).

var. ferrugi neum .
(Siberian forms.)

COMMON BREAD WHEAT 283

distributed and includes a large number of forms, most of which are Spring and
Winter wheats with lax, compressed ears, and grain of high quality. Over 200
examples of it have been collected and grown at Reading, these coming from
every country where wheat is cultivated.

As in the case of the corresponding white-chaffed var. erythrospermum, the
commonest forms possess lax, compressed ears ; dense-eared forms are excep-
tionally rare in this variety.

1 . Very early forms from Northern India and Persia ; they come into ear
at Reading about the end of May.

Young shoots, erect.

Straw, short, slender, 70-85 cm. (28-34 inches) high ; leaves yellow-green.

Ear, 7'5-9'5 long, almost square in section, 10-12 mm. across the face ; awns
4-7 cm. long, scout at the base, brittle; spikelets 16, elongated; D = 16-20
(Ear type i, Fig. 176).

Empty glume, rigid, 9-10 mm. long, keeled to near the base ; apical tooth
2-3 mm. long (14, 20, 21, Fig. 165).

Grain, flinty, well-filled, 7-5-7*9 mm. long, 3-7 mm. broad, 3-5 mm. thick.

2. Forms similar to the preceding, with ears 9-11 cm. long, spikelets over-
lapping, and density 20-25, are frequent in Persia, Central Asia, and India ; van
Niekirk's wheat received from the Transvaal belongs to these.

3. Spelt-like early forms from Bokhara, Turkestan, and other parts of
Central Asia.

Young shoots, erect.

Straw, stout, rigid, of medium height, 100 cm. (about 40 inches) high ;
leaves yellow-green.

Ear, long, lax, 10-12 cm. long, somewhat square, 10 mm. across the face
and side ; awns rigid, 6-8 cm. long ; spikelets 16-18, narrow, elongated, 2- to
3-grained ; D = 16-20 (Ear types i, 2, Fig. 173).

Empty glume, rigid, 10 mm. long, keeled to the base, apex usually broad with
prominent lateral nerve ; apical tooth acute (4, 6, Fig. 165).

Grain, flinty or semi-flinty, 6-8-7 mm. long, 3-6 mm. broad, 3 mm. thick.

4. Barletta. — An early form received from Spain, Argentina, and Para-
guay ; it comes into ear at Reading about the first week in June.

Young shoots, erect.

Straw, tall, 118-132 cm. (47-52 inches) high ; leaves glaucous.

Ear, long, lax, 10-12 cm. long, almost square in section, 10-12 mm. across
the face and side ; awns 8-9 cm. long ; spikelets 18-21, elongated ; D = 19 (Ear
type 2, Fig. 173).

Empty glume, pale red, 8-10 mm. long, rigid, keeled to the base ; apical
tooth or awns 2-5 mm. long ; lateral nerve prominent (14, 15, Fig. 165).

Grain, semi-flinty, apex narrow, 6-7 mm. long, 3-5 mm. broad, 3-2 mm. thick.

5. Somewhat similar are several forms received from Spain with more rigid
ears and glumes, and longer, flinty grain, 8 mm. long, 3-5 mm. broad, 3-25 mm.
thick.

284 THE WHEAT PLANT

6. Aka Yenidashi. — An early form received from Japan ; it comes into ear
at Reading about the end of May.

Young shoots, erect.

Straw, of medium height, about 96 cm. (38 inches) high ; leaves yellow-green.

Ear, lax, 8-9 cm. long ; awns 5-6 cm. long, spreading ; spikelets broad,
16-20 ; D = 20-23 (Ear types 3, 4, Fig. 172).

Empty glume, 8 mm. long, keeled to the base, apex broad ; apical tooth
acute, 3 mm. long (5, 6, 19, Fig. 165).

Grain, flinty, 6 mm. long, 3-1 mm. broad, 2-6 mm. thick.

Allied to this is Hosagara, also from Japan, with less erect young shoots,
slightly taller straw, and plumper, broader grain.

7. Daruma. — An early form received from Japan ; it comes into ear at
Reading about the end of May.

Young shoots, semi-erect.

Straw, stout, medium to tall, 100-115 cm. (40-46 inches) high; leaves
yellow-green.

Ears, 10 cm. long ; awns 5-6 cm. long, stout, spreading ; spikelets 25 ;
0 = 25 (Ear type i, Fig. 172).

Empty glume, short, inflated, 7 mm. long, apex broad ; apical tooth bent
inwards, 4 mm. long (19, Fig. 165).

Grain, flinty, short, plump, 5-2 mm. long, 3-3 mm. broad, 3-1 mm. thick.

Resembling this is a denser-eared Japanese form (0 = 27-31), Sunagawa
yanagi Kobu, with shorter, more slender awns, and somewhat longer grain.

8. Hsu Hsu Mai. — An early form received from Chungking, China ; it has
given rise to several semi-bearded forms, with short awns 1-5-2 cm. long.

Young shoots, erect.

Straw, short to medium height, 85-90 cm. (about 36 inches) high ; leaves
yellow-green.

Ear, lax, 10-11 cm. long, with very fine awns 7-8 cm. long ; spikelets broad,
21 ; D= 20-23 (Ear type i, Fig. 172).

Empty glume, 8 mm. long, apex broad, lateral nerve distinct ; apical tooth
acute, 3 mm. long, curved inwards slightly (19, Fig. 165).

Grain, mealy, apex narrow, dorsal ridge somewhat prominent, 6-3 mm. long,
3-45 mm. broad, 3-15 mm. thick.

9. Xeixa. — A late form received from Spain ; it comes into ear at Reading
about June 18.

Young shoots, erect.

Straw, slender, tall, about 127 cm. (50 inches) long ; leaves yellow-green.

Ears, lax, 10-12 cm. long, with awns 8 cm. long, 12-14 mm. across the face,
10 mm. across the side ; spikelets 16-18 ; D = 18 (Ear type i, Fig. 170).

Empty glume, 8 mm. long ; apical tooth or awn 2-15 mm. long (8, 15, Fig. 165).

Grain, flinty, apex somewhat truncate, 7-1 mm. long, 3-65 mm. broad, 3-35
mm. thick.

Romanella from Italy and many forms received from Spain under different
names resemble this.

Vwf < > ' ' \\

FIG. 176. — BREAD WHEAT (T. vulgar e, Host).

i. var. erythrospermum.

2. var. erythrospermum.
(Bearded Squarehead.)

COMMON BREAD WHEAT 285

10. La Pera. — An early form received from Spain ; it comes into ear at
Reading about the first week in June.

Young shoots, erect.

Straw, slender, tall, 110-120 cm. (44-48 inches) high ; leaves glaucous.

Ear, lax and flattish, 10-12 cm. long, 12 mm. across the face, 9-10 mm.
across the side ; awns 8-9 cm. long; spikelets 18-20; D = i8 (Ear types i, 2,
Fig. 170).

Empty glume, pale red, 10 mm. long, 3-55 mm. broad, 3-35 mm. thick (2, 8,
Fig. 165).

Grain, long, flinty, 7-8 mm. long, 3-55 mm. broad, 3-35 mm. thick.

11. Winter (Azima) wheats received chiefly from Russia, Roumania, and
Bulgaria ; the commonest forms of var. ferrugineum.

Young shoots, prostrate.

Straw, tall, slender, no cm. (44 inches) high ; leaves glaucous.

Ear, lax, 10-13 cm. long, often tapering towards both ends ; spikelets 22-24 ;
D = 19-20 (Ear types i, Fig. 170 ; 2, Fig. 171).

Empty glume, 10 mm. long, keeled to near the base, apex bluntish ; apical
tooth acute, 2-4 mm. long (2, 20, Fig. 165).

Grain, flinty or semi-flinty, 6-8 mm. long, 3-65 mm. broad, 3-1 mm.
thick.

12. Fine, narrow-eared forms, with very small, narrow, pointed grains ;
var. ferrugineum sibiricum of Flaksberger. Early to mid-season wheats received
from Russia in commercial samples of Ulka and " Fine North Russian Wheat,"
and from Canada under the name Onega.

Young shoots, semi-erect.

Straw, slender, of medium height, 100-110 cm. (40-44 inches) high ; leaves
glaucous and usually pubescent.

Ear, narrow, tapering, 6-10 cm. long, with fine awns 5 cm. long ; spikelets
23-27 ; D = 22-25 (Ear types i, 2, Fig. 175).

Empty glume, narrow, keeled to the base, 8-9 mm. long, apex truncate ;
apical tooth 1-1-5 mm- l°ng> acute (13, Fig. 165).

Grain, flinty, narrow, very small, 5-9-6-3 mm. long, 2-65-2-9 mm. broad,
2-3-2-9 mm. thick.

Allied to these forms are Rouge barbu de Mars from France.

13. Carman. — A dense-eared late form received from Haage and Schmidt,
Germany.

Young shoots, prostrate.

Straw, of medium height, about 100 cm. (40 inches) high ; leaves glaucous.

Ear, dense, 9-10 cm. long, square, 10-11 mm. across the face and side;
awns fine, 4-6 cm. long ; spikelets 22-25 > D = 25-30.

Empty glume, 9-10 mm. long, keeled to the base, apex broad ; apical tooth
stout, short.

Grain, plump, mealy, 7-2 mm. long, 4 mm. broad, 3-3 mm. thick.

Somewhat similar to this are Michigan Bronze and Golden Cross.

286 THE WHEAT PLANT

Early bearded ; glumes red, glabrous, awns black ; grain red.

T. vulgare, var. Sardoum, Korn. Handb. d. Getr. i. 46 (1885).

The varietal name was given by Kornicke to a wheat mentioned by Seringe
(Ce'r. eur. p. 127) as " variation F. Touzelle Saisette (brune chauvre, barbes
noires) or Touzelle de Sardaigne," a bearded form with lax, yellowish-red ears,
awns almost black, stiff straw, and soft grain tending to become hard ; the
colour of the grain is not given by Seringe.

I have seen no variety of T. vulgare with all the characters mentioned by
Kornicke.

Ear bearded ; glumes red, pubescent ; grain white.

T. vulgare, var. turcicum, Korn. Handb. d. Getr. i. 48 (1885).

A rare variety, which I have received only from Asia, i example from India,
i from Turkestan, and 1 1 from Persia.

Kornicke's examples came from Kastamoni, near the north coast of Asia
Minor, and from Turkestan, Central Asia.

1 . An early form received from Cawnpore, India.
Young shoots, semi-erect.

Straw, slender, of medium height, 90-110 cm. (36-44 inches) high ; leaves
pale yellowish-green.

Ear, short, somewhat dense, 7 cm. long, with spreading awns 5-6 cm. long ;
spikelets 16-18 ; 0 = 23-28 (Ear type 2, Fig. 170).

Empty glume, 8-9 mm. long, keeled to the base ; apical tooth durum-\ike,
stout, acute, 1-2 mm. long (16, 18, Fig. 165).

Grain, small, flinty, 6 mm. long, 2-7 mm. broad, 2-8 mm. thick.

2. Kar-i-safid — A very distinct early form received from Khorasan, Persia,
where it is grown on irrigated soils.

Young shoots, erect.

Straw, of medium height, no cm. (about 44 inches) high ; leaves blue-green

Ear, lax, 10-13 cm. long, with inflated spikelets arranged irregularly on the
rachis, and spreading awns 9-10 cm. long ; spikelets 20-21, 3- to 4-grained ; D =
18-20 (Ear type i, Fig. 179, with awns).

Empty glume, 10 mm. long, inflated ; apical tooth narrow, 4-10 mm. long
(14, 20, Fig. 165).

Grain, large, mealy, or semi-flinty, 7-2-7-4 mm. long, 3-8 mm. broad, 3-4 mm.
thick.

Allied to it are forms from Ispahan, Persia, and Turkestan, and a true Winter
wheat with prostrate young shoots from Tabriz under the name Kizil Goon
(golden colour), with amber, flinty grain.

Ear bearded ; glumes red, pubescent ; grain red.

T. vulgare, var. barbarossa, Korn. Handb. d. Getr. i. 48 (1885).
A rare variety, endemic only in Persia and India.

Kornicke received examples from Kastamoni, near the north coast of Asia
Minor, Turkestan, and North America.

FIG. 177. — BREAD WHEAT (T. vulgar e, Host).

i. var. lutescens.
(Granella di Carpegna.)

2. var. albidum.
(Bordier.)

COMMON BREAD WHEAT 287

Three examples were sent to me from India (the Punjab and United Pro-
vinces), 2 from Eastern Bokhara, 9 from various provinces of Persia, and an
isolated form from Rhodesia, South Africa.

An early form received from Karachi ; similar forms also from Cawnpore
and the Punjab, India.

Young shoots, semi-erect.

Straw, of medium height, 106 cm. (42 inches high) ; leaves pale yellowish-
green.

Ear, lo-n cm. long, with spreading, somewhat slender awns 5-7 cm. long ;
spikelets 19-21, large, well-filled, 3- to 4-grained ; D = 18-21 (Ear types 2, Fig.
170 ; i, Fig. 176).

Empty glume, 9-10 mm. long, with acute, apical awn, 4-6 mm. long (14, 20,
Fig. 165).

Grain, semi-flinty, narrowed at the apex, 6-5-6-9 mm. long, 3-4 mm. broad,
3-3 mm. thick.

A series of forms similar in all respects to Kar-i-safid wheat (var. turcicum),
but with red grains, were received from Tabriz under the name Sari Bogda
(yellow wheat), Ispahan and Seistan in Persia, and from E. Bokhara ; a single
example also from Rhodesia.

Ear bearded ; glumes grey-blue, glabrous ; grain red.

T. vulgare, var. caesium, Korn. Handb. d. Getr. i. 47 (1885).

A very rare variety, a specimen of which Kornicke obtained from Botanic
Gardens.

Metzger (Eur. Cer. p. 4, F.) describes a lax-eared form of this variety with the
following characters :

Straw, tall, 116-132 cm. (46-52 inches) high, hollow.

Ear, 8-1 1 cm. long, erect, flattish, tapered towards the apex.

Empty glume, pointed, with long, apical tooth ; upper portion of flowering
glume bluish, the base white ; awns spreading, rough, brownish, as long as, or
longer than, the ear.

Grain, mealy, oval, short, reddish-grey.

A Spring wheat, giving only a moderate yield.

Ear bearded ; glumes grey-blue, pubescent ; grain red.

T. vulgare, var. coeruleovelutinum, Korn. Syst. Uebers. 12 (1873).

A rare variety.

Kornicke's specimens of this variety were obtained from Botanic Gardens
and from Vernoe, Turkestan.

According to Werner (Handb. d. Getr. ii. 378), the form from Turkestan
possesses the following characteristics :

Young shoots, erect, young leaves densely hairy.

Straw, of medium height, 100-110 cm. (39-43 inches) high.

Ear, 9-12 cm. long, narrow, of medium density ; awns pale, 10 cm. long.

Empty glume, white, with bluish tinge.

288 THE WHEAT PLANT

Grain, mealy, yellowish-red, roundish, 7 mm. long, 4 mm. broad.

A winter form is also described by Werner, with upright young shoots
and slightly hairy, yellowish-green leaves.

Straw, 115-140 cm. high ; thick- walled.

Ear, very lax, narrow, 10-13 cm. long, dark and greyish-blue, with awns
8-10 cm. long.

Grain, flinty, long, and narrow, 8 mm. long, 3 mm. broad.

Both the above forms are very susceptible to rust. Many of their morpho-
logical characters suggest a close relationship to T. turgidum.

Ears bearded ; glumes black, pubescent ; grain red.

T. vulgare, var. fuliginosum, Korn. Handb. d. Getr. i. 48 (1885).

A very rare variety, which Kornicke found only in Botanic Gardens.

Straw, tall, 116-132 cm. (46-52 inches) high, solid.

Ear, 5-7 cm. long, flattish ; spikelets 16-20, broader than long.

Empty glume, inflated, black, clothed with fine hairs ; upper part of the
flowering glume black, lower part reddish.

Grain, mealy, plump, yellow.

Although Metzger refers this to T. vulgare, he notes that it possesses some of
the features of T. turgidum.

The solid straw, broad spikelets, inflated glumes, and plump, rounded,
starchy grains suggest close affinity with T. turgidum.

Werner (Handb. d. Getr. ii. 378) describes a winter form with erect, quad-
rangular ears 10 cm. long, pale yellow or reddish-yellow glumes, their margins,
keel, and apical tooth blue ; awns blackish, spreading, 9 cm. long ; grain starchy,
yellowish-red, oval, 7 mm. long, 4 mm. broad ; it appears to be closely allied to
T. turgidum.

Flaksberger (Bull. App. Bot. viii. 197) mentions forms of var. fuliginosum,
with glumes having black hairs on a yellow ground, occurring in Turkestan and
the Tiflis province of Transcaucasia.

BEARDLESS VARIETIES

As explained elsewhere (p. 104), there are comparatively few varieties of
wheats whose flowering glumes are totally devoid of awns, most of the so-called
beardless forms having awns varying in length from -5 to 2-5 cm., those with the
longest, which are always found near the apex of the ear, are conveniently
described as " tip-bearded."

It is among the beardless varieties that the most prolific wheats are found,
and it is largely on this account that these, rather than bearded forms, are grown
on the intensively cultivated lands of Western Europe.

The beardless varieties exhibit a much greater range of variability in all
their characters than bearded wheats. In regard to the density of the ear, all
grades are found in cultivation between the most open, lax ears and the dense
Squarehead types. By far the greater number of forms belong to the varieties
lutescens, milturum, and albidum.

FIG. 178. — BREAD WHEAT (T. vulgare, Host).

i . var. albidum.
(Russia.)

2. var. albidum.
(Semiretchensk .)

COMMON BREAD WHEAT 289

Ear beardless ; glumes white, glabrous ; grain white.

T. vulgare albidum, Korn. Handb. d. Getr. i. 45 (1885).

A widely distributed variety, much more numerous in its forms than the
corresponding bearded variety graecum. About 150 examples were collected
from England, France, Germany, Holland, Russia, South Africa, Persia, India,
Australia, New Zealand, and the United States.

They are most commonly cultivated at the present time in dry warm climates,
the grain of those from Australia being especially well developed and of a fine
white tint.

With the exception of some of the Squarehead type of ear, forms of this
variety are rarely grown in Britain or the countries of Western Europe. Like
all white-grained wheats, they are extremely liable to sprout in the ear when
cut and left in the field in a damp season.

1. Gentile bianco. — An early form received from Italy. It comes into ear
at Reading about the end of May.

Young shoots, erect.

Straw, slender, tall, 120-128 cm. (48-50 inches) high.

Ear, lax, 10 cm. long ; upper spikelets with awns -5-'7 cm. long ; spikelets
20 ; 0 = 19-20 (Ear type 2, Fig. 177).

Empty glume, 9-10 mm. long, apex truncate, apical tooth blunt, -5-1 mm.
long (2, Fig. 1 66).

Grain, large, semi-flinty, 7-3 mm. long, 3-65 mm. broad, 3-45 mm. thick.

Closely similar is Napoles from Spain and a form from Chili.

2. An early form, coming into ear May 20-26 at Reading ; received from
the Central Provinces and United Provinces, India.

Young shoots, erect.

Straw, slender, short to medium, height 75-95 cm. (30-36 inches) long.
Ear, lax, 9-10 cm. long, with awns 5-7 mm. long ; spikelets 19-21 ; D =
20-23.

Empty glume, 9 mm. long ; apical tooth blunt, -5 mm. long.

Grain, semi-flinty, large, plump ; 7 mm. long, 3-9 mm. broad, 3-75 mm. thick.

A similar form was received from Australia.

3. G-andum-i-Kaiseh. — An early form ; received from Persia. It comes into
ear at Reading about May 20.

Young shoots, erect.

Ear, erect, 9 cm. long, with awns -5-1 cm. long; spikelets 19; D=22
(Ear type 2, Fig. 178).

Empty glume, 8 mm. long, keeled to base, broad, apex with incurved claw-
like tooth, 2 mm. long (i, Fig. 166).

Grain, large, semi-flinty, plump ; 7-2 mm. long, 3-5 mm. broad, 3-55 mm.
thick.

4. Epp. — A mid-season variety received from Germany.
Young shoots, prostrate.

Straw, very tall, 142 cm. (about 56 inches) high.

U

290 THE WHEAT PLANT

Ear, ion cm. long, with spreading spikelets and short awns, 3-5 mm.
long ; spikelets 21-23 ; D=22 (Ear type 2, Fig. 177).

Empty glume, 9 mm. long, keeled to the base, apex broad ; apical tooth blunt,
•5 mm. long (2, n, Fig. 166).

Grain, semi-flinty, large, loosely held in the glumes ; 7 mm. long, 3-95 mm.
broad, 3-7 mm. thick.

A similar form with laxer ears (0 = 17) from Russia.
«

5. Fenton. — A late form coming into ear at Reading about June 20. Intro-
duced by G. Hope of Fenton Barns, East Lothian, Scotland, who found it in a
quarry on his farm in 1835. ^ was mucn grown down to 1870, about which
date it began to be supplanted by Squarehead wheat.

Young shoots, prostrate.

Straw, somewhat slender, tall, 127 cm. (about 50 inches) high.

Ear, lax, tapering towards both ends, about 10 cm. long, with a few awns
i cm. long near the apex ; spikelets 20-23 ; D = 21-23 (Ear type i, Fig. 178).

Empty glume, 8 mm. long, apex truncate, apical tooth blunt, -5-1 mm. long
(6, 9, Fig. 166).

Grain, flinty, plump, 6 mm. long, 3-5 mm. broad, 3-2 mm. thick.

The following forms completely resemble this variety in habit of growth,
time of ripening, and morphological characters of the straw and ear :

Chidham, a famous Old English wheat found in a hedge at Chidham in
Sussex in 1789 or 1790, Trump, Zealand White, Markischer, and some Russian
forms.

6. Solid - straw White Tuscan. — A form with solid straw and empty
glumes resembling those of T. Spelta ; received from New Zealand, the United
States, and Austria.

Young shoots, erect.

Straw, somewhat slender, very tall, 120-132 cm. (48-52 inches) high, solid.

Ear, 9-10 cm. long ; spikelets 20 ; D = 20-23 (Ear type 2, Fig. 181).

Empty glume, rigid, 8 mm. long, keeled to the base ; apex broad, truncate ;
apical tooth blunt, -5-1 mm. long (10, n, Fig. 166) ; lower and middle flowering
glumes with claw-like awns 5-6 mm. long ; awns of the apical spikelets 2-5 cm.
long.

Grain, amber, flinty, 7 mm. long, 3-6 mm. broad, 3-5 mm. thick.

7. Bobs. — An early form received from Australia. It comes into ear at
Reading about the end of May.

Young shoots, erect or semi-erect.

Straw, somewhat slender, of medium height, 109-114 cm. (about 43-45
inches) high.

Ear, 9-10 cm. long, completely awnless, tapered slightly ; spikelets 20-22 ;
D = 20-22 (Ear type i, Fig. 180).

Empty glume, rigid, 8 mm. long, apex truncate, apical tooth blunt, -5-1 mm.
long (22, Fig. 166).

Grain, flinty, 6-5 mm. long, 3-6 mm. broad, 3-25 mm. thick.

Closely similar Australian forms with semi-erect young shoots, completely

FIG. 179. — BREAD WHEAT (T. vulgare, Host).

i . var. lutescens.
(Persia.)

2. var. albidum.
(Semiretchensk .)

COMMON BREAD WHEAT 291

beardless ears with rigid truncate empty glumes resembling those of T. Spelta
are Warren, Thew, Comeback, and Jonathan, the latter with more hairs on
the young leaf-surfaces and somewhat denser ears (D = 24).

A form received from Rhodesia under the name Nobbs is similar to these.

8. White Victoria. — A mid-season form received from England and France.
It comes into ear at Reading about June 10.

Young shoots, semi-erect.

Straw, tall, 127 cm. (about 50 inches) high.

Ear, somewhat lax, 10 cm. long with a few awns, -5-1 cm. long near the apex ;
spikelets 22-25 '•> 0 = 22-25 (^ar tyPe T> Fig. 188).

Empty glume, 8 cm. long, apex rather narrow, apical tooth blunt, i mm.
long (Form 16, Fig. <66).

Grain, semi-flinty, plump, 5-9-6 mm. long, 3-7 mm. broad, 2-9 mm. thick.

Very similar is Blanc de Flandres from France with slightly longer awns
on the apical spikelets.

White Treasure from New Zealand, Hybrid Bordier from France, and
White Japhet from Cape Colony, South Africa, are somewhat similar forms
with larger grains (6-5 mm. long, 4-1 mm. broad, 3-7 mm. thick).

9. Hallett's Pedigree White. — An Old English mid-season form.
Young shoots, erect or semi-erect.

Straw, tall, 127 cm. (50 inches) high, pink.

Ear, 9 cm. long, slightly tapered with a few awns, -5 cm. long near the apex ;
spikelets 26 ; D =29-30 (Ear type i, Fig. 188).

Empty glume, 8 mm. long, apex narrow, apical tooth blunt, -5-1 mm. long
(7, Fig. 1 66).

Grain, semi-flinty, 6-25 mm. long, 3-9 mm. broad, 3-2 mm. thick.

10. Mansholt's Dikkop, No. 1. — A very late form from Holland.
Young shoots, prostrate.

Straw, stout, of medium height, 102 cm. (40 inches) high.

Ear, 9 cm. long with a few awns at the tip about -5 cm. long ; spikelets 23 ;
D =27 (Ear type 2, Fig. 188).

Empty glume, 9 cm. long, apex broad, truncate, apical tooth blunt, -5 mm.
long (7, Fig. 1 66, with broad apex).

Grain, semi-flinty, 6-85 mm. long, 3-8 mm. broad, 3-4 mm. thick.

11. Wilhelmina. — A late prolific form received from Holland ; grown also
in England.

Young shoots, prostrate.

Straw, of medium height, 96-102 cm. (38-40 inches) high.

Ear, dense, 7-8 cm. long, 15 mm. across the face, 12-14 mm. across the side ;
spikelets 24-26 ; 0 = 28-32 (Ear type 2, Fig. 189).

Empty glume, 8-9 mm. long ; apical tooth blunt, i mm. long ; lower flowering
glumes with short incurved tooth, apical flowering glumes with awns 1-1-5 cm-
long.

Grain, white, opaque, mealy ; 6-8 mm. long, 4 mm. broad, 3-3-2 mm. thick.

Similar to this is Willem I. from Holland, Victor, Stand-up, Essex Hybrid

292 THE WHEAT PLANT

from England, and Blanc a paille raide from France ; all come into ear at
Reading about the last week of June.

12. Million. — Received from Holland.

Young shoots, prostrate.

Straw, of medium height, 96-102 cm. (38-40 inches) high.

Ear, dense, 7-8 cm. long, 15 mm. across the face, 12-14 mm. across the side ;
spikelets 24-27 ; 0 = 34-38 (Ear type i, Fig. 190).

Empty glume, 8-9 mm. long, apical tooth blunt, i mm. long (7, n, Fig. 166) ;
flowering glumes of upper spikelets with awns '5-1*5 cm. long.

Grain, creamy-white, usually semi-flinty ; 6-8 mm. long, 4 mm. broad, 3-3
mm. thick.

White Monarch, Mark Lane, and Emperor, from England, closely resemble
this form. They are all very late prolific varieties, which do not come into ear
at Reading until about the last week in June.

Ear beardless ; glumes white, glabrous ; grain red.

T. vulgare, var. lutescens, Korn. Handb. d. Getr. i. 45 (1885).

This variety possesses the largest number of forms of any of the varieties
of T. vulgare.

About 250 forms were collected, examples being obtained from every wheat-
growing country. The majority, however, came from Western Europe,
Australia, and New Zealand, countries in which bearded varieties are not
esteemed.

Some of the forms have long lax ears, others are of the dense Squarehead
type, and between these extremes are found all grades of ear-length and density.

Similar gradation occurs among the numerous forms in respect of earliness
and lateness of growth, tallness and shortness of straw, and examples of this
variety are found in all the groups mentioned on pp. 270-272.

1. G-ranella de Carpegna. — An early form received from Spain, with similar
forms having empty glumes keeled to the base. These come into ear at Read-
ing at the beginning of June.

Young shoots, erect.

Straw, stout, of medium height, 112 cm. (about 44 inches) high.
Ear, very lax, 11-13 cm- l°ng 5 upper spikelets with awns 1-1-5 cm- ^On8 >
spikelets 20 ; D = 17 (Ear type i, Fig. 177).
Empty glume, 10 mm. long (6, 8, Fig. 166).
Grain, semi-flinty, 7 mm. long, 3-65 mm. broad, 3 mm. thick.

2. A common winter (Azima) wheat received from Russia with characteristic
velvety young leaves and empty glumes keeled to the base. It is early at
Reading, coming into ear during the first week of June.

Young shoots, erect.

Straw, somewhat slender, tall, 120-128 cm. (48-50 inches) high.
Ear, lax, 11-12 cm. long ; upper spikelets with awns i cm. long ; spikelets
19-21 ; D = 19-22 (Ear type 2, Fig. 181).

Empty glume, 10 mm. long, keeled to the base (Forms 5, n, 22, Fig. 166).

FIG. 1 80. — BREAD WHEAT (T. vulgar e, Host).

i. var. lutescens. 2. var. lutescens.

(Japan.) (Cedar.)

COMMON BREAD WHEAT 293

Grain, flinty, 7-3-7' 5 mm. long, 3-5 mm. broad, 3-15 mm. thick.
Similar forms were received also from Manchuria, Austria, Hungary, and
Germany.

3. Solid-straw Eed Tuscan. — A solid-strawed form of T. vulgare received
from New Zealand. It comes into ear at Reading about June 10.

Young shoots, erect or semi-erect.

Straw, slender, solid, of medium height, 96-102 cm. (38-40 inches) high.

Ear, lax, 9-10 cm. long ; upper spikelets with awns 1-2 cm. long ; spikelets
20 ; D =22 (Ear type i, Fig. 178).

Empty glume, 8 mm. long, keeled to the base, with broad apex ; apical tooth
i mm. long (2, Fig. 166).

Grain, mealy, 6-5 mm. long, 3-2 mm. broad, 3 mm. thick.

4. Frenisburg. — Received from Switzerland. It comes into ear at Reading,
June 10-11.

Young shoots, prostrate.

Straw, slender, tall, 125-135 cm. (48-52 inches) high.

Ear, lax, 9-10 cm. long ; upper spikelets with awns i cm. long ; spikelets
22 ; D = 22 (Ear type i, Fig. 178).

Empty glume, 8 mm. long, keeled to the base (Forms 9, 15, Fig. 166).
Grain, flinty, 6« mm. long, 3-25 mm. broad, 2-95 mm. thick.

5. fled Fife. — A famous wheat grown extensively in Canada and the United
States. Its grain is of high quality for bread-making. The original plant was
derived from a sample sent by a friend in Glasgow to David Fife in Ontario,
Canada, about 1842. The sample is said to have been taken from a cargo
shipped from Dantzig to Glasgow. Fife wheat very closely resembles some
forms from Galicia and Western Russia, and probably came from this region.

Young shoots, prostrate.

Straw, slender, medium height, 95-115 cm. (about 38-45 inches).

Ear, lax, 10-11 cm. long, upper spikelets usually with awns 10-12 mm. long ;
spikelets 20-22 ; D = 18-20 (Ear type i, Fig. 178).

Empty glume, 9-10 mm. long, apex narrow, tooth blunt (6, 9, Fig. 166).

Grain, flinty, apex blunt, 6-5-6-8 mm. long, 3-4 mm. broad, 3-3 mm.
thick.

Marquis wheat is supposed to be derived from a cross between an Indian
early wheat and Fife. It closely resembles the latter, but its straw, ears, glumes,
and grain are slightly shorter (Ear type 2, Fig. 181). It ripens its grain from
four to seven days earlier than Fife, and on this account has supplanted the
latter wheat in districts liable to early droughts and frosts.

6. Gandum-i-Kaiseh. — A distinct and early form received from Persia. It
comes into ear at Reading about May 22.

Young shoots, erect.

Straw, of medium height, 91-96 cm. (36-38 inches) high.
Ear, lax, 9-11 cm. long ; upper spikelets with curved hook-like awns about
i cm. long ; spikelets 20 ; D = 21-22 (Ear type i, Fig. 179).

294 THE WHEAT PLANT

Empty glume, 9 mm. long, inflated, apex broad with apical tooth 1-2 mm.
long, bent inwards (2, 3, Fig. 166).

Grain, large, semi-flinty, 8-4 mm. long, 3-6 mm. broad, 3-5 mm. thick.

7. Frankenstein. — A late form received from Germany. It comes into ear
at Reading about June 23.

Young shoots, prostrate.

Straw, somewhat slender, tall, 125 cm. (about 49 inches) high.

Ear, narrow, 10-11 cm. long; upper spikelets with awns -5-1 cm. long;
spikelets 27 ; D = 26 (Ear type i, Fig. 181).

Empty glume, 9-10 mm. long, apex broad (22, Fig. 166).

Grain, flinty, 7 mm. long, 3-7 mm. broad, 3-2 mm. thick.

Closely resembling this form are the tip-bearded Saumur de Mars from
France and the English Trump.

8. Greldersche. — A late form received from Holland. It comes into ear
about June 22.

Young shoots, prostrate.

Straw, somewhat slender, tall, 115 cm. (45 inches) high.
Ear, 10-11 cm. long ; upper spikelets with awns 1-1-5 cm. long ; spikelets
24 ; D = 24 (Ear types i, Fig. 178 ; 2, Fig. 181).

Empty glume, 8 mm. long (Forms 20, 22, Fig. 166).

Grain, flinty, 6-5 mm. long, 3-25 mm. broad, 3-1 mm. thtck.

9. Cedar. — An Australian form.
Young shoots, semi-erect.

Straw, medium height, 90-102 cm. (36-40 inches) high, hollow with thick walls.

Ear, lax, rigid, 10-12 cm. long, 12 mm. across the face, 10 mm. across the
side ; flowering glumes quite awnless ; spikelets 20-22 ; D = about 20 (Ear type
2, Fig. 1 80).

Empty glume, 10 mm. long, apex truncate (n, Fig. 166).

Grain, flinty, apex blunt, 6-4 mm. long, 3-8 mm. broad, 3-2 mm. thick.

10. Hybrid Inversable. — A late form received from France. It comes into
ear at Reading about June 20.

Young shoots, semi-erect.

Straw, stiff, of medium height, 94 cm. (about 37 inches) high.

Ear, 9 cm. long; upper spikelets with awns 1-2-5 cm- l°ng; spikelets 20 ;
0 = 23 (Ear type i, Fig. 187).

Empty glume, 9 mm. long (7, Fig. 166).

Grain, semi-flinty, 7 mm. long, 4 mm. broad, 3-2 mm. thick.

Bon Fermier from France, Hadmersleben from Germany, and the English
Dreadnought are similar to this.

11. Trigo Regadio. — An early form received from Spain.
Young shoots, erect.

Straw, of medium height, 96-102 cm. (38-40 inches) high.

Ear, 8-9 cm. long ; upper spikelets with awns -5-1 cm. long ; awns or teeth
of the lower flowering glumes curved inwards ; spikelets 22 ; D = 23 (Ear type
i, Fig. 187).

FIG. 181. — BREAD WHEAT (T. vulgare. Host).

i . var. lutescens.
(Frankenstein.)

2. var. lutescens.
(Chidham de Mars.)

COMMON BREAD WHEAT 295

Empty glume, 8 mm. long (i, 2, Fig. 166).
Grain, semi-flinty, 7 mm. long, 4-1 mm. broad, 3-6 mm. thick.
A similar form was also received from Germany under the name Tiichtiger
Landwirth.

12. Red Japhet. — An early form received from France. It comes into ear
at Reading about June 9.

Young shoots, erect.

Straw, of medium height, 115 cm. (about 45 inches) high.
Ear, 9-10 cm. long ; apical spikelets with awns -5 cm. long ; teeth of lower
flowering glumes curved inwards ; spikelets 20 ; D = 22 (Ear type i, Fig. 187).
Empty glume, 8 mm. long, apex truncate (22, Fig. 166).
Grain, semi-flinty, 6-6 mm. long, 4-2 mm. broad, 3-85 mm. thick.
Resembling it is Bed Marvel.

13. Victoria. — A late form received from France. It comes into ear at
Reading, June 20-24.

Young shoots, prostrate.

Straw, tall, 135 cm. (about 52 inches) high.

Ear, 10 cm. long ; upper spikelets with awns not more than -5 cm. long ;
spikelets 24-26 ; D =26 (Ear type i, Fig. 188).

Empty glume, 8-9 mm. long (6, Fig. 166).

Grain, mealy, 6-3 mm. long, 4 mm. broad, 3-4 mm. thick.

Similar forms are Essex Conqueror, Street's Imperial, Kinver Red, Model
Red, and Swan.

14. Tystofte Standhvete. — A very late form received from Denmark. It
comes into ear at Reading about June 23.

Young shoots, prostrate.

Straw, of medium height, 102 cm. (about 40 inches) high.

Ear, 7-8 cm. long ; upper spikelets with awns 1-2 cm. long ; spikelets 19-20 ;
0 = 27 (Ear type 2, Fig. 188).

Empty glume, 8 mm. long (9, n, Fig. 166).

Grain, semi-flinty, 6-25 mm. long, 3-25 mm. broad, 3-0 mm. thick.

Somewhat similar are Tystofte Smaahvete from Denmark, Gneisendorf
from Austria, and English White Straw Red.

15. Svalb'f Grenadier. — A late form received from Sweden. It comes into
ear at Reading about June 24.

Young shoots, prostrate.

Straw, of medium height, 96 cm. (about 38 inches) high.

Ear, 9-10 cm. long ; upper spikelets with awns about i cm. long, the short
awns of the lower spikelets curved inwards ; spikelets 23 ; D = 25-27 (i, 2, Fig.
1 88).

Empty glume, 9 mm. long (n, Fig. 166).

Grain, semi-flinty, 6*6 mm. long, 3-7 mm. broad, 3-1 mm. thick.

Similar to this are Mettel's Squarehead and others from Germany and
Austria.

296 THE WHEAT PLANT

1 6. Squarehead. — A late, very prolific English wheat very widely grown
throughout Western Europe since 1870 on account of its stiff straw and high
yield. It comes into ear at Reading about June 20. Breymann (Landw. Jahrb.
786, 1878) was informed by Samuel D. Shirreff that it was discovered by a Mr.
Taylor in a field of Victoria wheat in Yorkshire about 1868. From 1870 it
was very extensively grown and sold by C. Scholey, Eastoft Grange, Goole,
Yorkshire, who is sometimes credited with its discovery. It was introduced
into Denmark by a pupil of S. D. Shirreff in 1874, and from thence into
Germany. Later it spread to Holland, Belgium, and France.

Previous to 1870, typical Squarehead wheats were grown as early as 1839
under the names Suffolk Thickset and New Eed Norfolk or Hickling's
Prolific, and specimens of these collected in 1839 and 1841 are in the Collection
of the Royal Agricultural Society of England (Nos. 136 and 245). Hickling's
Prolific was discovered in 1830 by Samuel Hickling of Cawston, Aylsham,
Norfolk, and continued to be grown for some considerable time. Whether the
more modern Squarehead of Taylor, Scholey, and Shirreff was a derivative of
Hickling's wheat, or a new and independent mutation or product of hybridisa-
tion cannot be determined.

Forms resembling Squarehead appear in the F2 and subsequent generations
of hybrids of lax-eared vulgar e wheats with T. compactum.

Squarehead.

Young shoots, prostrate.

Straw, of medium height, 105 cm. (about 41 inches) high.

Ear, dense, square, 8-5 cm. long ; upper spikelets with awns 1-1-5 cm- l°ng !
spikelets 24 ; D = 32 (Ear type 2, Fig. 189).

Empty glume, 9 mm. long (6, 9, Fig. 166).

Grain, semi-flinty, 6-5 mm. long, 3-6 mm. broad, 3-25 mm. thick.

Numerous selections have been made of Squarehead wheat in all the countries
previously named. These differ slightly in morphological characters, such as
form of empty glume, the length of the awn on the flowering glumes near the
apex of the ear, and in ear density.

Nearest the original form are Browick Greychaff, Leutewitz, and Beseler's
Squareheads from Germany. Criewener Squarehead from Germany and
Austria and Essex Squarehead have shorter awns (-5 cm. long). Several of the
Swedish Svalof Squareheads, Strube's Squarehead, and English Kent Payer,
which closely resemble each other, have denser ears (D = 36-37).

Ear beardless ; glumes white, pubescent ; grain white.

T. vulgare, var. leucospermum, Korn. Syst. Uebers. 10 (1873).
A widely distributed variety.

i. An early form received from the Punjab, the United Provinces, and other
parts of India. It comes into ear at Reading at the end of May.

Young shoots, erect.

Straw, slender, short to medium height, 88-102 cm. (35-40 inches) ; leaves
blue-green.

FIG. 182. — BREAD WHEAT (T. vulgare, Host).

i. var. milturum.
(Russia.)

2. var. milturum.
(Chubut.)

COMMON BREAD WHEAT 297

Ear, lax, 8-9 cm. long ; spikelets 17-19, spreading ; D = 18-22 (Ear types 2,
Fig. 180 ; i, Fig. 181).

Empty glume, thin, 9 mm. long, narrow at apex ; apical tooth acute, 1-2 mm.
long ; flowering glumes often have slender awns 5-10 mm. long (23, Fig. 166).

Grain, flinty, apex truncate, 6-6 mm. long, 3-65 mm. broad, 3 mm. thick.

A similar early form with somewhat denser semi-bearded ears (D = 2i,
awns at the apex of the ear up to 2 cm. long) was obtained from the Transvaal,
South Africa, under the name Wit Wol Koren.

2. A spring form received from Persia under the name G-andum Kamisheh.
Young shoots, erect.

Straw, slender ; leaves glaucous.

Ear, lax, 9-10 cm. long; spikelets 17, inflated; 0 = 19 (Ear type i, Fig.

179)-

Empty glumes, oval, 9 mm. long ; apical tooth curved, slender, 1-3 mm.

long ; flowering glumes of upper spikelets with tortuose awns 1-2-5 cm- l°n£
(Form 3, Fig. 166).

Grain, long, semi-opaque, with highly arched dorsal side ; 7-8 mm. long,
3-6 mm. broad, 3-7 mm. thick.

3. Pearl. — A winter wheat received from New Zealand.
Young shoots, prostrate.

Straw, tall, 112-118 cm. (44-47 inches) high.

Ear, 8 cm. long; spikelets 21-23; 0 = 27-28 (Ear type i, Fig. 188, but
shorter).

Empty glume, 8 mm. long ; apical tooth blunt, i mm. long ; flowering
glumes of apical spikelets with awns 5-12 mm. long (Form 9, Fig. 166).

Grain, short, plump, semi-flinty; 5-8 mm. long, 3-65 mm. broad, 3-0 mm.
thick.

Closely similar are Snowdrop and Velvet Chaff, also from New Zealand,
and Pudel wheat from Sweden.

4. Blanc a duvet from France is a winter form similar to Pearl in the length
of its ear and straw and in the habit of its shoots, but possesses a somewhat
narrower empty glume 10 mm. long, the apical tooth 1-1-5 mm. long (15, Fig.
166), and a more elongated flinty grain 6-15 mm. long, 3-5 mm. broad, 3-0 mm.
thick.

The English Old Hoary, Essex Rough Chaff, and Velvet Chaff White,
which were formerly much grown in this country, closely resemble this form.

5. Early spring forms with erect young shoots, narrow tapering ears 9-10
cm. long, 0=23-25, and grain 6-5 mm. long, 3-65 mm. broad, 3-55 mm. thick
have been obtained from Rhodesia, Australia, and New Zealand. They come'
into ear at Reading in the first week of June.

6. A distinct early form from Asia Minor.
Young shoots, erect or semi-erect.

Straw, short, slender, 80-90 cm. (about 32-36 inches) long.
Ear, lax, glaucous, with awned tips 8-5-9 cm- l°ng '•> spikelets 16-18 ; D =
about 20 (Ear type 2, Fig. 181).

298 THE WHEAT PLANT

Empty glume, 10 mm. long; apex broad; apical tooth blunt and short;
flowering glumes of upper spikelets with awns 1-2 cm. long (20, Fig. 166).
Grain, flinty, 7 mm. long, 3-5 mm. broad, 3-4 mm. thick.

Ear beardless ; glumes white, pubescent ; grain red.

T. vulgare, var. velutinum, Korn: Handb. d. Getr. i. 45 (1885).
A rare variety. Considerable areas of one or two forms of it are cultivated
in the United States.

1 . Forms were received from India with characters similar to those of No.
i, var. leucospermum, but with red grain.

2. Haynes' Blue Stem. — A winter form, received from the United States
and Australia.

Young shoots, prostrate or semi-erect.

Straw, soft, weak, of medium weight, short, 90-110 cm. (about 36-44 inches)
long ; leaves glaucous.

Ear, lax, 10-12 cm. long ; spikelets 19-23 ; D = 19-22 (Ear type i, Fig. 178,
but usually shorter).

Empty glume, 9 mm. long ; apical tooth -5 mm. long, blunt (21, Fig. 166) ;
flowering glumes of upper spikelets often have an awn 1-2-5 cm- l°ng-

Grain, flinty, 6-4 mm. long, 3-2 mm. broad, 2-9 mm. thick.

Similar late forms also from Switzerland, Holland, and Sweden. Jones'
Winter Fife, Silver King, and Oakley from the United States are also similar
but somewhat earlier, with shorter awns at the tip of the ear.

3. De Haie, and some selections and hybrids from Germany and Sweden
with shorter and denser ear (7-8 cm. long ; D =24-30), belong to this variety.

They are all late forms with prostrate young shoots and stout straw of
medium height.

Ear beardless ; glumes red, glabrous ; grain white.

T. vulgare, var. alborubrum, Korn. Syst. Uebers. 10 (1873).

A widely distributed variety chiefly grown in warm climates.

About fifty named forms were obtained, the majority from Spain, Russia,
Turkey, India, South Africa, Australia, New Zealand, Canada, and the United
States.

Most of these forms are Spring wheats with lax tapering ears, some of them
markedly " tip-bearded." One or two forms with much denser ears came from
Holland and France.

i. Punjab 21. — A very early form received from India. It comes into ear
at Reading about May 20.

Young shoots, erect.

Straw, slender, of medium height, 88-115 cm- (about 32-45 inches) high.

Ear, lax, narrow, 8-9 cm. long, square, about 10 mm. across the sides ; D = 20.

Empty glume, 10 mm. long, keeled to the base ; apical tooth acute, i mm.
long (5, Fig. 1 66) ; apex of flowering glume narrow, acute.

FIG. 183. — BREAD WHEAT (T. vulgare, Host).

var. milturum.
(Abyssinia.)

I

FIG. 184.— BREAD WHEAT (T. vulgare, Host).

i. var. lutescens.
(India.)

2. var. Delfii.
(Sind.)

COMMON BREAD WHEAT 299

Grain, flinty, with prominent dorsal ridge ; apex truncate ; 7-2 mm. long,
3-6 mm. broad, 3-2 mm. thick.

A somewhat similar form, but ten to fourteen days later with denser ears
(0 = 23), is Punjab 20, received from India.

2. John Brown. — An early form received from Australia. It comes into
ear at Reading about June 2.

Young shoots, erect.

Straw, stiff, 90-102 cm. (about 36-40 inches) high.

Ear, lax, 10 cm. long ; upper spikelets with awns -5 to -75 cm. long ; spikelets
18-20 ; D =20-22 (Ear type 2, Fig. 180, with short awns).

Empty glume, 8 mm. long ; apex broad, truncate ; apical tooth -5 mm. long,
blunt (n, Fig. 166).

Grain, long, flinty, 7-3 mm. long, 3-4 mm. broad, 2-8 mm. thick.

Similar to this is Darling from the Transvaal.

3. Xeixa. — An early form received from Spain. It comes into ear at
Reading about June 8.

Young shoots, semi-erect.

Straw, somewhat slender, medium to tall, 122-132 cm. (48-52 inches) high.

Ear, tip-bearded, lax, 1 1 cm. long ; upper spikelets with awns i'5~3 cm. long ;
spikelets 19-20 ; D = 17-20 (Ear type i, Fig. 178).

Empty glume, 8 mm. long, keeled to the base ; apical tooth i mm. long (21,
Fig. 1 66).

Grain, flinty ; dorsal ridge somewhat prominent ; 7-7*3 mm. long, 3-5 mm.
broad, 3-55 mm. thick.

Very similar to this but with shorter awns at the apex of the ears are Nobbs,
G-luyas, and Early G-luyas from Australia, Union C from the Transvaal, and
Odessa from France and Spain. The Australian and Transvaal forms are
earlier than the rest, coming into ear at Reading about the end of May.

A somewhat . denser-eared series of prostrate forms (0 = 22-23) are Early
Windsor, Fortyfold, Prize Taker, Superlative, Mogul, and Abundance from
Canada, Bianchetta from the Cape, and Kizildaly from Turkey.

4. Red Federation. — An early form received from Australia. It comes
into ear at Reading about May 26.

Young shoots, erect or semi-erect.

Straw, stiff, short, 80-90 cm. (about 32-36 inches) high.
Ear, 10-12 cm. long ; spikelets 21-23 ; D = about 21 (Ear type i, Fig. 181).
Empty glume, coriaceous, 10 mm. long, somewhat inflated ; apical tooth 5
mm. long (10, n, Fig. 166).

Grain, flinty, 6-25 mm. long, 3 mm. wide, 2-8 mm. thick.

5. A mid-season form received from Russia. It comes into ear at Reading
about June 15.

Young shoots, prostrate.

Straw, slender, tall, 121 cm. (about 48 inches) high.

Ear, narrow, lax, 11-12 cm. long; upper spikelets with awns 1-3 cm. long ;
spikelets 23 ; D = 2o.

300 THE WHEAT PLANT

Empty glume, 8 mm. long, keeled to the base ; apex truncate ; apical tooth
•5 mm. long, blunt.

Grain, small, semi-flinty, 6 mm. long, 3-5 mm. broad, 3 mm. thick.

6. Dattel. — A mid-season to late form received from France. It comes
into ear at Reading about June 17.

Young shoots, prostrate.

Straw, stout, tall, 127 cm. (50 inches) high.

Ear, 10-11 cm. long ; spikelets 22 ; D = 2i (Ear type 2, Fig. 186).

Empty glume, 10 mm. long, narrow at apex ; apical tooth blunt, -5 mm. long.

Grain, flinty, 6-4 mm. long, 3-65 mm. broad, 3-1 mm. thick.

7. Sandomir. — A mid-season form received from Germany. It comes
into ear at Reading about June 17.

Young shoots, prostrate.

Straw, slender, tall, 112 cm. (about 44 inches) high.

Ear, 9-10 cm. long ; upper spikelets with awns -75-1 cm. long ; spikelets 21 ;
0 = 23 (Ear type 2, Fig. 181).

Empty glume, narrow, 8 mm. long ; apical tooth blunt, -5-1 mm. long (22,
Fig. 1 66).

Grain, flinty, 6-1 mm. long, 3-5 mm. broad, 3-3 mm. thick.

8. Dawson's Golden Chaff. — A mid-season form received from Canada. It
comes into ear at Reading about June 15, and is very susceptible to Yellow Rust.

Young shoots, prostrate or semi-erect.

Straw, somewhat slender, of medium height, 102 cm. (about 40 inches) high.

Ear, 8-9 cm. long; upper spikelets with awns 1-2 cm. long; spikelets 21 ;
D =26 (Ear type 2, Fig. 187).

Empty glume, 9 mm. long ; apical tooth -5 mm. long, blunt (n, Fig. 166).

Grain, flinty, 6-3 mm. long, 3-55 mm. broad, 3-1 mm. thick.

A similar form with darker red glumes was received from New Zealand and
from Germany.

9. White Matador Rood Kop. — A late form received from Holland. It
comes into ear at Reading about June 18.

Young shoots, prostrate.

Straw, stout, tall, 121 cm. (about 48 inches) high.
Ear, dense, 8 cm. long ; spikelets 22 ; D = 29 (Ear type 2, Fig. 188).
Empty glume, 8 mm. long; apex truncate ; apical tooth -5 mm. long, blunt
(5, 9, Fig. 166).

Grain, semi-flinty, 6-4 mm. long, 3-45 mm. broad, 3 mm. thick.

Ear beardless ; glumes red, glabrous ; grain red.

T. vulgare, var. mifturum, Korn. Handb. d. Getr. i. 45 (1885).

This is one of the most widely distributed varieties of T. vulgare, and em-
braces a large number of forms. About 180 forms have been grown at Reading,
the largest proportion being obtained from Western Europe, although examples
are found in almost every wheat-growing country.

\

FIG. 185.— BREAD WHEAT (T. vulgar e, Host).

i . var. Delfii.
(Sonora.)

2. var. milturum.
(Hongrie rouge.)

f.

FIG. 1 86.— BREAD WHEAT (T. vulgare, Host).

i. var. milturum.
(China.)

2. var. milturum.
(Lammas.)

COMMON BREAD WHEAT 301

Forms with lax and moderately dense ears are equally common ; those with
the Squarehead type of ear are much less numerous in this variety than in the
vars. erythrospermum and albidum.

1. Gallego rapado. — An early form with very lax tip-bearded ears and stiff
glumes, received from Portugal. It comes into ear at Reading about the end of
May.

Young shoots, erect.

Straw, strong, tall, 112 cm. (44 inches) high.

Ear, very lax, 10 cm. long, almost square with short inflated spikelets, often
irregularly arranged on the rachis, those near the apex having awns 3-4 cm.
long ; D = 17 (Ear type 2, Fig. 178).

Empty glume, 7 mm. long, strongly keeled to the base ; apex truncate ; apical
tooth blunt, -5-1 mm. long (2, Fig. 166).

Grain, flinty or semi-flinty, plump, dorsal arch prominent ; 7 mm. long, 3-75
mm. broad, 3-6 mm. thick.

2. Punjab 22. — A very early form received from India. It comes into ear
at Reading, May 20 to 25.

Young shoots, erect.

Straw, slender, short to medium height, 90-115 cm. (about 35-45 inches) ;
leaves generally yellow-green.

Ear, 8-9 cm. long, spikelets frequently arranged irregularly on the rachis ;
upper spikelets with awns i cm. long; spikelets 17 ; D = 18-20 (Ear type 2,
Fig. 181).

Empty glume, rigid, 8 mm. long, keeled to the base ; apex somewhat broad ;
apical tooth acute, 1-1-5 mm- l°ng (5» Fig. 166).

Grain, flinty, 6-8 mm. long, 3-5 mm. broad, 3-1 mm. thick.

Several similar forms ranging in density from 17 to 25 were obtained from
Northern India (Cawnpo're and the Punjab) ; also from Egypt under the name
Selected Hindi.

3. Bordeaux. — An early form received from France, Spain, and Austria,
and from Argentina under the name French. It comes into ear at Reading
about June 7.

Young shoots, erect.

Straw, stout, medium to tall, 112 cm. (45 inches) high ; leaves glaucous.

Ear, 9-5-10 cm. long, 15 mm. across the face, 10 mm. across the side ; upper
spikelets with awns 1-1-5 cm- l°ng '•> spikelets 19-20 ; D = 19-22 (Ear type 2,
Fig. 177).

Empty glume, stout, 8-9 mm. long, apex truncate, keeled to the base ; apical
tooth blunt, -5-1 mm. long (2, 7, Fig. 166).

Grain, flinty, apex blunt, dorsal side arched ; 6-6 mm. long, 4 mm. broad,
3-65 mm. thick.

Rinipau's Schlanstedt Summer wheat from Germany is similar.

4. Little Joss. — A prolific form raised by Professor Biffen at Cambridge.
It was obtained from the cross Squarehead's Master x Ghirka.

Young shoots, prostrate.

302 THE WHEAT PLANT

Straw, moderately stout, medium to tall, 100-1 15 cm. (about 40-45 inches)high.

Ear, 10-11-5 cm. long ; spikelets broad, 22-23 5 D =20-22 (Ear type 2, Fig.
177, with less tip beard).

Empty glume, about 10 mm. long, keeled to near the base ; apical tooth
blunt (7, 22, Fig. 1 66).

Grain, large, semi-flinty, somewhat narrow at the apex ; 7 mm. long, 3-9 mm.
broad, 3-45 mm. thick.

5. Touzelle rouge de Provence. — A widely distributed form received from
France. It comes into ear at Reading about June 1-9.

Young shoots, prostrate.

Straw, slender, tall, 120-140 cm. (48-55 inches) high.

Ear, 10-11 cm. long, tapering towards the apex and the base ; upper spike-
lets with awns 1-1-5 cm- l°ng> 2 or 3 Dasal spikelets often barren ; spikelets
21-23 ; D = 20-25 (Ear types i, Fig. 178 ; 2, Fig. 181).

Empty glume, 8-9 mm. long, keeled from tip to base ; apex truncate ; apical
tooth blunt, -5-1 mm. long (2, 21, Fig. 166).

Grain, flinty or semi-flinty, 6-35-7 mm. long, 3-45-3*75 mm. broad, 3- 1-3-3
mm. thick.

Resembling it are Winckler's and Horner's Hochfeldt and Utendorf from
Switzerland, and a few forms from Austria and Spain.

Closely similar, but with ears somewhat tapering at the apex and narrower
spikelets, are a large number of forms from Russia, Argentina, and Hungary,
Stanley from Canada, and one or two from the United States.

6. A common form included among wheat sent from Russia under the name
Ulka and North Russian.

Young shoots, semi-erect.

Straw, slender, of medium height, 90-105 cm. (35-42 inches).
Ear, slender, tapering ; many barren spikelets at base and apex, 10-11-5 cm-
long ; spikelets 22-25 ; 0 = 22-25 (^ar tyPes *> 2, Fig. 182).
Empty glume, 8 mm. long, keeled to the base (8, 9, Fig. 166).
Grain, semi-flinty, 6-7 mm. long, 3-45 mm. broad, 3 mm. thick.
Forms similar to this are frequent in some parts of Argentina.

7. Golden Drop. — A late Old English form coming into ear at Reading
about June 1 6 to 20.

Young shoots, prostrate.

«S^nm;,somewhatslender,tall, 120- 130 cm. (48-5 2 inches) high; leaves glaucous.

Ear, 10-11 cm. long, 15 mm. across the face, 9-10 mm. across the side,
tapering towards the apex ; upper spikelets with awns i cm. long ; spikelets
23-25 ; D =20-24 (Ear type 2, Fig. 186).

Empty glume, 9 mm. long ; apex somewhat truncate ; apical tooth blunt, i
mm. long (Form 20, Fig. 166).

Grain, flinty or semi-flinty, 5-85 mm. long, 3-5 mm. broad, 2-95-3 mm.
thick.

Very similar are a large number of forms which have been received from
England, France, Germany, Austria, Sweden, Holland, Switzerland, and Russia.

FIG. 187. — BREAD WHEAT ( T. vulgare, Host).

i . var. lutescens.
(Japhet.)

2. var. cyanothrix.
(Persia.)

FIG. 188. — BREAD WHEAT (T. vulgare, Host).

i. var. lutescens.
(Partridge.)

2. var. milturum.
(Squarehead's Master.)

COMMON BREAD WHEAT 303

8. Hongrie Rouge. — A distinct, long-glumed early form received from
France. It comes into ear at Reading about the end of May.

Young shoots, prostrate.

Straw, stout, tall, 137 cm. (54 inches) high ; leaves yellowish-green.
Ear, 9-10-5 cm. long, 18-20 mm. across the face, 22 mm. across the side ;
spikelets 25, large, overlapping each other ; D = 24 (Ear type 2, Fig. 185).
Empty glume, n mm. long, apical tooth blunt, short.
Grain, flinty, 6-5 mm. long, 3-75 mm. broad, 3-1 mm. thick (7, Fig. 166).

9. Teverson. — A prolific late wheat, grown in England and parts of France
and Germany. It comes into ear at Reading about June 18.

Young shoots, prostrate.

Straw, stout, tall, 127 cm. (about 50 inches) high ; leaves glaucous.

Ear, dense, 8-9 cm. long, 15 mm. across the face, 10 mm. across the side ;
upper spikelets with awns about i cm. long ; spikelets 22-24 5 D = 27-30 (Ear
type 2, Fig. 188).

Empty glume, 9 mm. long (6, 22, Fig. 166).

Grain, semi-flinty, plump, 6-8 mm. long, 3-75 mm. broad, 3 mm. thick.

Similar to it are Squarehead's Master, English Standard Red, and Red
Standup. Red Criewejier from Austria (glume form 7, Fig. 166) also resembles
these forms, but is about a week earlier.

Browick is a similar prolific wheat, with denser and somewhat shorter ears.

It was found in 1844 among Annat wheat by R. Banham on his farm* at
Browick, near Wymondham, Norfolk.

The name Browick is sometimes erroneously applied to Squarehead wheat,
a form of var. lutescens of similar ear-density.

10. Kwang T'ou mai. — A distinct form received from Chungking, China.
It yields a large number of small grains and is very early, coming into ear at
Reading about May 20.

Young shoots, erect.

Straw, stiff, of medium height, 85-95 cm- (33"3^ inches) high ; leaves
yellowish-green.

Ear, dense, 8-5 cm. long, 15 mm. across the face, 10 mm. across the side ;
spikelets 22, well filled, frequently containing 5 grains in each (Ear type i, Fig.
186). In some ears the density is uniform (D = 23), in others the lower half is
lax and the upper part densely crowded (i, Fig. 189).

Empty glume, thin, broad, 8-9 mm. long, with prominent keel from apex to
base and a fine awn 3-5 mm. long (3, 5, Fig. 166).

Grain, dark red with bluntish apex, flinty, small ; 5-7 mm. long, 3-6 mm.
broad, 2-8 mm. thick.

Closely allied to this form is Kyu Shu from Japan.

Ear beardless ; glumes red, pubescent ; grain white.

T. vulgare, var. Delfii, Korn. Handb. d. Getr. i. 46 (1885).
A comparatively rare variety confined to warm countries, being obtained
only from India, Khorasan (Persia), Egypt, South Africa, and California.

3o4 THE WHEAT PLANT

All the forms are very early with the erect spring habit, and belong to Groups
I., II., and V.

The Indian forms have yellowish-green leaves, slender straw, and lax ears
(D = 17-21), with the spikelets sometimes arranged irregularly on the rachis.

The South African and Californian forms are similar, but have denser ears
(D= 22-27).

The Persian forms are distinct from these, having bluish-green leaves, taller
straw and ears with broader inflated glumes, and tortuose or hooked awns
often 1-2 cm. long.

1. Sonora. — An early form received from California. It comes into ear
at Reading at the end of May.

Young shoots, erect.

Straw, slender, of medium height, 90-102 cm. (about 36-40 inches) high;
leaves pale yellowish-green.

Ear, lax, 9-10 cm. long, 15 mm. across the face, 10-12 mm. across the side ;
spikelets 21, spreading ; D = 22-26 (Ear type i, Fig. 185).

Empty glume, 8 mm. long ; apex narrowed ; apical tooth acute, 1-5 mm. long ;
flowering glumes with awns 3-8 mm. long (15, Fig. 165).

Grain, flinty, 6-2 mm. long, 3-65 mm. broad, 3-15 mm. thick.

Closely similar forms with more hairy leaf-surfaces were obtained from
Rhodesia under the name Victoria, and from the Transvaal under the name
Rooi Wol Koren.

A form resembling Sonora, but with laxer ears (D = 20-22) and slender straw
about 114 cm. (45 inches) high, was sent from Bombay, the Punjab, and other
parts of India.

FromSind, India, I obtained two lax forms (D = 17-1 8) of var.Z)e//z/ with ears
lo-n cm. long, one of them possessing awns i'5-2 cm. long on the flowering
glumes of the upper spikelets (Ear type 2, Fig. 184).

2. Kalkori. — Sent by Sir Percy Sykes from Khorasan, Persia, where it is
grown as a rain-fed or non-irrigated crop.

Young shoots, erect.

Straw, of medium height, 100 cm. (about 39 inches) high ; leaves bluish-
green.

Ear, lax, 9-10 cm. long ; spikelets swollen, 19-20, somewhat irregularly
arranged on the rachis ; D =21 (Ear type i, Fig. 179).

Empty glume, inflated, 8 mm. long, 4-5 mm. broad, more or less keeled to the
base ; apical tooth 2-3 mm. long, narrow and strongly curved inwards ; flower-
ing glumes with short tortuose and hook-like awns 1-2 cm. long (4, Fig. 165).

Grain, flinty, 6-5 mm. long, 3-5 mm. broad, 3-35 mm. thick.

Associated with it is a similar semi-bearded form with straighter awns 2-3

cm. long.

Ear beardless ; glumes red, pubescent ; grain red.

T. vulgare, var. pyrothrix, Korn. Handb. d. Getr. i. 46 (1885).
i. A very early form, received from the Punjab, the United Provinces, and
other parts of India. It comes into ear at Reading about May 20.

V

FIG. 189. — BREAD WHEAT (T. vulgare, Host).

I. var. milturum.
(China.)

2. var. lutescens.
(Squarehead.)

COMMON BREAD WHEAT 305

Young shoots, erect.

Straw, slender, short, 76-81 cm. (30-32 inches) high, hollow with thickish
walls ; leaves generally pale yellowish-green.

Ear, lax, 9-10 cm. long ; spikelets 17-19, spreading, often irregularly arranged ;
D = 19-21 (Ear type i, Fig. 185).

Empty glume, firm, 9-10 mm. long, keeled to base, apex narrow ; apical tooth
1-5 mm. long, narrow, curved ; flowering glumes with incurved awns 5-8 mm.
long (5, 12, Fig. 166).

Grain, flinty, 7-35 mm. long, 3-4 mm. broad, 3-45 mm. thick ; resembles the
grain of T. durum.

2. Seigle. — A winter form received from France, Germany, and Austria.
Young shoots, prostrate or semi-erect.

Straw, stout, tall, 114-127 cm. (45-50 inches) high.

Ear, 10-12 cm. long ; spikelets 23, spreading ; D =23 (Ear type i, Fig. 185).

Empty glume, 9 mm. long, apex narrowed ; apical tooth blunt, -5 mm. long
(22, Fig. 1 66).

Grain, flinty, dorsal ridge somewhat prominent ; 6-6 mm. long, 3-7 mm.
broad, 3-4 mm. thick.

A similar form also from New Zealand under the name Golden Gem.

3. Fox. — A selected sport from Ble a duvet, a French white velvet-chaffed
form of var. leucospermum.

It tillers extensively, and is especially suited to well-drained wheat soils.
Young shoots, prostrate.

Straw, short, 75-90 cm. (about 30-36 inches) high.
Ear, tapering, 8-9 cm. long ; spikelets 19-22 ; 0=25-27.
Empty glume, 8-9 mm. long, apex narrowed ; apical tooth blunt, i mm. long
(14, 15, Fig. !66).

Grain, semi-flinty, 6-2 mm. long, 3-6 mm. broad, 3-1 mm. thick.

4. Chinese. — A very early form received from Chungking, China. It
comes into ear about May 20 Forms with denser, shorter ears (8-9 cm. long ;
D = 23) have sported from it.

Young shoots, erect.

Straw, stout, of medium height, 84-96 cm. (33-38 inches) high ; leaves pale
greenish-yellow.

Ear, lax, 11-12 cm. long ; spikelets 20 ; D=*= 17-18 (Ear type i, Fig. 186).

Empty glume, oval, 9 cm. long ; apical tooth acute, -5 mm. long (4, 5, Fig.
1 66) ; flowering glumes sometimes with awns 5-10 mm. long.

Grain, flinty, small, with truncate apex ; 5-8 mm. long, 3-3 mm. broad,
3-0 mm. thick.

Ear beardless ; glumes blackish-yellow, glabrous.

T. vulgare, var. triste, Flaksb. Bull. App. Bot. Petrograd, iv. 2 (1911).
Flaksberger's type was obtained from Sunpan in the province of Sze-chwan,
China.

He describes (i) a spring form, var. triste Sunpani, with ears about 9 cm.

X

306 THE WHEAT PLANT

long, and short incurved awns; spikelets 14-20; 0 = 20-25; empty glume
inflated, yellow or yellow-blackish with dark brownish-black margins and
reddish grain, and (2) a winter form, var. triste, and with lax tapering ears (in
Bull. App. Bot. viii. 194, he states the grains of var. triste are white).

Ear beardless ; glumes bluish-black ; grain red.

T. vulgare, var. cyanothrix, Korn. Handb. d. Getr. i. 46 (1885).
A very rare variety represented only by one or two forms.

1 . Blue Velvet Chaff. — A winter late form received from Germany (Haage
and Schmidt). The bluish colour of the empty glumes is absent in some
seasons, the ear being then a foxy-red tint.

Young shoots, prostrate.

Straw, stout, tall, 127-140 cm. (50-55 inches) high ; glaucous.

Ear, tapering, 10-13 cm. long, spikelets 26-28; D = 22-24 (Ear type 2,
Fig. 187).

Empty glume, 9-10 mm. long, apex narrow ; apical tooth blunt (14, Fig. 166) ;
flowering glumes of the upper spikelets with awns 10-12 mm. long.

Grain, flinty, apex somewhat truncate ; 6-8 mm. long, 3-85 mm. broad, 3 mm.
thick.

2. A somewhat similar but earlier form received from Persia.
Young shoots, semi-erect.

Straw, stout, tall, very glaucous.

Ear, 8-10 cm. long, square, 10 mm. across the sides or 12 mm. across the
face, and 10 mm. across the 2-rowed side ; spikelets 18-20 ; D = 20 (Ear type 2,
Fig. 187).

Empty glume, 9-10 mm. long, apex truncate ; apical tooth blunt (10, n,'Fig.
1 66) ; upper half bluish-black, lower half yellow, in some seasons all yellowish ;
flowering glumes of the upper spikelets with awns 10-12 mm. long.

Grain, large, semi-flinty, 7-7-5 mm. long, 3-9 mm. broad, 3-5 mm. thick.

Ear beardless ; glumes black, pubescent.

T. vulgare, var. nigrum, Korn. Handb. d. Gert. i. 46 (1885).

This variety was only known to Kornicke through Seringe, who refers to it
in his Cereales europeennes (p. 139) as " Variation N. Touzelle Lammas (noire
veloutee)."

FIG. 190. — BREAD WHEAT (T. vulgare, Host).

i. var. milturum.
(Browick.)

2. var. anglicum.
(Benefactor.)

CHAPTER XX

CLUB, DWARF, CLUSTER, OR HEDGEHOG WHEAT

T. compactum, Host. Icon, et descr. Gram. Aust. iv. 4, t. 7 (1809).

T. vulgare compactum, Alef. Landw. Fl. 327 (1866).

T. sativum compactum, Hackel. Nat. Pfl. ii. 2, 85 (1887).

T. tenax, A. II., compactum, Asch. u. Graeb. Syn. ii. 686 (1901).

THE most ancient of wheats possessing grains loosely invested by the
chaff and readily separated by threshing belonged to this race, and were
widely grown by Neolithic man in many parts of Europe.

A remarkable wheat named by Buschan T. compactum, var. globiforme,
was common in the Neolithic and Bronze periods, being found in many
deposits of these ages in Hungary, Germany, Switzerland, Italy, Spain, and
Sweden. Only the grains are known ; these are small, more or less hemi-
spherical, resembling a miniature coffee-bean in shape, the apex blunt, and
furrow well defined ; the average dimensions about 4-6 mm. long, 3-4
mm. broad, and 3-3*3 mm. thick. Buschan's wheat appears to resemble
the Indian Dwarf wheat T. sphaerococcum (see p. 321) rather than T.
compactum.

More typical examples of T. compactum, with small oval or elongated
grains 5*5-7 mm. long, 3-4*5 mm. broad, like those of the common forms
of the present day, have only been found in the Neolithic and Bronze Age
deposits of Switzerland and Northern Italy.

Subsequently its cultivation decreased, its place being largely taken
by the larger-grained Bread wheat (T. vulgare), which became common
in the later prehistoric periods.

There is no evidence that T. compactum was known to the ancient
Egyptians, Greeks, or Romans.

The small-grained wheat referred to by C. Bauhin in 1623 (Ptnax
Theatri botanicf) as Triticum sylvestre creticum, and grown extensively
at that time in Crete, was doubtless a form of T. compactum.

As mentioned later, this wheat is now widely distributed more or less
sporadically among the longer-eared wheats, but entire crops of it are
rarely seen, a fact which may account for the somewhat remarkable

307

3o8 THE WHEAT PLANT

absence of it from English herbaria of the seventeenth and eighteenth
centuries in which representatives of most of the other races of wheat
are found.

In this species the ears are very short and dense, with stiff glumes
and well-filled spikelets closely packed and arranged almost at right
angles to the rachis. In some forms the ears are clubbed, the spikelets
being more crowded at the apex than the base, a character met with also
in T. vulgare.

These wheats are sometimes termed " Cluster " wheats and " Dwarf "
wheats, but the latter name is not particularly appropriate since many
of the forms possess tall straw.

In France the name " He'risson " (Hedgehog) is given to them ; in
Germany the beardless forms are designated " Binkel " wheats, the
bearded varieties being known as " Igel " wheats.

Host first gave them specific rank in 1807, and refers to their cultiva-
tion in Styria under the name " Binkel."

The closely packed, short, rigid ear is characteristic, but the morpho-
logical features of the leaves, culms, glumes, and grain indicate the closest
affinity with T. vulgare.

When grown from single ears many forms are quite constant ; others,
more especially those which possess " clubbed ears," i.e. dense at the
apex and laxer at the base, frequently exhibit extensive segregation, lax,
intermediate, and dense-eared forms, bearded and beardless with smooth
or pubescent chaff, being often found among their descendants.

The Club wheats are widely distributed at the present day, most
frequently mixed with various forms of T. vulgare or among T. durum
in Asiatic Russia ; pure crops of them are uncommon except in Central
Asia and the Pacific Coast States of North America.

In Asia they are prevalent in Turkestan and the western provinces of
the Transcaucasus and more sparingly in Siberia. I have obtained
forms also from China and Manchuria.

It is somewhat remarkable that true T. compactum, Host, is absent
from Persia and India, the examples attributed to this race by Howard
belonging to the very distinct race T. sphaerococcum, mihi (see p. 321).

In Europe they are met with in South-East Russia, Germany, France,
Italy, Switzerland, Spain, and Portugal, and were formerly grown in
England under the names " Club," " Cluster," or " Thickset " wheats.
I have also had specimens from Rhodesia, South Africa.
In North America they are cultivated on a somewhat extensive scale
in the States of California, Oregon, Washington, and Idaho.
In South America some forms are grown in Chili.
Most of the Club wheats resist frost, drought, and fungi well, and
grow freely on the poorer classes of soils ; they are chiefly spring forms,

CLUB, DWARF, CLUSTER, OR HEDGEHOG WHEAT 309

many of them exceptionally early ; only a few winter late-ripening forms
are known.

The straw is erect and usually stout, not liable to lodge, though a few
forms have soft weak stems. The ears are prolific in number of grains,
many of the spikelets ripening three or four each ; the yield in volume
or weight per acre is, however, comparatively low as the grains are in-
dividually of small size. The grains are firmly held by the glumes, a
character which renders these wheats particularly suited to cultivation in
districts where it is the practice to leave the crop on the field for some
considerable time before the harvesting operations can be completed.

The grain is soft or of medium hardness, its quality resembling that
of the softer varieties of Bread wheat (T. vulgar e).

GENERAL CHARACTERS OF T. compactum, Host.

The coleoptiles are pinkish or colourless.

In the majority of forms the young leaf-blades and shoots are of the
erect " spring " habit ; a few are semi-erect, others having the prostrate
" winter " habit.

The leaves are pale yellowish-green or glaucous blue-green, and the sur-
faces always more or less hairy, the arrangement of the hairs in most forms
resembling that of T. vulgare (Fig. 164) ; a few forms have leaves clothed
with soft hairs somewhat similar to those of T. turgidum and T. dicoccum.

The straws are erect, stiff, and generally hollow, with comparatively
thin walls, though one or two kinds have solid straws. They possess five
or six internodes above ground, and vary in height from 93 to 140 cm.
(35-55 inches).

The ears usually make their appearance from the side of the leaf-
sheath and not from the apex as in most wheats. They are short, stiff, and
compact, beardless or bearded, from 3-5 to 6 cm. in length, the majority
being about 5 cm. (2 inches) long with 17 to 25 closely packed spikelets,
the density of the ear being 40-50 or more.

Typical ears are of uniform density and oblong or somewhat oval in
outline as viewed from the 2-rowed side ; others, especially those of
hybrid origin, are " clubbed," the spikelets being crowded towards the
apex and loosely arranged at the base. In section the ears are square or
more frequently oblong, the width across the imbricated face being
usually 10-12 mm., and across the 2-rowed side 13-20 mm.

The rachis is fringed with short hairs along the side and across the
upper part of each notch immediately beneath the points of insertion of
the spikelets.

The spikelets possess 6-7 flowers, 3-5 of which frequently produce
grain ; they are 10-13 mm- l°ng> I3~I5 mm- broad, and about 3 mm. thick.

The empty glumes are whitish-yellow, red, dark brown or blue-black,

310

THE WHEAT PLANT

and either glabrous or clothed with soft hairs. Those of the lateral
spikelets are 8-9 mm. long, 5- to 6-nerved, and unsymmetrical, measuring
from 3 to 4 mm. from the midrib to the outer margin, and 1-1-5 mm- to tne
inner margin. In beardless varieties the apical tooth of the empty glume
is bluntish and from -5 to 1-5 mm. long ; in bearded varieties it is generally
prolonged into a fine awn usually not more than 5 mm. long, though
occasionally reaching a length of 3 cm. In most forms the midrib or keel

IMJ

9 10 ii 12 13 14

FIG. 191. — Empty glumes of Club wheat (T. compactuni) ( x 2).

is not prominent except in the upper half of the glumes, the basal half
of the latter being rounded with only a slight indication of the central
nerve ; some Asiatic forms, however, are keeled from the tip to the base.
The various forms of empty glume are illustrated in Fig. 191.

The flowering glumes are inflated, about 100 mm. long, 7- to 10-
nerved, the parts covered by the empty glume being thin and pale in
colour ; in bearded varieties the flowering glumes terminate in scabrid
awns 5-9 cm. long, which in some forms diverge widely.

The palea is of the ordinary bicarinate form.

CLUB, DWARF, CLUSTER, OR HEDGEHOG WHEAT 311

The grains, which are white, yellowish, or red, are small, oval, narrow
towards the apex and plump, the cheeks right and
left of the shallow furrow usually well rilled
(Fig. 192). In some forms they possess a prominent
dorsal hump similar to that found in T. turgidum.

The endosperm in the majority of forms is opaque ^ i, f

and starchy, but in some it is flinty. ^

In well-developed grains the breadth is usually ^ 9

greater than the thickness.

Ml MM

IIIMM

FIG. 192. — Grains of Club wheat (T. compac-
tum), front, back, and side views (nat. size).

* . , FIG. 193.— Grains of the
Below are given measurements ot grains taken spikelets of one side

from the spikelets near the middle of the ear of of an ear of Club wheat

. . (1 . compactum) mat.

twenty-six varieties, size).

Length.

Breadth.

Thickness.

mm.

mm.

mm.

Average
Limits
Ratio ....

6-19

5.7-7-0
IOO

S'31
2-9-3'75
53-47

3-J3

2-85-3-65
50-46

I. Ears bearded —

VARIETIES OF T. compactum

Glumes white, glabrous.

a. Grain white .

b. Grain red

Glumes white, pubescent.

a. Grain white .

b. Grain red
Glumes red, glabrous.

a. Grain white .

b. Grain red

var. splendens, Korn.
var. icterinum, Korn.

var. sericeum, Korn.
var. albtceps, Korn.

var. Fetisowit, Korn.
var. erinaceum, Korn.

312 THE WHEAT PLANT

4. Glumes red, pubescent.

a. Grain white ...... var. rubriceps, Korn.

b. Grain red ...... var. echinodes, Korn.

5. Glumes blue-black, glabrous.

Grain red ...... var. atriceps, Korn.

6. Glumes blue-black, pubescent.

Grain red ..... var. atrierinaceum, Korn.

II. Ears beardless —

1. Glumes white, glabrous.

a. Grain white ...... var. Humboldtii, Korn.-

b. Grain red ..... var. Wernerianum, Korn.

2. Glumes white, pubescent.

a. Grain white ...... var. linaza, Korn.

b. Grain red ..... var. Wittmackianum, Korn.

3. Glumes red, glabrous.

a. Grain white ...... var. rufulum, Korn.

b. Grain red . . . . . . var. creticum, Korn.

4. Glumes red, pubescent.

a. Grain white ...... var. crassiceps, Korn.

b. Grain red . . . . . var. rubrum, Korn.

5. Glumes bluish-red, pubescent.

Grain red ...... var. clavatum, Korn.

6. Glumes black, pubescent.

Grain red ...... var. atrum, Korn.

Ear bearded ; glumes white, glabrous ; grain white.

T. compactum, var. splendens, Korn. Handb. d. Getr. i. 53 (1885).

T. vulgare splendens, Alef. Landw. Fl. 328 (1866).

T. vulgare, Q., Metzger. Eur. Cer. 10 (1824).

Kornicke received from Chili, under the names Trigo de la Vuida and
Trigo Carbillo, a form of this variety with short thrck ears, broader across the
side than the face, with straight spreading awns ; he also states that the variety
is met in North America and Italy.

Flaksberger records its occurrence in Semiretchensk, Semipalatinsk, and
Turkestan.

Ear bearded ; glumes white, glabrous ; grain red.

T. compactum, var. icterinum, Korn. Han'db. d. Getr. i. 53 (1885).

T. vulgare icterinum, Alef. Landw. Fl. 328 (1866).

T. vulgare, P., Metzger. Eur. Cer. 10 (1824).

One of the most widely distributed varieties of T. compactum, forms of it
being sent to me from France, Germany, Spain, Rhodesia, Turkey, Asiatic
Russia (Bokhara and Semiretchensk), Manchuria, and the United States.
Flaksberger mentions its occurrence in Siberia, Turkestan, the Caucasus, Amur,
and China, and states that it was formerly found in European Russia.

FIG. 194. — CLUB WHEAT (T. compactum, Host).

i. var. icterinum.
(Manchuria.)

2. var. icterinum.
(Central Asia.)

FIG. 195. — CLUB WHEAT (T. compactum, Host).

i. var. icterinum.
(Semiretchensk.)

2. ennaceum.
(Heterozygote ?)

CLUB, DWARF, CLUSTER, OR HEDGEHOG WHEAT 313

1. Herisson barbu. — An early spring form received from France, and from
the United States under the name Walla Walla.

Vilmorin records its cultivation in the eastern departments of France from
Lorraine to Provence. It tillers little, is adapted to spring sowing, and succeeds
in cold mountainous districts on poor soils.

Young shoots, erect.

Straw, tall, 135 cm. (53 inches) high, soft, hollow, usually pink when unripe ;
leaves glaucous.

Ear, 5-6 cm. long, square, 11-12 mm. across sides ; spikelets 20-23, often
4~ or 5-grained ; 0=40 ; awns slender, divergent, 7-9 cm. long (Ear type i,
Fig. 194).

Empty glume, 8 mm. long, apex narrowed with a slender awn -5-3 cm. long
(2, Fig. 191).

Grain, semi-flinty, 6-6-2 mm. long, 3-2-3-5 mm. broad, 3-3-3-5 mm. thick.

2. Early forms similar to (i) but with empty glumes as in 7, 8, Fig. 191, were
obtained from Asiatic Russia (Bokhara and Semiretchensk).

3. Winter forms also resembling (i), but later, were obtained from Turkey,
Russia, and from a sample of Manchurian wheat.

4. A very dense-eared early form, coming into ear at the end of May at
Reading, received from Spain mixed with Trigo Canute, a form of var. erinaceum

(P- 3i5)-

Young shoots, erect.

Straw, of medium height, 104 cm. (about 41 inches) high, stout, hollow.

Ear, 4-5-5 cm. long, 11-13 mm- across the face, 15 mm. across the side ;
spikelets 18-19 ; 0=48-50 ; awns 5-7 cm. long (Ear type i, Fig. 195).

Empty glume, 8-8-5 mm- long, apex narrow and terminated by a slender awn
•5-3 cm. long (5, Fig. 191).

Grain, flinty, 6-2-6-7 mm. long, 3-3-3 mm. broad, 3-15-3-25 mm. thick.

Ear bearded ; glumes white, pubescent ; grain white.

T. vulgare sericeum, Alef. Landw. Fl. 328 (1866).
T. compactum, var. sericeum, Korn. Handb. d. Getr. i. 54 (1885).
Kornicke states that this variety is only known in Botanic Gardens ; his
specimens arose from a cross.

Flaksberger records its occurrence in the Caucasus.

Ear bearded ; glumes white, pubescent ; grain red.

T. compactum albiceps, Korn. Handb. d. Getr. i. 54 (1885).

Kornicke's type was of hybrid origin.

Flaksberger mentions its occurrence in Semiretchensk and Semipalatinsk.

Ear bearded ; glumes red, glabrous ; grain white.

T. compactum, var. Fetisowii, Korn. Handb. d. Getr. i. 54 (1885).
Kornicke's type was sent to E. Regel, Petrograd, by Fetisow from Vernoe,
Central Asia.

314 THE WHEAT PLANT

This distinct variety I have only seen from Central Asia, where it appears to
be common ; all its forms are early, producing ears at the end of May at Reading.

Flaksberger records its occurrence in Siberia, Turkestan, and S.E. Russia.

Received from Semiretchensk and East Bokhara, Central Asia.

Young shoots, erect.

Straw, of medium height, about 90-100 cm. (36-40 in.) high, hollow, rigid.

Ear, 4-5-5-5 mm. long, 10-12 mm. across the face, 12-15 mm. across the
side, usually clubbed at the apex ; spikelets 17-19, 4- to 5-grained ; D =40-50 ;
awns brittle, 5-7 cm. long (Ear type i, Fig. 196).

Empty glume, 8 mm. long, usually narrowed towards the apex, with slender
awn 1-3-5 cm- l°ng m the Semiretchensk forms (i, Fig. 191) ; in some forms from
Bokhara the empty glume is broader at the apex with a short awn about 4 mm.
long (2, Fig. 191).

Grain, flinty, 6-4-6-6 mm. long, 3-3-3-7 mm. broad, and 3-1-3-6 mm. thick.

Ear bearded ; glumes red, glabrous ; grain red.

T. compactum, var. erinaceum, Korn. Handb. d. Getr. i. 54 (1885).

T. sativum erinaceum, Desv. From. 161 (1833).

Kornicke found this variety mixed with var. Fetisowii from Central Asia.

Flaksberger records its occurrence in Turkestan, Siberia, Caucasus, Mid-
and South-European Russia, and China.

I have received it from Spain, the Transvaal, Manchuria, and as segregates
from Chinese wheats and from Herisson barbu (var. icterinurri). A glaucous
form obtained from a sample of Manchurian wheat ; also among French samples
of Herisson barbu.

Young shoots, prostrate.

Straw, tall, 135 cm. (53 inches) high, somewhat slender, hollow.

Ear, 5-6 cm. long, 10-12 mm. across the face, 12-13 mm. across the side;
spikelets 21-22 ; D =40-43 ; awns slender, divergent, 7-8 cm. long (Ear types i,
Figs. 194, 195).

Empty glume, 7-8 mm. long, apex narrow with awn -5-2 cm. long (2-6, Fig.
191). This form has given semi-bearded segregates (2, Fig. 195) with broader
empty glumes.

Grain, yellowish-red, mealy or semi-flinty, the flanks of the furrow plump ;
6-1 mm. long, 3 mm. broad, 2-9 mm. thick.

2. An early form with erect young shoots and yellowish-green leaves and ears,
found among the progeny of a " clubbed " ear of T. vulgar e (var. milturum) from
Chungking, China.

Straw, short, 60-80 cm. (24-30 inches) long ; leaves yellowish-green.

Ear, dense and short, about 4 cm. long, 12 mm. across the face and side ;
spikelets 22-24, 4-grained ; D =48-50 ; awns 5 cm. long (Ear type i, Fig. 194).

Empty glume, 8 mm. long, keeled to the base, with a curved awn 5 mm. long
(2, 3, Fig. 191).

Grain, semi-flinty, often compressed a little on one side ; 2-5 mm. thick,
5 mm. long, 3 mm. wide.

FIG. 196. — CLUB WHEAT (T. compactum, Host).

i. var. Fetisozci. 2. var. Fetisowi.

(Central Asia.)

FIG. 197. — CLUB WHEAT (T. compactum, Host).

i. var. echinodes. 2. var. echinodes.

(China.)

CLUB, DWARF, CLUSTER, OR HEDGEHOG WHEAT 315

Trigo canuto. — A glaucous early form received from Spain mixed with var.
icterinum (p. 312). A form closely resembling it was received also from Bokhara.
They come into ear at the end of May at Reading.

Young shoots, erect.

Straw, of medium height, 102 cm. (40 inches) high, stout, hollow.

Ear, 5-6 cm. long, 12-13 mm. across the face, 13-14 mm. across the side ;
spikelets 20-22 ; D =40 ; awns of flowering glumes 5-6 cm. long, stiff, divergent
(Ear type i, Fig. 194).

Empty glume, 7 mm. long, with broad apex; apical tooth acute, 1-2-5 mm-
long (6, 8, Fig. 191).

Grain, flinty, apex bluntish with prominent dorsal hump ; 7 mm. long, 3-4
mm. broad, 3-35 mm thick.

Ear bearded ; glumes red, pubescent ; grain -white.

T. compactum rubriceps, Korn. Handb. d. Getr. i. 54 (1885).
Kornicke's type was obtained from a cross.
Flaksberger mentions its occurrence in the Caucasus.

Ear bearded ; glumes red, pubescent ; grain red.

T. compactum, var. echinodes, Korn. Handb. d. Getr. i. 54 (1885).

Kornicke's type was obtained from a cross.

Flaksberger states that it occurs in the Caucasus.

I have received it from China.

An early form obtained among the progeny of a " clubbed " ear of T. vulgar e
(var. milturum) from Chungking, China ; the leaves, stems, and ears before
ripening are a yellowish grass-green tint.

Young shoots, erect.

Straw, short, 60-80 cm. (24-32 inches) high, stout, hollow.

Ear, 4-4-5 cm. long, sometimes " clubbed," 15 mm. across the face, 20-22
mm. across the side ; spikelets 22 ; 0 = 50 ; awns stiff, 3-5-4-5 cm. long (Ear
types i, 2, Fig. 197).

Empty glume, 7-8 mm. long, narrow, with slender apical awn 5-8 mm. long
(4, 5, Fig. 191).

Grain, flinty or semi-flinty with prominent dorsal hump ; 5-8-6-5 mm. long,
3-1 mm. broad, 3-15 mm. thick.

Ear bearded ; glumes blue-black or dark brown, glabrous ; grain red.

T. compactum, var. atriceps, Korn. Handb. d. Getr. i. 54 (1885).

Known only to Kornicke from Seringe.

Flaksberger records its occurrence in Semiretchensk and Turkestan, and
mentions an unnamed variety with white grain from Turkestan.

The examples I possess originated from an ear of Herisson barbu (var.
icterinum} ; the colour of the glumes is not constant.

Young shoots, prostrate.

Straw, tall, 135 cm. (53 inches) high, somewhat slender, hollow, pink when
unripe.

316 THE WHEAT PLANT

Ear, 5-6 cm. long, square, 10-11 mm. across the face and side ; spikelets
20 ; D =40-45 ; awns divergent, 4-5 cm. long (Ear type i, Fig. 194).

Empty glume, 7-8 mm. long, narrow ; apical tooth curved inwards, acute, 2-4
mm. long (4, 5, Fig. 191).

Grain, flinty, plump ; furrow shallow ; 6 mm. long, 3-5 mm. broad, 3-1 mm.
thick.

Ear bearded ; glumes blue-black, pubescent ; grain red.

T. compactum, var. atrierinaceum, Korn. Arch.f. Biontologie, ii. 400
(1908).

Ear, long, strongly awned.

Like var. atrum (p. 319), this appears to have been derived from the crossing
of a velvet-glumed form of T. compactum with T. turgidum, var. iodurum.

Ear beardless ; glumes white, glabrous ; grain white.

T. compactum, var. Humboldtii, Korn. Syst. Uebers. Landw. Cer.

Poppelsd. 12 (1873).

Kornicke first obtained this type from a Botanic Garden ; later he received
specimens from Chili and California.

Flaksbergef records its occurrence in the Western Caucasus.

1. A common spring form in samples of Walla Walla wheat from the
United States.

Young shoots, erect or semi-erect.

Straw, of medium height, 96-104 cm. (38-41 inches) high, stout, striate,
hollow.

Ear, 5-6 cm. long, side view somewhat oval, 10-12 mm. across the face,
15-20 mm. across the side ; spikelets 23-24 ; 0=46-50 (Ear type i, Fig. 198).

Empty glume, 7-8 mm. long, apex narrowed ; apical tooth blunt, -5 mm. long
(12, 14, Fig. 191).

Grain, semi-flinty, with prominent dorsal hump ; 6-6-2 mm. long, 3-25 mm.
broad, 3 mm. thick.

2. A spring form from Chili with coarse glumes, glaucous ears of moderate
density, approximating to the " Squarehead " forms of T. vulgare.

Young shoots, erect.

Straw, of medium height, 104 cm. (30-40 inches) high, stout, hollow with
thickish walls.

Ear, 5-7 cm. long, "clubbed," 10-12 mm. across the face, 15 mm. across
•the side ; spikelets 18-22 ; 0 = 33-36 (Ear type 2, Fig. 198).

Empty glume, 8-9 mm. long, apex broad ; apical tooth blunt, -5-1 mm. long
(14, Fig. 191).

Grain, flinty, 6-6-7 mm. long, 3-3 mm. broad, 3 mm. thick.

3. Little Club. — A common glaucous form grown extensively in the Pacific
Coast States of America.

Young shoots, semi-erect.

Straw, of medium height, 104 cm. (41 inches) high, hollow.

FIG. 198. — CLUB WHEAT (T. compactum, Host).

i . var. Humboldtii.
(Walla Walla.)

2. var. Humboldtii.
(Chili.)

FIG. 199. — CLUB WHEAT (T. compactum, Host).

i . var. ruf ulum.
(Walla Walla.)

x 2. var. creticum.
(Mocho de espiga quadrata )

CLUB, DWARF, CLUSTER, OR HEDGEHOG WHEAT 317

Ear, 4-5-5 cm. long, square, 10-12 mm. across the face and side ; spikelets
16-18 ; D = about 40-45 ; upper spikelets often with awns i cm. long (Ear type
i, Fig. 199).

Empty glume, 8 mm. long, apex narrowed ; apical tooth blunt, -5 mm. long
(12, 14, Fig. 191).

Grain, mealy or semi-flinty, 6 mm. long, 3-3 mm. broad, 3 mm. thick.

Ear beardless ; glumes white, glabrous ; grain red.

T. compactum, var. Wernerianum, Korn. Handb. d. Getr. i. 57 (1885).
Kornicke obtained forms of this variety from California and Chili.
Flaksberger mentions its occurrence in Kharkov and other parts of Russia,
Turkestan, and China.

1 . Kubb. — A very late glaucous form of winter habit received from Professor
Eriksson, Stockholm, and from European Russia.

Young shoots, prostrate.

Straw, of medium height, 114 cm. (45 inches) high, stout, striate, hollow.

Ear, 5 cm. long, 12 mm. across the face, 15-17 mm. across the side ; spikelets
24 ; 0 = 50 (Ear type i, Fig. 200).

Empty glume, 7 mm. long, apex narrowed ; apical tooth blunt, -5-1 mm. long
(12, Fig. 191).

Grain, flinty, 5-9-6 mm. long, 3-55 mm. broad, 3-25 mm. thick.

2. An early form of this variety, more or less sterile in some seasons and with
longer ears than the previous form, was obtained from commercial samples of
Chilian wheat.

Ear beardless ; glumes white, pubescent ; grain white.

T. compactum, var. linaza, Korn. Handb. d. Getr. i'. 52 (1885).
This variety, according to Kornicke, is found in Chili under the name Trigo
linaza.

According to Werner's description the young leaves are dark green and erect.
Straw, short, 75-85 cm. (about 30-34 inches) high, stiff.
Ear, pale yellow, very dense and short ; spikelets 3 -grained, 13 mm. across.
Grain, pale yellow, mealy ; 6 mm. long, 3-5 mm. broad.

Ear beardless ; glumes white, pubescent ; grain red.

T. compactum, var. Wittmackianum, Korn. Handb. d Getr i 53
(1885).

Kornicke's type was obtained from a Botanic Garden.

Velvet Kubb.— A very late glaucous winter form received from Professor
Eriksson, Stockholm. Similar but earlier forms also among commercial samples
of Walla Walla wheats from the United States, and from Vilmorin, Paris, under
the name Chili velu.

Young shoots, prostrate.

3i8 THE WHEAT PLANT

Straw, tall, 114 cm. (45 inches) high, soft, hollow.

Ear, 5-0-6 cm. long, 12 mm. across the face, 15-17 mm. across the side ;
spikelets 23-24 ; 0=40-45 (Ear type i, Fig. 200).

Empty glume, 7-8 mm. long, apex narrowed ; apical tooth blunt, i mm. long
(Form 12, Fig. 191).

Grain, flinty with prominent dorsal hump ; 6-2 mm. long, 3-5 -3-75 mm.
broad, 3-5 mm. thick.

Ear beardless ; glumes red, glabrous ; grain white.

T. compactum, var. rufulum, Korn. Handb. d. Getr. i. 52 (1885).

Kornicke's type appears to have been a segregate from a cross.

A glaucous form received in a sample of Walla Walla wheat from the United
States.

Young shoots, semi-erect.

Straw, tall, 127 cm. (50 inches) high, hollow.

Ear, 6-6-5 cm- l°ng> tip with a few awns, 10-11 mm. across the face, 12-14
mm. across the side ; spikelets 22-25 '•> D = 35-40 (Ear type i, Fig. 199).

Empty glume, 7 mm. long, keeled to the base, apex broad ; apical tooth blunt,
•5-1 mm. long (7, Fig. 191).

Grain, semi-flinty, prominent on the dorsal side ; 6-9 mm. long, 3-4 mm.
broad, 3-5 mm. thick.

Ear beardless ; glumes red, glabrous ; grain red.

T. compactum, var. creticum, Korn. Handb. d. Getr. i. 52 (1885).

A widely distributed variety received from France, Germany, Portugal, and
China.

Flaksberger records its occurrence in Siberia, Semiretchensk, Turkestan, and
the Pamir Plateau.

1 . He"risson sans barbes. — A glaucous early form received from France.
Young shoots, semi-erect.

Straw, tall, 132 cm. (52 inches) high, stout, hollow.

Ear, short and very dense, 4-5 cm. long, 12 mm. across the face, 18 mm.
across the side ; spikelets 23, 4-grained ; D =48-54 (Ear type 2, Fig. 201).

Empty glume, 7 mm. long, apex broad ; apical tooth blunt, i mm. long
(9, Fig. 191).

Grain, semi-flinty, 6 mm. long, 3-2 mm. broad, 3-1 mm. thick.

2. A glaucous form cultivated in Madeira under the names Mocho de

O

espiga quadrata and Rapado de espiga quadrata is similar, but the apex of the
empty glume is narrower (14, Fig. 191) and the ear often has a few awns
1-1-5 cm- l°ng at tne tip (2» Fig. 199) I resembling this also is Ble carre de
Sicile from France and Sicilian wheat from Germany.

3. A grass-green early form among the progeny of a " clubbed " ear of T.
vulgare, var. milturum, from Chungking, China ; glumes keeled to the base (10,
Fig. 191).

FIG. 200. — CLUB WHEAT (T. compactum, Host).

i. var. crassiceps.
(Walla Walla.)

2. var. rubrum.

FIG. 201. — CLUB WHEAT (T. compactum, Host).

I. var. rubrum.
(China.)

2. var. creticum.
(H^risson sans barbes.)

CLUB, DWARF, CLUSTER, OR HEDGEHOG WHEAT 319

Ear beardless ; glumes red, pubescent ; grain white.

T. compactum, var. crassiceps, Korn. Handb. d. Getr. i. 53 (1885).

Kornicke's type was of hybrid origin.

I obtained an exceptionally early form of it in a sample of Walla Walla wheat
from the United States. Its ears appear about the end of May at Reading and
ripen about the last week in July.

Young shoots, erect.

Straw, tall, 130-142 cm. (51-56 inches) high, somewhat slender, hollow.
In the early stages of growth the leaves, straw, and ears are of a yellowish grass-
green tint.

Ear, 4-5-5 cm. long, square, 12-15 mm. across the face and side ; spikelets
18-20 ; 0=42-45 (Ear type I, Fig. 200).

Empty glume, 7 mm. long, apical tooth blunt, i mm. long (9, Fig. 191)

Grain, flinty, 5-6-5-9 mm. long, 3-2-3-6 mm. broad, 3 mm. thick.

Ear beardless ; glumes red, pubescent ; grain red.

T. compactum, var. rubrum, Korn. Handb. d. Getr. i. 53 (1885).

A glaucous-eared form received in a sample of Walla Walla wheat from the
United States.

Young shoots, prostrate.

Straw, tall, 130 cm. (51 inches) high, stout, hollow, pink when unripe.

Ear, 5-6 cm. long, 12 mm. across the face, 12-15 mm. across the side >
spikelets 20-22 ; D =40-42 (Ear type 2, Fig. 200).

Empty glume, 7 mm. long, narrow at apex ; apical tooth blunt, i mm. long
(9, Fig. 191).

Grain, flinty, 6-2-6-4 mm. long, 3-2-3-6 mm. broad, 3-15 mm. thick.

A short, dense-eared form of this variety with grass-green leaves and ears
4-4-5 cm. long (D =48-50) was obtained among the progeny of a " clubbed " ear
of T. vulgare (var. milturum) from Chungking, China (2, Fig. 200).

Ear beardless ; glumes bluish-red, pubescent ; grain red.

T. compactum, var. clavatum, Korn. Handb. d. Getr. i. 53 (1885).

Kornicke states that the type of this variety received by him from Hohenheim
has somewhat elongated narrow ears, slightly awned at the tip, with blue-black
glumes on a reddish ground like those of T. turgidum, var. dinurum (p. 257).

Ear beardless ; glumes blue-black, pubescent ; grain red.

T. compactum, var. atrum, Korn. Archiv f. Biontologie, ii. 398 (1908).

Kornicke's type with more or less quadrangular ears was derived from
specimens shown in 1883 in an Agricultural Exhibition by Bestehorn, who
crossed various wheats ; this variety was apparently the result of crossing some
form of T. compactum with T. turgidum, var. iodurum.

A glaucous form received in a sample of Walla Walla wheat from the United
States.

Young shoots, prostrate.

320 THE WHEAT PLANT

Straw, tall, 118 cm. (47 inches) high, stout, hollow.

Ear, 4-5-5 cm. long, sometimes " clubbed," 12 mm. across the face, 15-17
mm. across the side ; spikelets 20-22 ; 0 = 50 (Ear type i, Fig. 200).

Empty glume, blue-black, and clothed with pale hairs except near the base,
where it is yellowish-white like the unexposed portions of the flowering glume ;
7 mm. long ; apex narrow ; apical tooth curved, blunt, i mm. long (9, Fig. 191).

Grain, flinty, 5-7 mm. long, 3 mm. broad, 3-1 mm. thick.

CHAPTER XXI

' INDIAN DWARF WHEAT

T. sphaerococcum, mihi.

THIS race, which I received from India and parts of Persia, is referred
by Howard to T. compactum. Host, but the latter so far as I am aware does
not occur in India.

T. sphaerococcum resembles T. compactum only in the possession of
a short dense ear ; the straw, however, is stouter, the leaves shorter, more
erect and rigid, and the glumes different in form and texture. The grain
also is very small and the typical form is of characteristic hemispherical
shape, resembling that of the prehistoric T. compactum, var. globiforme, of
Buschan.

When grown side by side with numerous forms of T. compactum from
all parts of the world, its specific differences are strikingly evident.

It is cultivated in the Punjab and Central and United Provinces of
India, and I obtained examples of it among Kallak ( = little head) wheat
from Persia. Doubtless the same race is referred to by Duthie and Fuller
in Field and Garden Crops of the North- West Provinces of Oudh (1882)
as " a curious round-berried wheat named ' Paighambari,' apparently
an introduction from Arabia," but I have been unable to trace its occur-
rence in the latter country.

This race resists drought well and, according to Howard, is generally
grown with inundation moisture and little rain. The Indian examples
are resistant to Yellow Rust at Reading.

GENERAL CHARACTERS OF T. sphaerococcum, mihi.

The young plants have upright shoots ; young leaves as in T. vulgare.

At Reading the straw is short, averaging not more than 65-70 cm.
(24-28 inches), but in India it sometimes reaches a height of 3 feet or
more. It is hollow, very stiff and erect with 4 to 5 internodes, which in
some varieties are pinkish. The successive internodes of ten well-grown
stems measured from below upwards -5 cm., 5-2 cm., 13-0 cm., 17-7 cm.,

321 Y

322

THE WHEAT PLANT

and 31-5 cm. respectively. Below the ear the straw is frequently bent in
a sinuous fashion.

The culm leaf-blades are somewhat rigid, comparatively short, 10-16
cm. long, and 1-2-1-5 cm. broad, tapering quickly towards the tip, their
upper surfaces scabrid with a few coarse hairs on the ribs. The auricles
are long, narrow, and fringed with a few hairs.

The ears are short, generally measuring 4-6 cm. (about ii-2^ inches)
in length, with 14 to 20 spikelets ; density usually =38-42.

Along the edges of the rachis is a fringe of white hairs -25 to -5 mm.
long, which also extends across the front of the rachis at the base of each
spikelet.

ttl

• •

• * I

FIG. 202. — Empty glumes of Indian
Dwarf wheat (T. sphaerococcum)

(X2).

FIG. 203. — Grains of
Indian Dwarf wheat
(T. sphaerococcum),
front, back, and side
views (nat. size).

t f

FIG. 204. — Grains of
the spikelets of one
side of an ear of
Indian Dwarf wheat
(T. sphaerococcum}
(nat. size).

The spikelets are about 10 mm. long, 10 mm. across,
and 4 mm. thick, often with 6 or 7 flowers, 4 or 5 of
which may produce grain.

The empty glumes, which have an inflated appear-
ance, are 8-9 mm. long and 4 or 5 mm. across the outer
half. They possess 6 or 7 nerves, and are generally
keeled in the upper part only, though in some forms
the midrib is somewhat prominent below. At the
apex is a broad curved scabrid tooth (Fig. 202).

The flowering glume is inflated, 9-nerved, and rounded on the back ;
in the beardless varieties the tip terminates in a short awn 3 or 4 mm.
long, the bearded forms possessing very stiff awns 1-5 to 2 cm. long,
which are frequently bent in a sinuous manner near the base and in ripe
ears spread outwards irregularly.

The palea is 7-8 mm. long, fringed with hairs along the two keels.

The grains are of very characteristic form, being shorter and rounder
than those of other wheats (Figs. 203, 204). They are often somewhat
angular and unsymmetrical on account of pressure of the glumes.

FIG. 205. — INDIAN DWARF WHEAT (T. sphaerococcum, mihi).

i . var. echinatum. 2. var. rubiginosum.

INDIAN DWARF WHEAT 323

The apex is truncate with a short " brush," the furrow shallow.

The grains measure from 4 to 5-5 mm. long, 3~3-7 mm. broad, and 3~3-y
mm. thick ; the ratio of length : breadth : thickness = about 100 : 75 : 75.

T. sphaerococcum is a very early wheat, the ear escaping from the
upper leaf-sheath about May 20 at Reading.

VARIETIES OF T. sphaerococcum, mihi.
Ear bearded —

1. Glumes white, glabrous ; grain white . . var. echinatum, mihi.

2. Glumes white, glabrous ; grain red . . var. spicatum, mihi.

3. Glumes red, glabrous ; grain red . . . var. rubiginosum, mihi.

Ear beardless —

4. Glumes white, glabrous ; grain white . . var. tumidum, mihi.

5. Glumes white, glabrous ; grain red . . var. rotundatum, mihi

6. Glumes white, pubescent ; grain white . . var. globosum, mihi.

Ear bearded ; glumes white, glabrous ; grains white.

T. sphaerococcum, var. echinatum, mihi.

T. compactum, var. splendens, Howard and Howard. Wheat in India, 193
(1910).

This variety I received from the United Provinces, India, through H. M.
Leake, Esq. It is recorded from the Central Provinces by Howard and Howard.

Young shoots, erect.

Straw, stout, rigid, hollow, very short, at Reading 45-60 cm. (18-24 inches)
long.

Ear, erect, stiff, 4'5-6'5 cm. long, almost square, 10-11 mm. across the face
and side ; spikelets 15-23, 3- to 4-grained ; 0 = 35-40 (Ear type i, Fig. 205).

Empty glume, 4-6 mm. long, 3'5-4'5 mm. across, with a broad curved scabrid
tooth (i, Fig. 202) ; awns of flowering glumes spreading, stout, rigid, 1-2-3 cm-
long, often more or less curved.

Grain, very short, prominent on the dorsal side, flinty ; apex blunt, 4-5 mm.
long and 3 mm. broad.

Ear bearded ; glumes white, glabrous ; grain red.

T. sphaerococcum, var. spicatum, mihi.

T. compactum, var. icterinum, Howard and Howard. Wheat in India, 214
(1910).

Mentioned by Howard and Howard as a wheat of the Central Provinces, but
not described. I have not seen it.

Ear bearded ; glumes red, glabrous ; grain red.

T. sphaerococcum, var. rubiginosum, mihi.

T. compactum, var. erinaceum, Howard and Howard. Wheat in India, 178
(1910).

Received from India as " Punjab Type 4 " (2, Fig. 205).

324 THE WHEAT PLANT

In morphology and habit it closely resembles var. echinatum, but has dark
red glabrous glumes often with a glaucous bloom. The grain is small, pale
red, and flinty.

Ear beardless ; glumes white, glabrous ; grain white.

T. sphaerococcum, var. tumidum, mihi.

T. compactum, var. Humboldtii, Howard and Howard. Wheat in India, 179
(1910).

This variety I received from the United Provinces, India, and from the
Punjab as " Type 7 " (i, Fig. 206).

Young shoots, erect.

Straw, stout, very stiff, hollow, short, and pinkish especially in the upper
part, 60-72 cm. (24-28 inches) long at Reading.

Ear, erect, stiff, 4-5-6 cm. long, almost square, 10-12 mm. across the face
and side ; spikelets rigid, 15-21, 3- to 4-grained ; 0 = 32-45.

Empty glume, 7 mm. long, 4-5 mm. broad, with strong curved scabrid tooth
(3, Fig. 202).

Grain, small and short, flinty or mealy sometimes, often irregular from
pressure of glumes ; 5-5-5 mm. long, 3-3 mm. broad, 3-3 mm. thick.

Ear beardless ; glumes white, glabrous ; grain red.

T. sphaerococcum, var. rotundatum, mihi.

T. compactum, var. Wernerianum, Howard and Howard. Wheat in India,
179 (1910).

Briefly described by Howard and Howard as a Punjab wheat, " Type 6 "
(2, Fig. 206).

Ears, 4-4 cm. long, sometimes slightly bearded ; D = 39 ; glumes with pinkish
tinge and slightly longer curved keel tooth ; grain small, round, pale red, mealy.

Ear beardless ; glumes white, pubescent ; grain white.

T. sphaerococcum, var. globosum, mihi.

T. compactum, var. linaza, Howard and Howard. Wheat in India, 179
(1910).

Received from India as " Punjab Type 5 " (3, Fig. 206).

^oung shoots, erect.

Straw, short, erect, stiff, often pinkish and wavy beneath the ear ; about
61 cm. (24 inches) long at Reading.

Ear, 5-6 cm. long, almost square, 10-11 mm. across the face and side ; spike-
lets 13-16, well filled, 3- to 4-grained ; D = about 30.

Empty glume, 6-7 mm. long, 4 mm. broad with strong curved scabrid tooth.
(3, Fig. 202).

Grain, very short, dorsal hump prominent, often flattened on one side ;
5 mm. long, 3-5 mm. broad, 3-3 mm. thick.

t;.

FIG. 206. — INDIAN DWARF WHEAT (7\ sphaerococcumy mihi).

i. var. tumidum. 2. var. rotundatum. 3. var. globosum.

CHAPTER XXII

s

SPELT OR DINKEL

Triticum Spelta, L. Sp. PL 86 (1753).
Triticum Zea, Host. Gram. Austr. iii. 20, t. 29 (1805).
Spelta vulgaris, Scringe. Cer. eur. 76 (114) (1841).
Triticum vulgare spelta, Alef. Landw. Fl. 334 (1866).
Triticum sativum Spelta, Hackel. Nat. Pfl. ii. 81 (1887).

THE word spelt in the genitive singular — speltae — occurs first in an edict
of the Emperor Diocletian (A.D. 301), where it is given as synonymous with
the Greek oXvpa.

According to Hieronymus (A.D. 414), it is of Germanic origin, and was
used by the early inhabitants of the province of Pannonia (Hungary) for a
cereal which he considered the same as far of Latin authors. It is not
until the thirteenth century that a distinction is made between far and
spelta ; Crescentius, writing then, says that " far resembles spelt, but is
larger in the blade and grain " (Cres., lib. iii. cap. de Farre).

There is little doubt that it is much more recent than Emmer (T.
dicoccurri) and Small Spelt (T. monococcuni), for no remains of it have been
found of Neolithic Age and none of Bronze Age, with the exception of a
doubtful example mentioned by Heer from the pile-dwellings of the
latter period in Switzerland. It is absent from Egyptian tombs, and all
the evidence at present available points to the conclusion that it was not
known to the ancient Greeks and Romans. I regard it as a segregate of
a hybrid between a wheat of the Emmer series (p. 342) and Aegilops
cylindrica (see p. 343), probably first obtained by the ancestors of the
ancient Germanic races who settled in the districts north of the Alps
where the cereal is still grown.

At the present time in Italy Emmer is spoken of as farro or grano
farro, Common Spelt as spelta or spelda.

Of the wheats with a brittle rachis Common Spelt or Dinkel is the
race most extensively grown. The greatest area is devoted to this crop
in Germany, where in 1901 it was grown on 314,671 hectares (over 777,000
acres), the largest proportion being found in Wiirtemberg, Bavaria, and

325

326 THE WHEAT PLANT

Baden, in which districts it has been cultivated from the earliest times
in place of Bread Wheat (T. vulgar e). On a smaller scale T. Spelta is
grown in parts of Prussia, Hesse, and Alsace.

In 1900 over 39,000 hectares (96,330 acres) were grown in Switzerland
and about 5000 hectares (12,350 acres) in Austria.

It is also cultivated in considerable amount in the northern parts of
Spain (Asturias) and on small areas in Switzerland, France, and Italy.

It does not appear to be a farm crop in any other countries of Europe,
and I have no authentic record of its occurrence in Africa nor in Persia,
India, China, or other parts of Asia.

Spelt will grow wherever Bread Wheat will thrive ; it yields best on
good wheat land, but will succeed on soils which are too dry and light
for the commonly cultivated varieties of T. vulgare.

On land not quite suited to it, rye is sometimes mixed with it in the
proportion of A rye to 4 spelt.

It is one of the hardiest of cereals, being rarely affected by frosts
which destroy other wheats, and grows at all elevations, from about 300
to 3000 feet above sea-level in Germany and Switzerland.

Although it is slightly less productive than ordinary wheat and
possesses the disadvantage of brittle ears from which the true grain
cannot be obtained without special mills, it has advantages which enable
it to compete successfully with the more prolific forms of T. vulgare in
the districts where its cultivation is established. In respect of these
advantages its winter hardiness is of the greatest import. Stoll states that
after the winter of 1900-1901 38 per cent of the winter wheat of Germany
(about 1,800,000 acres) had to be ploughed up in May 1901 on account
of the damage done by frost, whereas less than i per cent of the Dinkel
sown (about 700 acres) was destroyed.

Other points in its favour are its greater resistance to smut, bunt, and
rust fungi, and freedom from the attacks of birds. Its straw is stiffer
than that of most ordinary wheats and the crop not easily " laid." In
addition, it can be harvested sooner than ordinary wheat, and the grain
in the ear does not sprout so readily when the reaped crop is left out in
wet seasons.

The majority of the kinds of Dinkel are winter forms, though a few
less hardy spring varieties are cultivated ; Emmer (T. dicoccum), however,
takes the place of T. Spelta where a spring " Spelt " is grown.

The former are sown at the end of September or during the first
week of October, while the spring kinds are sown in February and March.

Awned varieties are less frequently cultivated than the beardless
forms. Red-chaffed varieties are preferred, as these are hardier than
those with white chaff and possess stronger straw.

The viability of the grain is retained longest when it is enclosed in

SPELT OR DINKEL 327

the glumes and the " Vesen " or husked grain is almost always sown,
the germinating capacity of samples of the free caryopses being usually
not more than 30 or 40 per cent on account of the bruising sustained in
the process of removing them from the husk.

The seed is commonly broadcasted at the rate of 5*5-6-1 hectolitres
or 220 to 250 kg. of husked grain per hectare (about 190-220 Ibs. per
acre), the smaller amount being employed in the case of the spring
varieties which tiller well.

It is usually drilled at a depth of i| to 2^ inches in rows from 7 to 8
inches apart.

On account of the ease with which the ears break off the straw, the
crop is best cut with the sickle before the ears are fully ripe after the
straw is white. It is harvested during the first half of August in South
Germany. The yield is very variable ; in poor mountainous districts
it may be as low as 10-15 hectolitres per hectare (about 400-600 Ibs. per
acre) ; on better soils it varies from about 40 to 80 hectolitres per hectare
(1600-3200 Ibs. per acre), the average for Winter Dinkel in a good season
being about 40 hectolitres per hectare ; for spring-sown Dinkel 25 hecto-
litres per hectare is a fair crop.

The " Vesen " or pieces of the broken ears obtained by thrashing
the crop are subjected to a light grinding process in order to separate
the grain from the closely investing glumes, two rough sandstone mill-
stones without the usual grooves being used for the purpose. Where
the cultivation of. Dinkel is prevalent such mills are common, and the fact
that the growth of this cereal does not spread into new districts is attri-
buted to the absence of suitable apparatus for removing the grain from
the chaff rather than to the unsuitability of soil or climate.

The grain when ground yields a flour of medium gluten content
especially valuable to the confectioner and pastrycook. Some of it is
employed in the preparation of farinaceous foods and puddings, a little
only being made into bread. A flour suitable for the latter is sometimes
made by grinding Dinkel and Bread Wheat grain together.

A considerable amount of Dinkel grain is used in South Germany in
soups. When wanted for this purpose the ears are cut from the straw
when the grain has just passed the milk-ripe stage ; the ears are then
dried in an oven and after thrashing are sent to the mill for the removal
of the grain from the chaff.

Occasionally the grain in the husk is fed to horses, and the straw,
although somewhat stiffer and coarser than that of ordinary wheat, is
useful in a chopped state for horses or as long fodder for cattle ; the
chaff can be fed to cattle also, but it should not be given to horses except
in small quantities.

The following comparative composition of (i) Spelt " Vesen " or

328

THE WHEAT PLANT

grain in the husk as thrashed, (2) the true grains or caryopses, (3) straw,
and (4) chaff of Dinkel and Bread Wheat is given by Wolff :

Water.

Ash.

Crude

protein.

Fibre.

N. free
extract.

Fat.

Spelt Vesen .
Spelt grain

14-8
H'5

37
1-7

10-0
I3'5

16-5
i'S

52-3
67-2

i*S

1-6

Bread Wheat .

14-4

1-7

I3-0

3-0

66-4

i'5

Spelt straw
Bread Wheat straw

H-S
I4-3

5'0
5'3

2'5

4-5

45'0
37-8

3I-8
36-7

1-4

1-4

Spelt chaff
Bread Wheat chaff

H'3
I4-3

8'3
9-2

3-5
4-5

40-0
36-0

32-6

34'6

i'3

1-4

The grain has a thinner .pericarp than wheat and yields less bran.
Spelt flour has less sugar and dextrin and a correspondingly higher
starch content than that of Bread Wheat, the nitrogen -content being
similar in both cereals.

GENERAL CHARACTERS OF T. Spelta, L.

T. Spelta is one of the most distinct of the cultivated wheats, having
a characteristic lax ear and a broad truncate empty glume the lateral
nerve of which ends in a blunt secondary tooth some distance away from
the keel tooth.

The leaves of the young plants of autumn and early spring are dark
green, and except in the case of a black-bearded form (var. coeruleum)
lie close to the ground ; they are not much more than about half the
width of those of T. dicoccum of the same age, being only 3 to 4 mm.
across. The older culm leaves, however, are broader than those of
T. dicoccum. They are comparatively smooth, there being in most
varieties only a row of long hairs on the summit and a few smaller ones
on the flanks of the longitudinal ridges of the leaves, as in T. vulgar e
(Fig. 164).

The straw is erect, harder and stiffer than that of Bread Wheat, from
100 to 120 cm. (40-48 inches) high and hollow, with thin walls.

The blades of the upper leaves of the straw are from 12 to 20 mm.
across, pale greenish-yellow, almost glabrous ; the auricles are very large
and more or less fringed with long hairs.

The ears are straight or slightly curved, white, red, grey-blue or
blue-black, and bearded or beardless, relatively long and lax, the spikelets
being generally well separated from each other on the rachis.

They are usually from 10 to 15 cm. (4-6 inches) long, and square or
oblong in section. Each ear possesses from 16 to 23 spikelets, the
density (D) being 15 to 22.

SPELT OR DINKEL 329

The notched rachis, which is easily seen between the spikelets, is
smooth with more or less hairy margins, the frontal tuft being very small
or absent. It is convex on one side, and flat or slightly concave on the
other. It is also brittle, breaking into short pieces each 5-6 mm. long,
narrow and thin at the base, becoming much thicker and wider at the
apex.

The rachis at the point of attachment of each spikelet is broad and
often hollow and weak, and when thrashed breaks transversely at this
point more easily than at the narrow basal portion of the rachis internode,
which is the natural point of disarticulation so clearly seen in the fragile
ears of the wild wheats, and the cultivated T. monococcum and T. dicoccum.
In thrashed samples of T. Spelta each separate spikelet generally carries
with it the internodal portion above its point of insertion, whereas in

T. dicoccum and other fragile-eared
wheats the internode below it is
found attached to the thrashed
spikelet (Fig. 207).

Although the axis is fragile as

FIG. 207. — Spikelets (" Vesen ") of thrashed 4

ears of T. dicoccum (upper row) and T. FIG. 208. — Empty glumes of Spelt wheat
Spelta (lower row) (nat. size). (T. Spelta) ( x 2).

indicated it resists disarticulation proper, and on this account I include
this wheat among the tough-eared races.

The spikelets are oval, convex in section, each 'consisting of 3 or 4
flowers, two of which generally produce well-developed grains, except in
the case of 4 or 5 of the upper spikelets, which often ripen only one grain
each ; occasionally the central spikelets contain three grains.

The empty glumes are half boat-shaped, with a broad truncate apex
and ii nerves (Fig. 208). The narrow margins are membranous, the
broad outer part strong and coriaceous with 6 or 7 nerves, the narrow inner
side possessing only i or 2. The keel, which is less prominent than in
T. dicoccum, ends in a short blunt tooth. The strong lateral nerve of the
glume ends in a blunt projection, which is always much farther away
from the base of the apical tooth than is the corresponding secondary
tooth in T. dicoccum, T. turgidum, or T. durum.

33°

THE WHEAT PLANT

The flowering glume is boat-shaped and comparatively thin, with
9-11 nerves and no distinct keel. In some varieties it terminates in a
short claw-like projection 2-5 mm. long, in others in a scabrid stiff awn
6-8 cm. long ; sometimes the flowering glume of the
third imperfect flower bears a short awn 2 cm. long.

The palea is about as long A.

as the flowering glume, oval,
blunt, and bicarinate, the two ^ A

keels being fringed with short
hairs.

The caryopses, which usually
have flinty endosperm, are red-
dish, long and pointed at both
ends, the apex covered with a
conspicuous brush of whitish
hairs (Figs. 209, 210) ; they
are most firmly enclosed in

those ears which are most

FIG. 209. — Grams of Spelt , . , , , ,

wheat (T. Spelta), front, brittle and have the stoutest

back, and side views (nat. glumes. The ventral surface is

flattened or hollowed slightly,

the furrow of the grain shallow. The cross section is
less angular and the breadth proportionately greater
than that of a grain of T. dicoccum. The caryopses
measure 7-10 mm. long, 3-3*5 mm. broad, and 2-7-3
mm. thick ; the average ratio of length, breadth, and
thickness = 100 : 40-7 : 35'7-

The produce of the thrashed ears, i.e. the husked
grain or " Vesen," consists of 68 to 79 per cent of
caryopses, and 21-32 per cent of chaff (pieces of
rachis and glumes), different varieties varying con-
siderably in the relative proportions of grain and chaff.
One hundred caryopses weigh from 5 to 5-8 grams.
The weight of a hectolitre of " Vesen " varies from
40 to 48 kg.

FIG. 210.- — Grains
of the spikelets of
one side of an ear
of Spelt wheat
(T. Spelta) (nat.
size).

VARIETIES OF Triticum Spelta, L.
I. Ears bearded —

1. Glumes white, glabrous .... var. Arduini, Korn.

2. Glumes white, pubescent . . . var. albovelutinum, Korn.
Glumes red, glabrous ..... var. vulpinum, Korn.
Glumes red, pubescent .... var. rubrovelutinum, Korn.
Glumes grey-blue, glabrous .... var. Schenkii, Korn.

3-
4-
5-
6. Glumes blue-black, pubescent

var. coeruleum, Korn.

SPELT OR DINKEL 331

II. Ear beardless —

1. Glumes white, glabrous .... var. album, Korn.

2. Glumes white, pubescent .... var. recens, Korn.

3. Glumes red, glabrous . . . . var. Dukamelianum, Korn.

4. Glumes red, pubescent ..... var. neglectum, Korn.

5. Glumes grey-blue, glabrous .... var. amissum, Korn.

6. Glumes grey-blue, pubescent . . . var. Alefeldii, Korn.

Ear bearded ; glumes white, glabrous.

T. Spelta, var. Arduini, Korn. Handb. d. Getr. i. 80 (1885).
T. Arduini, Mazz. Sopra ale. sp. d. frum. 50, t. iv, Fig. i (1807).
T. Spelta, A., Metz. Eur. Cer. 26, t. vi, A (1824).
T. Spelta, L., Krause. Getr. Heft iv. 13, t. 5, D., E. (1836).
T. vulgare Arduini, Alef. Landw. Fl. 335 (1866).

A common variety, rarely pure, but often occurring among other varieties.
Winter and Spring forms are met with.

1. Bearded White Winter Spelt (i, Fig. 212). — A hardy winter form of
moderate yielding capacity, cultivated in Switzerland, South Germany, France,
Italy, and Spain.

Young shoots, prostrate.

Straw, tall, 115-130 cm. (about 44-50 inches) high, hollow, thin-walled but
firm ; leaves glaucous.

Ear, glaucous, lax, 12-15 cm. long, spikelets' 18-20, each often bearing 3
grains ; D = 15-17.

Empty glume, somewhat thin (2, Fig. 208).

Grain, red, flinty or semi-flinty ; 9-10 mm. long, 3-5-4 mm. broad, 3 mm.
thick.

2. Bearded White Spring Spelt. — A common spring form often grown
where the White Winter Spelt is cultivated ; its yields are small except on the
best soils. The plant tillers well and the straw is very strong and never
lodges. It is somewhat shorter than that of the winter form, and the ears
ripen 10-12 days later than those of the latter. The ears are very brittle,
narrow, and lax, 11-13 cm. long, spikelets 16; 0 = 13-14; grain 8 mm. long,
3-3 mm. broad, 2-9 mm. thick. Allied to this is a glaucous early form.

Ear bearded ; glumes white, pubescent.

T. Spelta, var. albovelutinum, Korn. Handb. d. Getr. i. 80 (1885).

Kornicke's type was derived from an ear of var. vulpinum, which he suggests
has been crossed with a velvet-chaffed variety of T. vulgare.

Under this variety Werner describes a White, Velvet-chaffed, Winter Spelt
with tall straw, 140-150 cm. (55-60 inches) high ; ears long, lax, narrow (15-20
cm. long) ; awns 7-8 cm. long ; grains red, mealy, 8 mm. long, 3 mm. broad.

Ear bearded ; glume red, glabrous.

T. Spelta, var. vulpinum, Korn. Handb. d. Getr. i. 80 (1885).
T. spelta, B., Metzger. Eur. Cer. 27, t. vi, A. (1824) ; Landw. Pfl. 94 (1841).

332 THE WHEAT PLANT

T. spelta, L., Krause. Getr. Heft iv. 13, t. 5, A., B., C. (1836).

T. vulgare vulpinum, Alef. Landw. Fl. 335 (1866).

Metzger states that this variety occurs frequently in fields of var. Duhameli-
anum, but Alefeld and Kornicke knew it only from Botanic Gardens.

Werner describes two forms of it : (i) Bearded Eed Winter Spelt, with
very tall, hollow, soft straw, 130-150 cm. (52-60 inches) high, liable to lodge and
rust. Ears lax and narrow, 12-17 cm. long ; spikelets 21 ; awns 7-8 cm. long ;
grain long and narrow, 8 mm. long, 3-5 mm. wide.

2. Bearded Dark Eed Winter Spelt with upright young leaves, straw 110-120
cm. high, narrow ears 12-16 cm. long, and glumes reddish-brown.

Ear bearded ; glumes red, pubescent.

T. Spelta, var. rubrovelutinum, Korn. Handb. d. Getr. i. 80 (1885).

Kornicke 's type originated from an ear of var. vulpinum. Werner describes
one spring and two winter forms of this variety.

The winter forms have bluish-red, velvet-chaffed ears, one with lax, the other
with somewhat denser ears.

The summer form (T.pretiosum, Jessen h. Dresden, 1872) possesses very long,
lax, pale red ears (14-18 cm. long), narrowed towards the apex, spikelets i cm.
broad, 3-awned, frequently containing 3 long flinty grains, 9 mm. long, 4 mm.
broad.

Ear bearded ; glumes grey -blue, glabrous.

T. Spelta, var. Schenkii, Korn. Handb. d. Getr. i. 80 (1885).
T. Spelta, L., Krause. Getr. Heft iv. n, t. 3 and 4 (1836).
Krause mentions this variety and a form of it with greyish-black glumes,
T. Spelta nigrescent, Schubl.

Kornicke states that he has not seen specimens.

Ear bearded ; glumes blue-black, pubescent.

T. Spelta, var. coeruleum, Korn. Handb. d. Getr. i. 80 (1885).
T. Spelta, C. and D., Metzger. Eur. Cer. 27, 28 (1824) ; Landw. Pfl. 95
(1841).

T. vulgare coeruleum, Alef. Landw. Fl. 335 (1866).

The colour of the glumes of this variety in hot seasons is blue-black on a
reddish ground ; in wet years the reddish tint predominates.

Bearded Blue Spelt (2, Fig. 211). — A very late form with prostrate or semi-
erect young shoots.

Straw, strong, 100-120 cm. (40-48 inches) high ; leaves broad, glaucous.

Ear, lax, 14-18 cm. long, narrowed towards the tip, rachis somewhat tough ;
spikelets 2-9 mm. broad, often with 3 grains in each ; 0 = 15-16.

Empty glume (3, Fig. 208), blue-black on a red ground, but in dull damp
seasons pale-ash grey with a dark line along the margins.

Grain, pale red, flinty, 8*5-9 mm- l°ng> 3'5~4 mm. broad, 3 mm. thick.

Werner mentions a very early spring form with taller straw and shorter ears
(11-16 cm. long).

FIG. an.— SPELT or DINKEL (T. Spelta, L.).

i. var. Arduini. 2. var. coeritleum.

SPELT OR DINKEL 333

Ear beardless ; glumes white, glabrous.

T. Spelta, var. album, Korn. Handb. d. Getr. i. 79 (1885).
T. Spelta, E., Metzger. Eur. Cer. 28, t. vi, B. (1824) ; Landw. Pfl. 95 (1841).
T. vulgar e album, Alef. Landw. Fl. 335 (1866).
T. Spelta, L., Krause. Getr. Heft iv. 15, t. 6, A. (1836).
One of the varieties most frequently grown and often found mixed with the
red-glumed var. Duhamelianum.

The following are common forms of this variety :

1. Beardless White Winter Spelt (i, Fig. 212). — A form widely cultivated
in South Germany and also met with in Switzerland and Southern France.
It requires a warm loam for the best yields.

Young shoots, prostrate with narrow leaves.

Straw, reddish, tall, 120-140 cm. (48-55 inches) high ; leaves glaucous.
Ear, narrowed towards the apex, pale yellow, very lax, 14-16 cm. long ;
spikelets narrow, 2-grained ; D = i6.
Empty glume (i, Fig. 208).
Grain, pale, brownish-red, mealy, narrow ; 7. mm. long, 3 mm. broad.

2. Beardless White Spring Spelt. — T. vulgare fringillarum of Alefeld has
short pale ears 11-12 cm. long ; spikelets 19-20 ; 0 = 15.

Grains, red, flinty, 8 mm. long, 3-2 mm. broad, 3 mm. thick. It ripens 10-12
days later than the winter form above.

3. Schlegel Dinkel (3, Fig. 212). — A less hardy form than (i), ripening
about a week earlier. It tillers well with straw about 100-1 10 cm. (40-44 inches)
high, and somewhat dense, pale yellow ears 10-14 cm. long ; . D =20-22. The
spikelets are broad, the grain reddish and mealy or semi-flinty, very long and
narrow (10 mm. long, 3-5 mm. broad).

4. Vogeles Dinkel. — A prolific winter form originally selected in 1836 by
A. Miinchenmaier, Hengenberg, in the Neckar Valley, Germany. The straw is
tall and somewhat weak, 120-130 cm. (48-52 inches) high, ears 12-17 cm. long,
of similar density to those of Schlegel Dinkel (D = 20-22) ; spikelets short, 19-20,
8 mm. broad ; grain angular and red ; 7 mm. long, 3 mm. broad.

Ear beardless ; glumes white, pubescent.

T. Spelta, var. recens, Korn. Handb. d. Getr. i.,8o (1885).
Kornicke's type was obtained from an ear of var. vulpinum.
Probably crossed with a velvet-chaffed variety of T. vulgare.

Ear beardless ; glumes red, glabrous.

T. Spelta, var. Duhamelianum, Korn. Handb. d. Getr. i. 74 (1885).
T. Duhamelianum, Mazz. Sopra ale. sp. d.frum. 55, t. iv, Fig. 2 (1807).
T. Spelta, F., Metzger. Eur. Cer. 29, t. vi, B. (1824) ; Landw. Pfl. 97 (1841).
T. Spelta, L., Krause. Getr. Heft iv. 15, Taf. vi, B., C. (1836).
T. vulgare rufum, Alef. Landw. Fl. 335 (1866).

The most frequently cultivated variety, grown extensively in Switzerland
and South Germany.

334 THE WHEAT PLANT

The following are the chief forms of it :

1. Tyrol Dinkel (2, Fig. 212). — One of the most prolific winter forms of
large Spelt.

Young shoots, prostrate or semi-erect ; young leaves narrow.

Straw, tall, stout, 120 cm. (about 48 inches) high.

Ear, very lax, 12-15 cm. long, rachis very brittle ; spikelets long and narrow,
7 mm. across with stiff glumes ; 0 = 17-18.

Empty glume (4, Fig. 208).

Grain, pale red, semi-flinty, usually not plump ; 10 mm. long, 3-5 mm.
broad.

2. Beardless Ked Winter Spelt. — One of the forms of Spelt widely cultivated
in South Germany and Switzerland. It is very hardy and yields well.

Straw, tall, strong, 125-130 cm. (about 50-53 inches) high.
Ear, lax, 15-17 cm. long; spikelets long, 9 mm. broad ; D = i6.
Grain, pale red, flinty ; 8 mm. long, 3-5 mm. broad.

3. Beardless Bed Spring Spelt is cultivated resembling (2), but with shorter
straw.

Ears, 11-12 cm. long ; spikelets 17 ; 0 = 15.
Grain, 8 mm. long, 3 mm. broad, 2-7 mm. thick.

Ear beardless ; glumes red, pubescent.

T. Spelta, var. neglectum, Korn. Handb. d. Getr. i. 80 (1885).
Kornicke's type was obtained from a Botanic Garden.

Ear beardless ; glumes grey-blue, glabrous.

T. Spelta, var. amissum, Korn. Handb. d. Getr. i. 79 (1885).
T. Spelta, L., Krause. Getr. Heft iv. 15, t. 6, D. (1836).
T. vulgare Duhamelianum, Alef. Landw. Fl. p. 335 (1866) (not Mazzucato).
A rare variety which Seringe states was much cultivated at the beginning of
the nineteenth century in the Canton of Bern, Switzerland.

Ear beardless ; glumes grey -blue, pubescent.

T. Spelta, var. Alefeldii, Korn. Handb. d. Getr. i. 80 (1885).
Kornicke received his type from Hohenheim.

According to Werner a form of it described below is cultivated in Italy.
Young leaves, narrow.

Straw, very tall, 135-145 cm. (about 53-57 inches) high, strong, hollow.
Ear, narrow, 10-15 cm. long ; spikelets 16-18 ; 7-8 mm. wide ; D = 2O.
Empty glume, pale reddish-blue, in damp seasons grey-blue with a dark line
along the margins.

Grain, long, narrow, flinty ; 8 mm. long, 3-5 mm. broad.

FIG. 212.— SPELT or DINKEL (T. Spelta, L.).

I. var. album.
(White Winter Spelt.)

2. var. Duhamelianum.
(Tyrol Dinkel.)

3. var. album.
(Schlegel Dinkel.)

CHAPTER XXIII

THE ORIGIN AND RELATIONSHIPS OF THE RACES OF WHEAT

ALTHOUGH the cultivation of wheat has not been traced to Paleolithic
times, stores of its ears or grain belonging to Neolithic man have been
discovered in many parts of Central and Eastern Europe, and the fact
that among these are specimens of several kinds suggests that its cultiva-
tion even at that period was of considerable antiquity. To the ancient
nations of Egypt, Greece, and Rome, the origin of wheat was as much a
mystery as it is to-day. The Greeks ascribed the gift of this precious
grain to Demeter, whom the Romans identified with their goddess Ceres.

Among the ancient Chinese, wheat was a grant from heaven, and the
institution, retained to recent times, of the annual sowing of the five
grains — wheat, rice, sorghum, millet, and soy — by the Emperors of
China was introduced by the semi-mythical Shen-nung or Chin-nong.

The wild prototypes from which the cultivated wheats have been
derived have been diligently sought, and records of their discovery exist
in all ages.

According to the legend of Diodorus Siculus (i. 4) the Egyptian
goddess Isis discovered " wheat (771730?) and barley growing promiscu-
ously about the country along with other plants, and unknown to man-
kind."

The country to which the reference alludes was Nysa, " a high
mountain of Phoenicia far away " (Diodorus i. 15), possibly the region
of Northern Palestine in which Wild Emmer (T. dicoccoides] is found at
the present day.

The Chaldean priest Berosus mentions the occurrence of wild wheat
(•nvpos) in the land of the Babylonians between the Tigris and Euphrates
(Syncellus, Frag. Hist. Grace, vol. ii. p. 416), and Strabo states that,
according to Aristobulus, " a wild corn similar to wheat (Trvpos) grows
in the land of Musicanus," the " most southern part of India " (Strabo,
Rerum Geog. lib. xv. t. i. p. 124; t. ii. pp. 776 and 1017, Amstel. 1707).

In more recent times Andre" Michaux in 1787 saw " Spelt " wheat
wild in Persia, north of Hamadan (Lamk. Ency. ii. 560), and Olivier
(Voy. dans Vemp. othoman, vol. vi. p. 338, 1807) says that when he was

335

336

THE WHEAT PLANT

north-west of Anah on the right bank of the Euphrates " we found near
the camp, in a sort of ravine, wheat, barley, and Spelt, which we had
already seen many times in Mesopotamia."

From the context these cereals were presumably uncultivated. Since
specimens have not been preserved, it is not possible to identify the wheat
and " Spelt " to which Olivier refers ; it is, however, highly probable
that the " Spelt " was the fragile-eared Wild Emmer ( T. dicoccoides) found
at the present day widely distributed throughout Asia Minor as far as
the western borders of Persia.

Beijerinck, Hausknecht, and others held the view that all the races
of cultivated wheat were derived from the single wild ancestor T. aegilo-
poides, and in the early half of last century Fabre concluded that Aegilops
ovata was the prototype (see p. 342). The majority of botanists,
however, who have studied the problem agree that the wheats are poly-
phyletic, and that at least two species are concerned in their origin. They
are also in accord regarding the main outlines of the relationship of the
chief races to each other and to the two wild species T. aegilopoides and
T. dicoccoides.

Schulz arranged the wheats in three separate series, the constitution
and relationship of which are indicated in the following table :

Series.

Wild
Prototype.

Groups of Cultivated Forms.

" Spelt "
Grain.

Naked Grain.

Normal Ears.

Monstrous Ears.

I. Small Spelt

T. aegilopoides

T. monococcum

None

None.

II. Emmer

T. dicoccoides

T. dicoccum

T. durum
T. turgidum

T.polonicum.
Not known.

III. Large Spelt
or Dinkel

Not known

T. Spelta

T. compactum
T. vulgare
T. compactum
x T. vulgare -
— T. capitatum

Not known.

Flaksberger's scheme of relationship given on p. 337 also divides the
wheats into three similar series or " conspecies," each with its corre-
sponding prototype.

Vavilov discovered that T. monococcum is quite immune to Puccinia
triticina, Eriks., and resistant to Erysiphe graminis, D.C. He also found
that T. durum, T. polonicum, and T. turgidum constitute a group resistant
to these fungi, while T. Spelta, T. vulgare, and T. compactum are all
susceptible.

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338 THE WHEAT PLANT

T. dicoccum has both immune and susceptible varieties. At Reading
the Indo- Abyssinian Emmers are slightly susceptible to rusts, the Russian
and Serbian forms more resistant, the late-ripening European repre-
sentatives being immune.

From a study of " serum " reactions Zade placed T. dicoccum , T.
durum, T. polonicum in one class, T. vulgar e^TTcompactum, and T. Spelta
in another, T. monococcum being distinct from these, and the results of
Sakamura's investigations into the chromosome number of the several
races suggests a similar grouping^

There is, I think, no doubt that the classification of the races of wheat
into (i) Small Spelt, (2) Emmer, and (3) Dinkel or Bread Wheat series,
supported as it is by so many independent lines of research, indicates in
a broad way their genetic affinities.

In regard to the phylogenetic relationship of the several races of
wheat, my investigations have led me to formulate the scheme given on

P- 339-

So far as concerns the races which are included in the three series

and the prototypes of two of them, it is in substantial agreement with
the conclusions of Schulz, Flaksberger, and others ; it differs, however,
in the view taken of the origin and relationship of the Bread Wheat
series.

The evidence upon which the scheme is based is discussed later.

SERIES I. — The Small Spelt series includes the wild T. aegilopoides,
Bal., and the cultivated T. monococcum, L.

Even from a cursory glance at the mature plants the relationship
between the wild T. aegilopoides and the cultivated T. monococcum is
evident.

In all morphological characters of the culm, ear, glumes, and grain
there is little or no difference between the two, the only modification
being a reduction in the hairs on the leaves and rachis.

Hybrids of T. monococcum with wheats of the dicoccum and vulgare
series are quite sterile, and the results of the determination of the chromo-
some number, immunity to rusts, and the " serum " reactions strikingly
support the conclusion that this series is distinct from the rest of the
cultivated wheats.

SERIES II. — The Emmer series comprises the Wild Emmer (T. her-
monis, Cook) and the cultivated Emmer, Khorasan, Macaroni,
Polish, Rivet, and Egyptian Cone wheats.

THE CULTIVATED EMMER WHEATS (T. dicoccum, Schiibl.) constitute
an ancient race agreeing closely in almost all its characters with the wild
T. dicoccoides. In both, the hairs on the surface of the leaf-blades are
similar in^form, length, and arrangement ; the ears also are flat, with

ORIGIN AND RELATIONSHIPS OF RACES OF WHEAT 339

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S?

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340 THE WHEAT PLANT

narrow spikelets containing two grains, and the empty glumes alike in
shape and texture. The wild plant has longer grains, and the conspicuous
fringe of silky hairs on the margins of the rachis is missing or much
reduced in the cultivated race.

The fragile-eared Indo-Abyssinian forms exhibit the closest re-
semblance to the prototype, possessing, like the latter, easily disarticulated
ears, short culms, yellow-green foliage, and the remarkable anatomical
character of four to six vascular bundles in the coleoptile, as well as the
early habit.

The European Emmers are a taller, later-ripening mutation, with
glaucous leaves, the usual pair of vascular bundles in the coleoptile, and
a fragile rachis.

KHORASAN WHEAT (T. orientale,mihi) is a small race having pubescent
leaves, short almost solid culms, very lax ears, strong awns, long flinty
grain, and early habit, all of which characters are found in T. dicoccoides,
from which I consider it has originated.

THE MACARONI WHEATS (T. durum, Desf.) exhibit obvious affinities
with T. dicoccoides and T. dicoccum, and I regard them as mutations derived
in some instances directly from the wild species, in others indirectly from
the latter by way of the cultivated Emmers. Like these the Macaroni
wheats possess solid culms, erect rigid ears with regularly arranged
spikelets, long, narrow, keeled glumes, and narrow, pointed grain ; many
forms also have a somewhat fragile rachis.

The Indian examples of the race, particularly the Kathias of the
United Provinces, with their comparatively small ears and slightly hairy
leaf-blades, appear to have arisen from the Indo-Abyssinian group of
Emmers, the European durums, with their large, coarse ears, glabrous
leaves, and smooth-based awns, being derived from the large-eared, tall-
strawed European Emmers.

POLISH WHEAT (T. polonicum, L.), the most recent of the races of wheat.
I have no doubt is a mutation of T. durum, as first suggested in 1884 by
Beijerinck. Like T. durum, its leaves are glabrous, its straw tall, striate
and almost solid, and the grain long and flinty. Both are also similar in
habit, tillering capacity, and resistance to rusts. The only point of
difference, namely the excessively long, thin glumes, I regard as a heredi-
tary teratological variation. I have seen occasional specimens of Indian
durums with elongated glumes suggestive of incipient polonicum, and
Kornicke states that Schweinfurth sent him from Upper Egypt a transition
form between durum and polonicum.

The lax ear, long spikelet, glumes, and grain of Khorasan wheat
(Race III.) suggest a variation towards the production of a polonicum
from the pubescent-leaved dicoccoides or Indo-Abyssinian dicoccum.

THE RIVET OR CONE WHEATS (T. turgidum, L.) have the characters
of a hybrid race produced probably within historic time by the crossing
of the tall European T. dicoccum with T. compactum or a dense-eared
form of T. vulgare.

Its affinity with the European Emmer can be traced in its morpho-
logical characters and habit. The two races agree in the characteristic
pubescence of their young leaves. Both have tall, solid or nearly solid
culms, and ears with spikelets very regularly arranged along the rachis.
They tiller very little and have a similar late-ripening period.

Moreover, the tendency to produce branched ears is strongly evident
in these two races and rare in others.

The square ear of T. turgidum , its many-flowered spikelets, and
plump, blunt-ended grain are characters derived from the dense-eared
compactum or vulgare parent, the dorsal hump of the grain being derived
from the Emmer parent.

In its typical form the Rivet race is endemic only in the area in which
T. dicoccum and T. compactum are present, namely, along the northern
side of the Mediterranean from Portugal eastward to the Caucasus, where
it is found sporadically distributed and hardly typical. It is absent from
Asia Minor, Persia, India, and China, in which countries either one or
both of the presumed parents are missing.

The solitary case from Baluchistan reported by Howard requires
further investigation. Forms bearing some resemblance to T. turgidum
I have received from the Central Provinces and Bombay ; those, of which
" Bansi " of Hoshangabad is a good example, are only endemic in the
provinces where T. dicoccum is grown and are probably hybrids of this
race with T. durum.

EGYPTIAN CONE WHEAT (T. pyramidale, mini) is a small but distinct
race confined to Egypt and Abyssinia. I regard it as an endemic dense-
eared mutation derived from the Abyssinian form of T. dicoccum. It
exactly agrees with the latter in the pubescence of its young leaves, yellow-
green culm-leaves, short culms, very early habit, and the shape of its
grain, and only differs from it in having short, dense ears with a tough
rachis.

SERIES III. — The Bread Wheat series includes Bread Wheat, Dinkel
or Large Spelt, Club and Indian Dwarf wheats.

THE BREAD WHEATS (T. vulgare, Host). — It is in regard to the origin
and relationship of the members of this series that the greatest diversity
of view is found among those who have studied the subject. The search
for a prototype possessing the characters of the vulgare race has been
diligently pursued from the earliest times, but nothing approximately
like it has been found growing wild. That another primitive species,

342 THE WHEAT PLANT

distinct from the prototypes of the Small Spelt and Emmer series, is
needed to account for the differentiating characters of the Bread Wheat
series is, however, generally recognised.

The discovery by Fabre of peculiar plants arising from the ears of the
wild grass Aegilops ovata, which became gradually transformed in suc-
cessive generations into wheat indistinguishable from that ordinarily
cultivated, led him to conclude that the wheats have been derived by
cultivation and selection from this wild species of grass. Fabre's view
was shared by many eminent botanists of last century.

In 1889 Kornicke announced that in the wild 7\ dicoccoides he had
found the prototype of his T. vulgare, which included all cultivated wheats
except T. monococcum.

Solms-Laubach decided that the ancestral wild wheat is extinct and
only the cultivated forms descended from it exist at the present day.

The belief that T. Spelta represents an ancestral type, and that the
rest of the vulgare series have been developed from it, has been somewhat
generally accepted, and in the phylogenetic schemes of Schulz and Flaks-
berger it is placed in the position of a primitive form whose wild proto-
type is yet undiscovered, although Schulz believes it probably exists in
the mountainous parts of the Euphrates-Tigris region.

From a study of the rachis and glumes of the two plants, Stapf states
that he is " almost convinced " that Aegilops cylindrica, a wild species
indigenous in Eastern Europe and Asia Minor, is the prototype of T.
Spelta.

While the characters of the wheat of the Small Spelt and Emmer series
can be traced with reasonable certainty to their respective prototypes,
from which they appear to have been derived chiefly by mutation, selection,
and cultivation, investigation of the morphological features of practically
all known forms of T. vulgare has convinced me that there is not, nor has
there ever been, a prototype of the Bread Wheat series. The characters
of T. vulgare and its allies appear to me to be those of a vast hybrid race,
initiated long ago by the crossing of wheats of the Emmer series with
species of Aegilops, and that T. Spelta is a segregate of this hybrid.

Crossing between mutants of the same specific prototype is, I think,
sufficient to account for many, if not all, of the moderate number of forms
found among the races of the Emmer series, but the extraordinary com-
plexity and almost endless number of varieties and intermediate forms
of the vulgare race can only be satisfactorily explained by the assumption
of its hybrid origin from two or more distinct species.

There is scarcely a single character, morphological or physiological,
which is not subject to very wide variation, and in respect of any pair of
its contrasting characters, such as lax and dense, bearded and beardless
ears, different forms of empty glume, early and late ripening period,

ORIGIN AND RELATIONSHIPS OF .RACES OF WHEAT 343

prostrate and erect habit of its young shoots, flinty and mealy grains,
immunity and susceptibility to attacks of fungi and other features, there
exists an almost continuous series of transition forms between their
extreme limits.

Any interpretation of the origin of the vulgare race and with it the
rest of the Bread Wheat series must account for the racial characters
mentioned below, which differentiate T. vulgare from the Emmer series.

(i.) The presence of a single line of long hairs on the summit of the
longitudinal ridges of the young leaf-blades, with shorter
ones or none at all on the sides of the ridges,
(ii.) Thin-walled, hollow culms.

(iii.) The exceptionally tough, non-disarticulating rachis.
(iv.) The rounded back and absence of keel on the lower part of the

empty glume of a large proportion of the race.

(v.) The comparatively short awns of the fully bearded ears, and the
occurrence of beardless and semi-bearded ears.

The presumed prototypes of the Emmers possess none of these ;
they are, however, all found in Aegilops ovata, L., or A. cylindrica, Host,
and I have no doubt that both these species have entered into the con-
stitution of the vulgare race of wheats.

The species named are wild annual grasses, the former widely dis-
tributed in the Mediterranean region from Portugal to Egypt, Syria and
Transcaucasia, the latter appearing in Italy, the Balkans, South Russia,
and Asia Minor. As already indicated, the long hairs on their leaf-blades
are like those of T. vulgare in form and arrangement but longer. The
culms are hollow and thin- walled. The rachis is non-fragile, but the
ear falls off the straw as a whole without disarticulating into separate
spikelets. The grains are small, resembling in form and general mealiness
those of T. vulgare, and like the latter are convex on the dorsal side without
the prominent ridge found on grains of the Emmer series.

In A. ovata the culms are 20-35 cm. high and the ears 2-3 cm. long,
with 2-5 spikelets, one or two at the base and apex being rudimentary
and sterile. The fertile spikelets are 3- to 4-flowered, each ripening one
or two grains.

The empty glumes are convex, inflated, and truncate at the apex,
without a keel, but having several prominent lateral nerves, and terminating
generally in three or four awns, 2-4 cm. long (i, Fig. 223). The flowering
glumes terminate in 2 or 3 short awns, usually less than 2 cm. long.

A. cylindrica is a taller species with more erect culms 30-60 cm. high,
and longer, more slender, cylindrical ears 6-12 cm. long, and composed
of 5-10 spikelets (Fig. 223). The axis breaks below the spikelets as in
T. Spelta when roughly handled (Figs. 214, 217). The empty glume

344

THE WHEAT PLANT

of the lateral spikelet is oblong, broad and truncate at the apex, with a
short terminal awn of variable length, and a much shorter lateral tooth,
that of the terminal spikelet bearing one or two very long awns. The
flowering glumes are awnless or only shortly bearded.

il

FIG. 214. — Empty glumes and spikelets of
(i) Aegilops cylindrica and (2) T. Spelta,
illustrating mode of fracture of the rachis

It is conceivable that the heterogeneous race
of T. vulgare wheats with its numerous Aegilops
characters may have arisen from the hybridisa-
tion of these two species, or from one, or
both, by mutation, selection, and cultivation,
but I think this is much less possible than
the view I have just expressed. The greater
length of ear, frequently keeled empty glume,
larger grain, and occasional solid culm appear to me to need the intro-
duction of another species like that of T, dicoccoides and its derivatives.

Both species of Aegilops mentioned have been artificially hybridised
with Emmers or with T. vulgare. Cook also refers to a Syrian specimen.

FIG. 213. — Ear of Aegilops
cylindrica, with two empty
glumes (nat. size).

ORIGIN AND RELATIONSHIPS OF RACES OF WHEAT 345

intermediate in its ear characters between A. ovata and T. dicoccoides,
which Aaronsohn considered a natural hybrid, and the photographs of the
spikelets undoubtedly support this conclusion.

It is highly probable that such natural hybrids are not infrequent,
and although the Fl generation produced and grown under European
experimental conditions is generally sterile, the vulgar e hybrid is fertile
with vulgare pollen. It is also possible that the natural hybrids between
the wild Aegilops and the Emmer prototype — the suggested parents of
the vulgare race — are more fertile under their native climatic conditions
than in Central or Western Europe.

The particular contribution made by the two species of Aegilops to
the vulgare race can at present only be surmised. Either of them wrould
account for the keelless empty glume, but the beardless character of half
of these wheats has most likely come from the short-awned beardless
A. cylindrica.

Many forms of T. vulgare are glaucous, others are non-glaucous and a
green or a yellowish-green tint ; the former agree with A. ovata, the
latter with its sub-species A. triaristata, which differs from the type
ovata chiefly in its colour, more robust growth, and the possession of
three instead of four awns on the empty glume.

Empty glumes with a single long terminal awn instead of the usual
short apical tooth are not infrequent among vulgare and compactum
wheats (Figs. 165, 191), especially those from Central Asia. This
character I regard as evidence of the presence of Aegilops in these races,
for it appears in hybrids of A. ovata with T. vulgare, though neither of
the parents show it. The three or four long awns on the empty glume
of A. ovata are reduced to one in the second or third generation of the
cross (p. 382 and Fig. 223).

DINKEL OR SPELT (T. Spelta, L.). — That T. Spelta is in some way
closely related to the Bread wheats (T. vulgare) is usually admitted. The
characteristic arrangement and form of hairs on the young leaves, the
hollow culms and broad but slightly keeled empty glumes which they
have in common are evidences of their affinity.

Speltoid mutations and transitional speltoid forms are not uncommonly
seen, more especially among the vulgare wheats coming from Central
Asia, Argentina, and Spain.

T. Spelta is frequently grouped with the fragile-eared Emmers, but
this classification is based on imperfect knowledge of the nature of its
rachis. In T. dicoccoides the ear falls in pieces as it ripens (cf. Fig. 77), the
rachis disarticulating at the nodes, and a similar though less easy dis-
articulation is found in the brittle-eared forms of T. dicoccum. The
rachis of T. Spelta is also brittle, and when roughly treated breaks into
short lengths, but . the transverse fracture occurs below the point of

346 THE WHEAT PLANT

insertion of the spikelets and not at the exact nodal point ; indeed, dis-
articulation at the node is not always easy to effect, even when force is
used, and on this account T. Spelta should be included among the tough-
eared wheats.

Exactly the same mode of fracture and at the same point of the rachis
is found in Aegilops cylindrica, and Stapf's conviction that these two are
closely related is, I think, undoubtedly correct. My view, however,
differs somewhat from his in that I regard A. cylindrica not as the proto-
type of T. Spelta but as one of its parents.

The experiments of Fabre, Godron, and others with Aegilops x T. vulgare
hybrids have shown that the complete or even approximate segregation
of an Aegilops never occurs in any succeeding generation of their descend-
ants, a result almost unique and hitherto unexplained.

T. Spelta, however, which very frequently appears among the progeny
of hybrids of vulgare wheats with other races, I regard as a segregate
nearest to the cylindrica parent.

CLUB WHEAT (T. compactum, Host). — In the chief morphological
characters of its young leaves, culms, glumes, and naked grain this race
resembles T. vulgare, and there is little doubt that both have had a
common origin. The great density of the ears of the race is a character-
istic hereditary feature, and such constant, dense, dwarf-eared forms
frequently arise apparently de novo among the progeny of hybrids between
widely different wheats with longer lax ears. I think it highly probable
that the compactum race arose in this way and made its appearance con-
temporaneously with the vulgare wheats.

It is one of the most ancient of races, archaeological evidence showing
that it was widely distributed in prehistoric times, and apparently more
commonly cultivated at first than the larger-grained vulgare wheats.

INDIAN DWARF WHEATS (T. sphaerococcum, mihi). — Like its ally
T. compactum, this race probably arose as a mutant among the progeny
of an early hybrid. Its peculiar small round grains very closely agree
in form and size with Buschan's T. compactum globiforme, the naked-
grained wheat most commonly grown in various parts of Europe in
Neolithic times, and it is possible that the race now confined to India is a
remnant of an ancient stock.

CHAPTER XXIV

VARIATION

ONE of the most obvious peculiarities of living things is their variability.
No two human beings are alike in all their features, and no two plants
of the many thousands present in a field of wheat are exactly alike in all
their characters, even where the entire crop has descended from a single
ancestral grain.

The form, size, and colour of the glumes, the height, strength, and
diameter of the straw, the length, weight, and number of spikelets of the
ear, and all other characters of the plant, morphological and physiological,
are subject to variation.

Although variations may be classified and investigated from many
points of view, they fall naturally into two distinct groups, namely :

(1) Fluctuations, or racial variations, and

(2) Discontinuous variations or sports.

FLUCTUATIONS

Fluctuations pass by regular gradation from a lower to an upper
limit, and are distributed at random among the individuals forming a
crop. When measured, weighed, or counted, the results can be plotted
on frequency curves, which conform to the laws of chance.

Variations of this class are produced by»changes in the environment
of the plant ; they belong to the individual and are not inherited.
. Examples of fluctuating variability are the lengths and numbers of the
spikelets of the ears of a crop of wheat.

Of 500 ears taken at random from a field of " Swan " wheat, the
length varied between 50 and no mm. Measured and arranged in
groups differing from each other by 5 mm. the frequencies or numbers
of individuals in each class were —

LENGTHS OF THE EARS (mm.)

Class . . {

5°
54

55
59

60
64

65
69

70
74

75
79

80
84

85
89

90
94

95
99

100
104

105
109

Frequency

i

9

28

30

93

94

98

77

49

10

9

2

Percentage in
each class .

•2

1-8

5'6

6-0

18-6

18-8

19-6

i5-4

9-8

2-0

1-8

•4

347

348

THE WHEAT PLANT

Similarly, the number of spikelets per ear varied between 14 and 24,
and the number of internodes per straw between 4 and 7, with the
following frequency distributions :

NUMBER OF SPIKELETS PER EAR

Class

H

15

16

I?

18

J9

20

21

22

23

24

Frequency

i

O

6

18

35

103

141

I4I

48

6

I

Percentage in each

class

•2

o

1-2

3-6

7-0

20-6

28-2

28-2

Q-6

1-2

•2

NUMBER OF INTERNODES PER STRAW

Class ....

4

5

6

7

Frequency

4

103

366

28

Percentage in each class

•8

20-6

73-2

5-6

In dealing with quantitative variations the application of statistical
methods is of great service in obtaining accurate expressions for the
amount and range of the variability of each character, and the determina-
tion of the mean (M), the standard deviation (<r), and the coefficient of
variability (C), or ratio of the standard deviation to the mean of all
fluctuating characters of plants, the pedigree of which is known, would
be of much value.

Some of these, obtained <from investigations of " Swan " wheat, are
given below :

c

M.

<r.

per cent.

Length of ear . . . mm.

77-72 ± -184

9'95 ± -13

12-8

Length of straw . . . cm.

99-4 t -30

14-06 ± -212

14-1

Length of upper internode cm.

39-97 — -142

6-69 ± -io

16-7

Length of lowest internode cm.

2-737± -029

I -21 ~I -02

4f2o

No. of internodes per straw

5-836

•513

8-78

No. of spikelets per ear

21-78 ± -029

1-59 ± -021

7-3

No. of grains per ear .

33-64 ±3-29

7'55 i2-27

22-4

Weight of ears per plant . gr.

16-81 ~ -49

9-03 ± "34

53-7

No. of ears per plant

5-46 ± -145

2-7 ™- -103

49-0

The following results obtained by Oetken refer to a Squarehead
wheat :

VARIATION

349

M.

<r.

c,

per cent.

Ear length, 2 good ears . . cm.

7-96 ± -04

•56 --03

7-i- '3

Ear length from longest straw cm.

7-90- -04

•6Sn:-03

8-2± -4

Ear length from shortest straw cm.

7'i7± 'OS

•8o = -O4

n-i± '5

Straw length (the longest straw) cm.

126-1 - -38

5-56±-27

4'4± '2

Straw length (the shortest straw cm.

108-0 = -60

8-88 ±-42

8-2± -4

Straw length (the average straw) cm.

118-2 ± -37

5 -04 ±-26

4'3 ± '2

Difference between longest and

shortest . . . cm.

18-4 ± -66

8-95 ±-46

48-6±2-s

Tillering

5-8 ± -19

2'79±'i3

47-8±2-3

Total yield per plant gr.

33'5 ±1-16

i6-53±-8o

49'3 ±2-4

Grain yield per plant gr.

1 1-2 = -43

5'30±-28

47'4±2-s

Grain yield to total yield %

32-7 ± -24

3'3i±'i7

io-i± -5

Weight of ear of longest straw gr.

2-89 ± -04

•58rb-o3

20- 1 ± i-o

Weight of ear of shortest straw gr.

i'95± -°4

•6i±-03

3i'5±i'5

Average yield of grain per ear gr.

i-8g± -03

•38 ±-02

20'0±I-0

Grains per spikelet .

2-37± -02

•29±'OI

I2'4± '7

looo-grain weight for whole plant gr.

40-43 ± -23

3-56--I9

8-8± -5

looo-grain wreight, 2 good ears gr.

4i-57= '35

4-63 ±-25

u-i± -6

Although all characters of the wheat plant fluctuate between a higher
and lower limit, there are great differences in the extent and range of their
variability. Of the characters mentioned above, the number of spikelets
per ear, length of straw and ear, number of internodes per straw, and
looo-grain weight vary much less than the yield per plant, weight and
yield of grain per ear, and the " tillering " or number of straws produced
by each plant.

As already stated, individual variations or fluctuations are not in-
herited, and are considerably influenced by soil and season ; nevertheless,
the mean and range of variation of the different characters are hereditary
features of the group or population constituting the pedigree cultures
raised from a single plant. For example, the grains from an ear 60 mm.
long do not produce plants with ears of this length only, but a fluctuating
series, having the same mean and standard deviation as those of the
pedigree culture from which the original grain was derived.

CORRELATIONS.— The different organs of a plant are so co-ordinated
with each other that variation in one part involves a simultaneous variation
in another. In wheat, for example, variation in the number of straws
or tillering is associated with an alteration in the total weight of the
plant, and length of growing period with the total yield of grain per
plant.

Such correlated variation may be positive ( + ), an increase in one of the
characters carrying with it an increase in the other, or it may be negative
(-), in which case an increase of one is linked with a decrease in the
other.

While in many instances the facts of correlation are undoubted, the

35°

THE WHEAT PLANT

exact nature and causes of the relationships are not understood in all cases,
and satisfactory explanations of them cannot always be given.

Where the variations are capable of being quantitatively expressed,
the amount of the correlation between a pair of characters can be repre-
sented as a coefficient calculated from a correlation table, in which is
displayed the frequency distributions of the simultaneous variations of
the two characters concerned.

The coefficient of correlation (r) lies between o and i . It is a measure
of the extent to which one character varies in unison with another.

When r=i complete correlation is indicated, the variations being
controlled by exactly the same causes ; this degree of correlation is,
however, rarely, if ever, found between any of the characters of living
organisms.

When r = > -9 correlation is almost complete.
,, r= -6-'9 there is very high correlation.
,, r = '3-'6 there is distinct correlation.
r =o there is no correlation.

CORRELATION BETWEEN THE NUMBER OF SPIKELETS AND THE LENGTH
OF THE EAR IN " SWAN " WHEAT

Length of the Ear (mm.).
Average Lengths of Classes.

50

55

60

65

70

75

80

85

90

95

100

i°5

f

54

59

64

69

74

79

84

89

94

99

104

109

I

I

z^

O

16

2

2

2

6

CL>

17

I

i

-}

18

-1 /

18

3

IO

IO

IO

2

35

o,

CC

19

6

17

34

32

II

3

103

e

20

i

2

37

40

37

*7

7

141

6

21

6

19

43

49

19

3

2

141

22

i

2

5

6

20

5

7

2

48

2-j

2

2

2

6

24

*

I

i

A

I

9

28

30

93

94

98

77

49

IO

9

2

500

VARIATION

351

Below is given a list of some correlations which have been observed in
wheat. The extent of the correlation varies more or less with the race
and variety of wheat and the external conditions of growth :

Subject Character.

Co-ordinated or Relative
Character.

Positive or Negative
Correlation.

Length of straw

Number of straws

Diameter of straw .

7»
»

Length of the ear .

»
Length of growing period

>»
>»
»
»
Weight of the grain

»»

Brittleness of rachis
Long upper internode .

Length of ear

Weight of ear

Weight of grain in ear

No. of grains per ear

Grain yield per plant

Grain yield per ear

Density of ear

Total weight of plant

Length of straw

Weight of straw

Length of ear

Density of ear

Weight of grain per plant

No. of grains per ear

Resistance to frost

Late ripening

Length of upper internode

Length of lower internode

Tillering power

Number of spikelets

Density of the ear

Total yield of plant

Total weight of grain per plant

Average size of the grain

Starch content of grain

Relative nitrogen content

Size of the grain

Nitrogen content

Close-fitting glumes

Heavy ear

352

THE WHEAT PLANT

The following are examples of the size of the correlation coefficient
of a number of characters of wheat, based chiefly on data obtained from a
few pedigree cultures :

Subject Character.

Relative Character.

r.

Length of straw .

Length of ear

•439 ±-017

Percival

»

»

•292

Roberts

»

No. of grain per ear

•668

»

»

Average yield of grain per

•44 ±-06

Oetken

ear

»

Yield of grains per plant

•08 ±-07

»

»>

»

•294 ±-032

Love

»

Average weight of grains

•278 ±-033

) )

No. of straws per plant

Weight of plant

•94 ±-oi

Oetken

>»

Weight of grains per ear

•834

Love

>»

»

•04 ±-07

Oetken

»

Length of straw

•281

Roberts

»>

No. of grains per ear

•180

»

»

Average weight of grain -013 ±-032

Myers

per plant

Length of ear

No. of spikelets

•757 ±-013 i Percival

»

,,

•735 ± -008

»

>j

Average length of rachis

•909

Parker

internodes

No. of grains per plant

Average weight of grains

•251 ±-033

Love

»

Yield of grain

•985 ±-ooi

J5

No. of spikelets per ear

No. of grains per ear

•230 ±-09

Percival

»

>j

•515 ±-07

»

Weight of straw and ear

Weight of grain per straw

•925 ± -004

Myers

»

»

•890 ± -006

»

Weight of ear

Weight of grain per ear

•777 ± -008

Percival

Average weight of

grains sown

Length of ear

-•5ii±-oi5

Waldron

»

Length of straw

-•404 ±-017

»

Yield of grain

Average weight of grains

•327 ±-03 1

Love

SPORTS OR DISCONTINUOUS VARIATIONS

It is a common experience of all who have made a study of pedigree
wheats raised from a single grain or ear of a plant that, sooner or later,
there appear among the progeny varieties which do not belong to the
fluctuations of the particular form, but differ from these in the possession
of one or more new characters. They are only occasionally found in the
small plots constituting the first or second generations of a selected ear,
but turn up with a considerable degree of regularity so soon as from 20
to 100 square yards of the pedigree culture is obtained.. Such sports or
discontinuous variations arise suddenly, and are not connected by inter-

VARIATION 353

mediates with the parent form. In some cases the new characters are
not constant, while in others they are hereditary sports of the latter class,
being usually designated mutations or mutants. There is scarcely a
single character normally present in wheat which may not thus arise
spontaneously among forms of wheat in which it has not previously
been observed, and the new feature may be either a Mendelian dominant
or a recessive.

Among beardless wheats bearded forms appear, and among bearded
forms beardless plants arise.

In regard to the colour of the glumes, red-chaffed varieties arise
among white-chaffed wheats, and white-chaffed plants among red.

Varieties with black awns sport to white- or red-awned forms, and
vice versa.

Among lax-eared wheats plants suddenly appear with compact ears,
and in dense Squarehead wheats varieties with long lax ears are found.

Glabrous glumed forms give rise sometimes to forms with pubescent
glumes, and vice versa.

Investigations to determine the nature and origin of the sports
enumerated have proved that many of them are natural hybrids, and their
progeny exhibits simple Mendelian segregation. Breeding tests of
others show that they are segregates — heterozygotes, homozygous domin-
ants and recessives — of natural hybrids which have escaped observation
in previous seasons. The majority of such examples generally involve
only one or two allelomorphic pairs of characters. A few sports, however,
which I have observed in pedigree cultures of T. vulgare, T. compactum,
and T. Spelta were much more complicated and gave rise to an extra-
ordinarily heterogeneous mixture of descendants. In Fig. 215 is illus-
trated the remarkable variety of forms obtained from a sport of a Chinese
beardless smooth-chaffed bread wheat, received from Chungking under
the name " Kwang T'ou Mai." The grains from a single ear were sown
in 1911, and from the progeny, which was apparently uniform, a single ear
clubbed at the apex and lax at the base, as in ear i, was selected and its
grains sown in 1912. These yielded 57 plants in 1913, one of them a
compactum form (ear 3) like the upper half of the original clubbed ear,
but with pubescent glumes, the rest smooth-chaffed and of uniform
density (ear 2).

From the compactum sport 36 plants were raised, among which 12
distinct forms were found (line A). Some of these had long lax ears,
others possessed intermediate or short and compact ears. Clubbed as
well as uniformly dense-eared plants were obtained, the majority beard-
less, but a few with short awns at the tips of the ears. In regard to the
pubescence of the glumes, some were densely hairy, others only slightly
pubescent, while several were quite glabrous.

2 A

354 THE WHEAT PLANT

The grains from single ears of each of the 36 " sported " plants were
sown separately, the range of the forms of the ears of their progeny being
illustrated in line B. In addition to the morphological characters already
mentioned, plants with fully-bearded ears appeared in this generation.
Examples such as this, if they are cases of Mendelian segregation, involve
several pairs of allelomorphic characters.

Some varieties of wheat which have remained constant for a con-
siderable period suddenly give rise to one or more sports.

These cases I regard as hybrids, which are stable under one set of
climatic and soil conditions, but have been induced to sport by cultivation
in a new environment. Examples which I consider support this
view, I observed in two pedigree cultures of Huguenot wheat a peculiar
beardless form of T. durum (Fig. 146), selected from a crop of Medea
(T. durum) about 1898 by J. Currell of Arthur River, Western Australia,
and supposed to be a hybrid between Medea and Purple Straw (a
vulgar e wheat).

For four years two short rows of 10-12 plants of each culture were
raised at Reading annually from single ears of the previous season. These
showed no variation until the fifth year, when both cultures sported in the
same manner, giving fully-bearded, semi-bearded, and beardless plants
of T. durum ; the numbers of each, however, were too small to admit of
any reliable ratios being established.

It may be argued that such an example, like those already mentioned,
is merely a case of segregation of a natural hybrid, the crossing of which
took place in the season immediately preceding that in which the sporting
occurred. This explanation, however, is not satisfactory, for it is very
improbable that two separate cultures should have been accidentally
crossed in the same season with apparently the same second parent.

From a bearded red-chaffed Squarehead wheat which had been
constant for several seasons, Riimker observed the production of a very
variable beardless sport under circumstances which excluded mixing of
sorts or natural hybridisation as explanations of the appearance of the
new form.

The original bearded form was a selection from the progeny of the
hybrid Landwheat $ x Squarehead $ .

I have had similar experience with Badger wheat (a selected bearded
Squarehead).

Other examples are known among wheats, and Blaringhem states
that the hybrid Svanhals barley, constant when grown at Svalof, reveals
its hybrid origin when grown in Flanders and Picardy.

These and similar sports are usually described as mutations, and
their new characters are assumed to be due to germinal variation arising
independently of segregation and recombination of the hereditary units

H
P

O

w
«
o

Q
—

eu

id

<£-2

T3 2

II
8 I

.2*c
ti o

Q C

Z NO.

o

OH

CO

o a

en co

s s

.00

— —

VARIATION 355

of the plants. Without at present being able to offer any explanation of
the processes involved, I am inclined to look upon them as the product
of hybrids, the ordinary segregation of which has been delayed or in-
hibited for a time.

The various races of wheat differ considerably in their tendency to
sport or mutate ; those in which evidence points to their being of hybrid
ancestry are much more subject to this kind of variation than the purer
races. For example, sports are much more frequent in T. vulgare and
T. compactum, two races which I regard as hybrid (pp. 342, 346), than in
T. durum or T. dicoccum. I have never seen an example in T. monococcum,
the race which gives no fertile hybrids. Certain forms, such as the Square-
head vulgare wheats, are also especially liable to vary in this manner.
Changes in the environment appear to stimulate the production of sports.
In some seasons, especially those with an abnormally high average summer
temperature following severe winters, sports are more frequent than usual
in autumn-sown wheat. Rimpau records great variation among Square-
head wheats throughout Germany in 1903, especially among the bearded
forms which had before been very constant and pure.

Buffum obtained sports in Black Emmer and a Turkey Red type of
vulgare wheat immediately after sowing in rich " sagebrush " soil at an
altitude of 4000 feet in Wyoming.

SPELTOID MUTATIONS. — These sports observed among the descendants
of pedigree cultures and hybrids of T. vulgare have the narrow open ears
and truncate empty glumes of T. Spelta ; the culms are tall and the ears
erect, rigid, and harsh.

Most of them are heterozygotes, which segregate into homozygote
normal forms and heterozygote speltoids in the ratio i : i , which Nilsson-
Ehle explains by the assumption of the total elimination of male speltoid
gametes.

In a few cases segregation occurs into normal forms, heterozygous
speltoids and homozygous speltoids in the ratio i : 2 : i, but more often
the proportion of speltoid forms is smaller than this owing to the partial
failure of male gametes.

I have found considerable numbers of speltoid forms (2, Fig. 173)
among mixed samples of vulgare wheats from Persia, Central Asia, Spain,
and Argentina ; these, however, are usually homozygous.

DWARFS. — Dwarf plants with short culms and compact short ears
are found in the F2 and subsequent generations of hybrids of certain
races of wheats. Some of them are fertile, with normal glumes and
grain ; others are monstrous forms with deformed sterile ears.

I have never observed the production of dwarf mutants among pedigree
lines of wheats, nor have I met with clear references to any. Farrer,

356 THE WHEAT PLANT

however, observed the occurrence in Australia of what he termed " grass-
clump plants " among the progeny of many of his hybrid wheats. Richard-
son also reports their appearance among his crosses in the same country,
and Cutler obtained a single specimen, which he describes as a " dwarf
wheat," in a plot of Marquis wheat in Saskatchewan. These forms differ
from the ordinary crop in producing dense, compact clumps of leaves,
from which arise only a few short culms with small ears. Farrer states
that they appear among crosses between widely different types of T.
vulgar e only and are first seen in the F2 generation, though in one instance
(a cross between " Bokhara desert " wheat and a Fife-Indian variety) all
the Fj plants (four in number) were " grass-clumps."

There is, I think, little doubt that these are segregates of the ordinary
" winter " type, requiring a longer period for completion of their growth
than the Australian climatic conditions allow, and are not true " dwarf "
plants ; they' behave under these conditions exactly as " winter " forms
of Bread Wheat do in this country when sown too late in spring or summer
to produce ears in the same year (see p. 90), and if protected from drought
would doubtless produce normal culms and ears in the following season.

BUD SPORTS. — This class of variation is exceptionally rare among
wheats.

Akerman describes four examples discovered among the descendants
of hybrids of T. vulgare. Larger or smaller portions of the ears of these
sports have empty glumes which resemble those of T. Spelta in form,
nervation, and close investment of the grain.

In one of them found in the F2 of the hybrid Iron x Thule II. wheats,
the ear had speltiform empty glumes on one side and normal vulgare
glumes on the other.

A similar sporting ear was obtained on a plant with five ears, four of
which were normal, the plant being one of the F5 progeny of the hybrid
Brown Schlanstedt x Borsum (a Norwegian spring wheat). From the
same cross another plant arose having three ears, two normal and one
with the upper part speltoid, and the lower portion speltoid on one side
and normal on the other.

In the fourth example all the spikelets of the upper part of the ear
possessed speltiform empty glumes, but in the four lowest spikelets one
of the empty glumes of each spikelet was speltiform and the other normal
vulgare.

The speltoid empty glumes in all these cases are erect and nearly
parallel to the rachis, the corresponding opposite glumes being arranged
almost at right angles to them. The affected spikelets are turned to one
side in a manner similar to some of those in Fig. 179.

Akerman regards these as periclinal or sectorial chimaeras. Three of
the sports, it was suggested, were normal plants surrounded by a hetero-

VARIATION 357

zygote speltoid epidermis, since they gave only normal progeny. From
the fourth example some of the grains produced speltoid heterozygote
plants which segregated in the ratio of i : i. In this type one or more
subepidermal cell-layers from which the gametic cells arise were evi-
dently heterozygote.

I have never observed white-chaffed and red-chaffed ears or white-
grained ears and red-grained ears upon the same plant, but plants are not
infrequently seen bearing fully-bearded and almost beardless ears upon
the same individual, the fully-bearded ears being always found on the
first- formed culms.

Plants possessing both " clubbed " and lax ears of uniform density
are not infrequently met with at Reading, the lax ears being borne on the
youngest straws. Rimpau observed both long lax ears and short denser
Squareheads upon an Fx plant of the hybrid

Frankenstein (lax-eared vulgar e) x Igel (compaction), and Caron-
Eldingen also describes and figures a plant possessing both lax and
" clubbed " ears of two different types which he explains as resulting
from a twin grain (p. 359).

Possibly some of these variations may be manifestations of natural
reduction of vigour or inadequate nutrition of the later-formed " tillers "
of the plants and hardly to be classed as examples of bud sports.

Ohlmer, Preul, and others attribute the clubbing of the ears of Square-
head wheats to an excess of nitrogen and diminished water-supply in the
early stages of development of the culm and ear, and my own observations
support this view.

TERATOLOGICAL SPORTS. — Besides the sports which are concerned
with the discontinuous variation of the normal characters of the wheat
plant, there are teratological sports, " monstrosities " or " freaks " pos-
sessing abnormal features, such as twin ears, supernumerary spikelets,
and other examples mentioned below.

They are comparatively rare and their cause unknown.

i. BENT AND SINUOUS UPPER INTERNODES. — In some individual culms
the upper internodes are bent completely round so that the ear is parallel
with the straw and its apex downwards. In others the upper internode
is sinuous just beneath the ear. Both these malformations are more
frequent in T. turgidum and T. durum than in other races, and are often
seen in the deformed progeny of hybrids between distinct races (Fig.
220).

ii. MALFORMED UPPER INTERNODE AND LEAF. — A common deformity is
the arrested development of the upper internode of the culm, the first
node being less than an inch from the base of the ear. In such cases the
ear is usually normal, but the uppermost leaf is more or less malformed,

358 THE WHEAT PLANT

its sheath and reduced blade sometimes appearing like a spathe of the
inflorescence.

This deformity is met with in all races of wheat and is caused by a
parasitic fungus.

iii. VARIEGATED LEAVES. — Plants with white longitudinal stripes along
the leaves are occasionally found in T. dicoccoides and T. vulgare. In the
examples I have seen, the abnormality is hereditary and chiefly confined
to the young leaves, those of the culm being uniformly green. The
edges of the variegated leaves are usually pink.

iv. BRANCHED EARS. — Many forms of T. turgidum have branched ears,
abnormal secondary axes of variable length arising from notches of the
rachis in the place of ordinary spikelets. The character is also seen in
T. dicoccum, from which I consider that T. turgidum has arisen. It not
infrequently appears among hybrids of T. dicoccum, and is seen occasion-
ally in T. polonicum, but is uncommon in other races. The development
of secondary ears is hereditary in certain forms, but the degree of branch-
ing is very largely dependent upon climatic and soil conditions. The
morphology of these ears is discussed in the description of Miracle or
Mummy wheat (p. 256).

v. TWIN EARS. — Two ears are occasionally found at the apex of a single
straw. Shirreff records such a variation, both the twin ears being perfect
in form, one containing forty-five grains, the other thirty-six. In all the
cases I have seen, it is the rachis which bifurcates, as in Fig. 216, and not
the culm. An example with the rachis divided near the base is figured in
the Gardeners' Chronicle and Agricultural Gazette for 1849 (p. 363).

The production of twin ears I have only seen in T. vulgare. It is not
hereditary and is much less common, and quite a different phenomenon
from the branching of the ears so frequently seen in forms of T. turgidum.

vi. SHORT EARS. — Short stumpy ears consisting of three to seven
spikelets are found occasionally in T. vulgare and T. turgidum.

The spikelets are of normal size and form and develop well-filled
grain. The terminal spikelet, unlike the arrangement in normal ears, is
parallel to the rest. Possibly the true apex of the rachis is destroyed in
its early development by a parasite, although there is no evidence of such
damage in the mature ears.

The variation is not hereditary.

vii. SMALL MALFORMED EARS, generally on short sinuous culms, appear
not infrequently among the progeny of hybrids between distinct races of
wheat, more especially when T. dicoccoides or T. dicoccum is one of the
parents (see Fig. 220). A number figured by Buffum were obtained
among the Fx of a cross between a " mutating " T. dicoccum and a " sport "
of Turkey Red wheat (T. vulgare).

OS

W £

H g

UJ ro

as

OS C
O '?

« 5

N U3

a
a ^

-- --a.,'"-

VARIATION

359

viii. SUPERNUMERARY SPIKELETS. — These sometimes arise singly by the
side of a normal spikelet and arranged at right angles to it. Both spring
from the same notch of the rachis.

Additional spikelets are also frequently found growing from the rachis
immediately below the points of insertion of the normal spikelets and
generally arranged parallel to them (i, Fig. 216). In some cases they
produce one or two grains, but are often rudimentary, consisting only of
minute misshapen glumes.

These variations, which I have found most often
in Chinese vulgar e wheats, occur chiefly in ears of
the latest tillers, the ears which develop first being
usually quite normal ; the variation does not appear
every season, though a form which has once shown it
may produce it again.

ix. AWNS. — I have not observed many variations
in the structure or general morphology of the awns
of wheat.

In a Persian form of T. vulgar e the short awns of
the flowering glumes of some of the spikelets some-
times have a thin membranous outgrowth on each
side, about half-way between the tip and base of the
awn (Figs 179, 217). The peculiarity is not constant.

In several Central Asiatic forms of beardless T. vulgar e the short awns
at the apex of the ears are often strongly curved and sometimes bent into
the form of a hook.

x. TWIN GRAINS. — Malformations of the parts of the flower are rare
in wheat. The only case recorded appears to be that figured by Caron-
Eldingen, of a twin grain with two embryos, developed from two united
ovaries.

FIG. 217. — Flowering
glumes of a Semi-
retchensk form ( T.
vulgare) (see 2,
Fig. 179)-

CHAPTER XXV

HYBRIDISATION AND WHEAT HYBRIDS

WHILE self-fertilisation may be considered the rule, examples of natural
hybrids among wheats are much more frequent than is generally assumed.

Howard states that cross-fertilisation is common among wheats in the
hot dry climate of Northern India.

Kornicke observed numerous hybrids among his cultures in Germany,
and Godron, Vilmorin, and others note the occurrence of cross-fertilisa-
tion in France.

Nilsson-Ehle and Kajanus state that in Sweden a small amount of
crossing takes place at Svalof and elsewhere, and renders it difficult to
preserve the purity of select varieties when these are multiplied and grown
on a large scale.

Shirreff records the crossing of some of his wheats, and natural hybrids
are found in small numbers annually at Reading both in experimental
pedigree cultures and in the open fields.

Many of the " sports " discovered among wheats in all parts of the
world have proved to be natural hybrids or their segregates.

A few wheats are distinctly protogynous, but in the majority the repro-
ductive organs ripen at the same time.

There are considerable differences among them in regard to their
tendency to hybridise. While many forms are probably always self-
fertilised, others cross more or less readily when grown in proximity to
another kind, and in such instances it is found that crossing sometimes
takes place in one way only, the reciprocal hybrid being never obtained.
This is most obviously the case with the wild T. dicoccoides, but Nilsson-
Ehle records the natural hybridisation of Swedish Velvet Chaff ( $ ) and
Pudel wheats (?) when grown in mixtures with another form of T.
vulgar e (Old Brown " Landwheat " c?)> the reverse crosses being never
found.

Since natural hybridisation occurs among widely different races and
varieties when grown on small plots, it may be assumed that it takes
place also in the field between plants of the same crop, and it is possible

360

HYBRIDISATION AND WHEAT HYBRIDS 361

that the greater vigour and yield of some wheats may be due to the fact
that they cross more frequently than the less prolific types.

The production of artificial hybrids of wheats appears to have been
first attempted in this country by Thomas Andrew Knight about the end
of the eighteenth century. After successfully crossing peas he tried the
hybridisation of wheat, but states in his Essay on the Fecundation of
Vegetables (1804) that it " did not succeed to my expectations," and goes
on to say : " I readily obtained as many varieties as I wished by merely
sowing the different kinds together : for the structure of the blossom of
this plant (unlike that of the pea) freely admits the ingress of adventitious
farina, and is thence very liable to sport in varieties."

In 1846 Mr. Maund of Bromsgrove exhibited hybrid wheats at a
meeting of the English Agricultural Society (Gardener's Chronicle, 1846,
p. 601), and in the same year Hugh Raynbird produced the hybrid Piper's
Thickset $ x Hopetoun <$ , the female parent a red-chaffed wheat with
short dense ears, the pollen parent a lax-eared form with white chaff.
The few hybrid grains obtained were sown in September and the " roots "
divided in January. ' The produce," he states, " was many kinds both
of red and white wheat ; some of the ears bore a perfect resemblance to
the Piper's Thickset, others partook of the character of the Hopetoun in
everything except in the colour of the chaff, others had half the ear thin
and open, and the rest close set, thus in the same ear showing the char-
acteristics of each kind."

One of Raynbird's hybrids was awarded a gold medal in 1848 by the
Highland and Agricultural Society of Scotland, and several produced by
Maund and Raynbird were exhibited at the Great Exhibition of London
in 1851.

These were the first hybrid wheats grown on a large scale.

From about 1850 to 1870, Patrick Shirreff of Haddington in Scotland
paid especial attention to the improvement of cereals and produced
several hybrid wheats.

In the early 'seventies of the nineteenth century Pringle and Blount
in the United States, Vilmorin in France, and Rimpau and Heine in
Germany were all actively engaged in hybridising wheats. Since that
date the crossing of this cereal has been extensively practised in a great
many countries.

Little is known of the conditions upon which success or failure depends
in the artificial hybridisation of wheats. Some crosses are difficult, while
others are exceptionally easy. As a rule, wheats of different races do not
cross so readily as varieties and forms of the same race, but it is occasion-
ally observed that pollinations carried out at the same time upon different
ears of the same plant are not all equally productive of grain.

Very slight differences in the state of development of the styles and

362 THE WHEAT PLANT

age of the pollen, the temperature and moisture of the atmosphere at the
time of pollination, and the time of day at which the operation is per-
formed, all have an influence upon the result.

While pollinations may be effective at all times of the day, I have had
the best returns from those made in the morning before 10 A.M. At that
time the tendency to rapid drying of the temporarily exposed parts of the
flower is much less than during the hotter part of the day.

The hybrid grains are frequently smaller and more shrivelled than
self-fertilised grains of the same ear, more especially when the two parents
belong to different races, and small imperfectly formed grain sometimes
results after cross-fertilisation between two nearly related forms, although
plump grains are equally frequent.

The embryos of shrivelled hybrid grains are usually delicate, and
special care is required in rearing them. It is best to sow the grain in
pots under glass rather than in the open soil, where the risk of destruction
of the young plants by frosts, rain, and pests is greater.

The shrivelled endosperm of hybrid grains is probably connected
with the fact that, like the embryo, it is of hybrid origin, or possibly its
feeble development is attributable to the failure of the second male gamete
to unite with the fusion-nucleus of the embryo sac.

The method of the artificial hybridisation of wheat is simple, never-
theless there are certain important points of detail to which it is useful to
direct attention.

Since anthesis commences usually in two to five days after the escape
of the ear from the upper leaf-sheath, crossing must be undertaken during
that period or the chances of finding flowers in which the anthers have
not already shed their pollen on the adjacent stigmas will be small.

After selecting the grain-bearing female parent, the following is the
plan of operations which I adopt :

1 . Cut off the upper third of the ear with fine pointed scissors and pull off
completely four or five of the lowest spikelets.

2. Of the remaining spikelets remove completely every alternate one on both
sides of the rachis.

3. Take hold of the tips of the few spikelets left and pull outwards and down-
wards ; this removes the upper florets of the spikelets, and leaves for crossing
only the two lowest and largest florets of each.

4. Gently press the glumes of the florets to be emasculated, and insert the
point of a pair of closed forceps between the upper edges of the glume and palea.
On releasing the forceps the glumes are separated and the anthers are exposed
to view.

5. If the anthers are green and erect they can often all be removed together,
but if further advanced in their development they should be removed singly,
care being taken not to burst them in the process.

HYBRIDISATION AND WHEAT HYBRIDS 363

In some wheats, two of the anthers are^hidden behind the overlapping edges
of the palea and require great care in their removal. Somewhat blunt-pointed
forceps are less liable to break the anthers than those with very fine points.

6. Select an ear of the pollen parent, and with scissors cut off the upper half
of the glumes of a few of its median spikelets. With forceps remove any thus
exposed anthers which are bright yellow, and after pinching them dust the
escaping pollen on the styles of the emasculated flower ; leave the broken anther
in contact with the styles and allow the glumes to close.

7. Enclose the crossed ear in an envelope of thin waterproof paper, and with
cotton tie the lower open end of the envelope round the upper part of the straw
so as to prevent the access of foreign pollen or insects .

Where waterproof envelopes are not available, take a strip of thin soft paper
six inches wide and wind it three or four times round the crossed ear and upper
part of the straw ; tie above and below the ear to prevent foreign pollination.

8. Label the crossed ear indicating the respective parents and date of the
cross.

9. After a week examine the crossed ear, and if the stigmas are withered
remove the covering altogether.

(If the stigmas are receptive, fresh pollen applied to them germinates im-
mediately and fertilisation is effected in forty-eight hours or less.)

Raynbird, Shirreff, Pringle, and Vilmorin, among the earlier hybridists,
observed the extraordinary multiplicity of forms among the descendants
of hybrid wheats, but it is only since the re-discovery of Mendel's laws
of inheritance that any explanation has been given of the cause of the
diversity.

In recent years there has been an accumulation of more or less pre-
cise knowledge of the mode of inheritance of the chief characters of the
species and races of wheat, which is summarised in the following pages.
In making clear some of the points which were previously obscure, the
Mendelian concepts of heritable factors and gametic purity have been
invaluable, but much remains to be discovered before all the facts of
inheritance in the highly complex hybridised wheat plant can be satis-
factorily explained.

INHERITANCE OF THE CHIEF CHARACTERS OF THE WHEAT PLANT

ERECT AND SPREADING HABIT (p. 69, Figs. 65, 66). — Hybrids of
wheats with these characters are intermediate, the F2 segregating in the
1:2:1 ratio.

EARLY AND LATE RIPENING PERIOD. — These characters are correlated
with the foregoing allelomorphic pair, early wheats always having upright
young shoots, while in the latest forms the young shoots are prostrate.

Hybrids are intermediate with semi-erect young shoots (B, Fig. 66),
and segregate in F2 in the 1:2:1 ratio.

364 THE WHEAT PLANT

HOLLOW AND SOLID STRAW. — In the majority of wheats of the vulgar e
and compactum races, the culms are comparatively wide and hollow with
thin walls ; those of T. durum, T. polonicum, and T. turgidum are smaller
in diameter and full of pith, or thick- walled with a very small central
cavity.

The hollow form is dominant over the solid straw, and F2 segregation
is in the 3 : i ratio.

PUBESCENT AND GLABROUS YOUNG LEAVES. — The surfaces of the
young leaves of T. durum are nearly or quite glabrous, those of other
races being more or less densely clothed with hairs.

In general the pubescent leaves of T. dicoccum and T. turgidum are
dominant over the less hairy and glabrous leaves, but the segregation
of the characters in the F2 generation has not been adequately in-
vestigated.

NORMAL AND BRANCHED EARS.- — According to Tschermak, the normal
condition is dominant over the branched ear of " Mummy " wheat
(T. turgidum, var. mirabile) (Fig. 160).

FRAGILITY AND TOUGHNESS OF THE RACHIS. — In the two wild wheats
T. aegilopoides and T. dicoccoides the rachis of the ripe ear separates
transversely at the nodes, each spikelet with an internode of the rachis
attached falling to the ground when the ear is slightly shaken.

In three cultivated wheats, T. monococcum, T. dicoccum, and T. Spelta,
the rachis is also fragile, but in a lesser degree.

In contradistinction to these are the rest of the cultivated wheats in
which the rachis is tough and does not become disarticulated even under
the rough treatment of thrashing.

i. Brittle-eared x Brittle-eared.

ii. Brittle-eared x Tough-eared. — The crossing of two brittle - eared
wheats gives a constant brittle-eared progeny, and fragility of rachis
behaves as a dominant character when a brittle-eared wheat is crossed
with a tough-eared form.

iii. Tough-eared x Tough-eared. — Hybrids of varieties belonging to the
same tough-eared race usually have constant tough-eared descendants, but
where the two plants crossed belong to different tough-eared races, brittle-
eared forms sometimes appear in the F2 and subsequent generations.

H. Vilmorin observed brittle-eared forms resembling T. Spelta among
the F2 generation of the hybrids, T. vulgare (Ble Seigle) x T. turgidum
(Poulard Ble' Buisson), and T. vulgare ? (Chidham d'automne) x T.
durum $ (Ismael).

Love and Craig record the appearance of two plants with fragile ears
similar to those of wild T. dicoccoides out of 113 plants of the F2 genera-
tion of the cross T. vulgare $ (Early Red Chief) x T. durum <$ (Marouani).

HYBRIDISATION AND WHEAT HYBRIDS 365

The numbers suggest the 63 : i ratio of a Mendelian three-factor
hybrid.

LAX AND DENSE EARS. — Extensive investigations have been made
regarding the inheritance of the lax open ears and the denser-eared
" Squarehead " forms of T. vulgar e as well as the short dense ears of T.
compactum.

The researches of Nilsson-Ehle have shown that the lax character of
T. vulgare is often dependent upon two (or possibly more) independent
factors (Lj and L2) over which the single factor (C) for short ear or T.
compactum is dominant.

i. Lax x Lax. — Hybrids of two lax-eared forms in which both factors
for laxity are present give rise to lax-eared descendants only. Where
the lax-eared wheats have the constitution L^ and 1XL2 respectively, in
the F2 there arise plants (Lx L2) with more open ears than either of the
grandparents and Squareheads (lx 12) in which both factors are absent.

ii. Lax vulgare x T. compactum. — In most cases the compactum char-
acter is dominant and the F2 segregates approximately in the ratio 3
dense-eared : i lax-eared.

In some instances, however, the dominance is incomplete, and group-
ing into two classes is not possible owing to the existence of an almost
continuous series of intermediates of varying ear-density between the lax
and compact ears of the grandparents.

For further discussion of these characters see pp. 401-405.

COLOUR OF THE AWNS. — The heredity of awn colour has not been
closely examined. It probably agrees with that of glume colour, black
and red being dominant to white and black dominant to red.

The development of the dark pigment in the awn like that of the
glumes is greatly influenced by climatic conditions, a fact which makes
the determination of its inheritance specially complicated.

PRESENCE AND ABSENCE OF AWNS. — The possession of awns or beards
is a primitive character and is common to all races of wheats. In addition
to fully-bearded plants, so-called " beardless " wheats are abundant in
the four races T. vulgare, T. compactum, T. sphaerococcum, and T. Spelta ;
in these the awns are absent or reduced to i cm. or less in length.

The following are the chief results of investigations concerning the
inheritance of the fully-bearded and beardless characters :

i. Bearded x Bearded. — The crossing of two bearded forms gives a
bearded ¥l generation, and the character is continued in all subsequent
generations so long as the wheats belong to the same race. Where the
plants belong to different races the Fx generation is also usually bearded,
but in some cases beardless as well as bearded forms appear in the F2
and later generations : an example of the latter is recorded by Vilmorin,

366 THE WHEAT PLANT

who obtained a beardless vulgare form among the descendants of the
hybrid T. polonicum x T. turgidum, var. lusitanicum (Pe'tianelle blanche),
both of which are bearded.

ii. Bearded x Beardless. — In the cross between the bearded T. dicoccum
and the beardless T. sphaerococcum (p. 392) the bearded character was
completely dominant in F1} and plants as beardless as the T. sphaerococcum
parent were not found in the F2 or F3 generations. In other crosses,
especially where the wheats belong to the same race, the beardless char-
acter is dominant, the bearded recessive.

As explained elsewhere (p. 104), the term " beardless " includes the
rare wheats in which awns are entirely absent from the ears, and the very
common group with awns from -3 cm. to about i cm. in length on the
spikelets of the upper fourth of the ear.

In the majority of records of crosses between bearded and beardless
wheats the latter belonged to the short- tipped group. The Fl generation
of such hybridisation has frequently been described as beardless, the
beardless character being dominant, and subsequent segregation in F2
was stated to occur in the ratio of 3 beardless dominant : i bearded
recessive.

This view of the inheritance of awns requires modification, since the
complete dominance of beardlessness is rarely seen. C. E. Saunders
(Report of the Third International Conference on Genetics , 1907, p. 370)
referred to the production of semi-bearded plants in the Fl generation
of crosses of bearded and beardless forms ; out of some 300 of these
hybrids he found that :

15 per cent were more than one-quarter bearded.
59 per cent were one-quarter bearded.
20 per cent were nearly beardless.
6 per cent were beardless.

Fully-bearded Fx plants were not observed, and he says " the number
of plants which could fairly be called beardless was very small."

In F2 he obtained fully-bearded and beardless plants with an unclassi-
fiable series of intermediates.

No doubt the particular length of awns of the heterozygote F1 genera-
tion depends on the difference of the parent forms employed in the cross,
but the fact that beardlessness is but imperfectly dominant is now quite
clear. The early inaccurate conclusion of complete dominance of beard-
lessness and the 3 : i segregation in the F2 generation was due to the
confusion of what may be termed " semi-bearded " heterozygotes with
the so-called " beardless " homozygous plants possessing short awns at
the tips of the ears.

When the latter " beardless " forms of T, vulgare are crossed with

F.

L

FIG. 218. — HYBRID (Fx) OF BEARDED x BEARDLESS WHEAT,
with F.2 and F3 segregates.

HYBRIDISATION AND WHEAT HYBRIDS 367

typically bearded forms of the same race having awns 6-9 cm. long, the
Fx plants have ears which on careful examination are readily distinguished
from those of the beardless parent, the awns of the spikelets of the upper
quarter of the ear being generally from 1-5 to 3 cm. long ; the influence
of the bearded parent is also visible in the points on the flowering glumes
of the lower portion of the ear, for these, though short, are distinctly
longer than those of the corresponding glumes of the beardless parent used
in the cross.

It is true that in the majority of cases the " semi-bearded " heterozy-
gotes are not exactly intermediate, but have awns much nearer in length
to those of the " beardless " than to the fully-bearded homozygous
parent ; they are nevertheless distinct from " beardless " plants and can
be readily separated from them in nearly all cases by eye inspection alone.

The F2 generation from these " semi-bearded " ¥1 hybrids consist of
"beardless," semi-bearded, and bearded plants in the ratio 1:2:1.
This has been established by numerous workers, and I have repeatedly
investigated natural hybrids between bearded and " beardless " forms of
T. vulgar e with the same result.

A. and G. L. C. Howard have made observations on several crosses
between bearded and quite beardless wheats belonging to T. vulgare.

The F! generation of these crosses had ears with very short tips to the
apical spikelets.

In the F2 generation were plants with completely beardless ears and a
series of forms with awns of various lengths, the ratio of the more or less
bearded to the quite beardless examples being 15 : i.

The character and distribution of the bearding among 986 plants of
an F2 generation is indicated below :

Fully bearded.

Almost
fully bearded.

Half
bearded.

Tips of
varying Length.

Quite
beardless.

61
Ratio . i

65

I

I24
: 2

673
: 10-7

63
I

14-7

I

The dihybrid ratio observed suggests that two factors are concerned in
the production of the fully-bearded ear, one factor being responsible for
short or " tip " beards, the other when combined with this giving rise to
long awns.

The fully-bearded plants, according to the Howards' view, contain
two factors, B and T, the former representing the " long," the latter the
" short " factor.

368 THE WHEAT PLANT

The gametic constitution of fully-bearded plants would be represented
by BBTT, the short-tipped plant by bbTT, and the strictly beardless by
bbtt.

Crossing a fully-bearded (BBTT) with a beardless plant (bbtt) would
give an Fl of the constitution BbTt, which in F2 would produce :

i BBTT (fully bearded) +4 BbTt (very short tips) +2 BbTT (nearly
bearded) +2 BBTt (half bearded) + 1 bbTT (short tips) +2 bbTt (minute
tips) + 1 BBtt (long tips) + 2 Bbtt (very short tips) + 1 bbtt (quite beard-
less).

The fully-bearded, quite-beardless, " long-tipped," and " short-
tipped " plants of the F2 generation just mentioned bred true in F3 ;
later the two latter were proved to contain the respective constituent
factors which on combination gave rise to the fully-bearded character.

These results, although probably of limited application, are of much
interest and help to explain the existence of semi-bearded " long-tipped "
forms (BBtt), which breed true, as well as the production of fully-bearded
wheats from the crossing of forms which would ordinarily be classed as
" beardless " (see below).

A repetition and extension of the work of crossing bearded and the
various " beardless " forms with accurate measurement of the awn
lengths of the descendants and mathematical analysis of the results is much
to be desired.

Tschermak states that in the two hybrids Fiirst Hatzfeldt (beard-
less) x Galician (bearded) and Mold's white (beardless) x Hungarian
(bearded) the bearded character was lost and did not appear among the
descendants.

iii. Beardless x Beardless. — In almost all cases the crossing of two
" beardless " forms results in " beardless " offspring both in the Fx and
all subsequent generations.

Rimpau, however, observed a small number of constant bearded plants
among the progeny of the second generation of a cross between two
beardless wheats, Saxon " Land " wheat and Squarehead, and Spillman
obtained bearded forms from the cross Little Club x Farquhar, both
beardless parents.

The Howards also record the production of fully-bearded plants in
F2 from the crossing of " long-tipped " and " short-tipped " " beardless "
wheats. Assuming the gametic constitution of the parents to be BBtt
and bbTT respectively, the Fj would be represented by BbTt, and, as
explained above, some of the descendants of this hybrid would be fully
bearded in the F2 generation.

It is, however, improbable that this explanation covers all cases of the
appearance of fully-bearded plants among the progeny of crosses of
" beardless " wheats.

HYBRIDISATION AND WHEAT HYBRIDS 369

COLOUR OF THE GLUMES. — The glumes of wheat are generally described
as some shade of black, red, or white, the red embracing various shades of
brown or red, and the white various shades of yellowish- white. Black
and red are more or less dominant over white, and black also dominates
the red tints.

i. Black* Red. — According to Tschermak, white-chaffed individuals
sometimes appear among the descendants of this cross, and Rimpau
obtained white-glumed forms in the F2 generation of a cross between a
bluish-black chaffed T. turgidum and a red-chaffed T. vulgare (p. 400).

ii. Black x White. — Tschermak states that the F2 of this cross is of
complicated character, but the ratio of the pigmented to the white forms
is 3 : i.

From a cross of Black Emmer (T. dicoccum. var. atratum) xa white-
chaffed T. sphaerococcum I obtained in F2 forms with the following chaff
colours : uniform black, red, and white, and chaff of these tints with black
outer margins. A similar series was obtained by Kezer and Boyack in
the F2 of the hybrid Black Emmer xa white-chaffed T. vulgare (p. 393).

iii. Red x Red. — The descendants of this cross are generally some shade
of red, though, as indicated later, white individuals may arise in F2 from
the hybridisation of two red forms.

iv. Red x White. — Very numerous hybrids have been raised between
red-chaffed wheats and white-chaffed wheats : in the majority of these
cases the Fx is intermediate in colour, and the segregation in the F2 genera-
tion of the ordinary monohybrid type 3 red : i white, or i red : 2 pale
red : i white, the colour being dependent upon a single pair of Mendelian
factors. In such wheats each of the many shades of red appears to have
its own special factor.

In crosses made between white wheats and some red-chaffed Swedish
" Landwheats " Nilsson-Ehle found that the segregation in F2 was that
of a dihybrid cross, viz. 15 red : i white.

He showed that the red colour in these cases is not due to a simple
factor but to two different and separately inherited red factors ; each of
these by itself produces a red tint, not necessarily of the same intensity,
and when acting together they have an additive or cumulative effect, the
colour increasing with the number of doses of the colour factors present
in the zygote.

If the two red factors are represented by Rx and R2 and their absences
by r, and r2 respectively, a red-chaffed wheat (RiR2) when crossed with a
white-chaffed variety (r1r2) results in a heterozygote Fj of the constitution

R1R2 rir2-

The sixteen possible zygotes of the F2 generation with their respective
doses of red factors are shown in the following diagram :

2E

37°

THE WHEAT PLANT

Gametes. R^. RI^. riR2- rxr?.

RXR2

RiR2

Rlf2

rlR2 rlr2

(4)

(3)

(3)

(2)

Rlr2

(3)

(2)

(2)

(2)

R!^

Rlr2

rxR2

RiR2

Rtr2

riR2

rlr2

(3)

(2)

(2)

(0

rxR2

r!R2

V*

(2)

(i)

(I)

(o)

From only one (r^ r^) out of the sixteen zygotes are all the red
factors absent : this will be white, the rest, having from one to four colour-
factors, will be of several shades of red.

When a wheat with deep red chaff, due either to one or two factors, is
crossed with a white-chaffed variety, the glume colour of the heterozy-
gotes in F2 is always an easily recognised red, and homozygous reds even
of the lighter shades are found to be distinct from white.

Where the coloured parent of the cross is of a light reddish tint the
heterozygotes are very pale and sometimes difficult to distinguish from
the homozygous whites.

Some of the F2 individuals classed as white were found by Nilsson-
Ehle to be in reality heterozygotes which split in F3 into red, pale red,
and white-chaffed plants. Owing to the confusion between these hetero-
zygote white and the pure white, the experimental results of segregation
may appear not to conform to the ordinary 3 : i or 15 : i ratios.

In Nilsson-Ehle's crosses of red and white spring wheats the total
numbers obtained in F2 were 2625 red : 987 white —2"j red : i white, and
in some of the individual crosses the apparent splitting was 2-3 or 1-5 red
to i white, the number of plants classed as white-chaffed being too high.

In consequence of the cumulative effect of two red factors the cross
Rjra x R2rx would, of course, lead to the production of forms containing
both Rj and R2, and therefore darker in colour than either parent,
together with some white individuals (r^ rnr2) lacking the colour factors
altogether.

v. White x White. — The Fx of this cross has invariably white chaff,
but the occurrence of red-chaffed plants is recorded by Shirreff in the F4
generation of the hybrid ShirrefFs Bearded white ? x Talavera $ , both
white- chaffed wheats.

HYBRIDISATION AND WHEAT HYBRIDS 371

PUBESCENT AND GLABROUS GLUMES. — Pubescence and smoothness of
glumes usually constitute an allelomorphic pair of characters, pubescence
being dominant, and segregation of the F2 is in the ratio 3 pubescent : T
smooth-glumed form.

In some instances, however, the hybrid (F5) between a velvet-chaffed
and smooth- chaffed wheat is more or less intermediate, the degree of
pubescence varying considerably ; in these hybrids segregation in F2
follows the 1:2:1 ratio.

In the hybrid between a pubescent and beardless T. vulgare (Velvet
Kolben) and a pubescent and bearded form of T. compactum (Igel) Riimker
obtained both glabrous and velvet-chaffed plants in the F2 generation in
.the ratio i glabrous : 3 velvet (p. 403), and a similar ratio of forms with
glabrous and pubescent glumes appeared in the F2 of my hybrid between
two pubescent-chaffed wheats, T. dicoccum 9x7". sphaerococcum $

(P- 392).

Pubescence was found to be linked with black chaff in hybrids of
Black Emmer with T. vulgare (p. 394) and T. sphaerococcum (p. 393).

CLOSE-FITTING AND LOOSE GLUMES. — In T. dicoccum and T. Spelta
the glumes firmly invest the grains, while in T, vulgare, T. turgidum, T.
durum, and some other wheats the glumes loosely enclose the grain.

The close-fitting glumes of T. Spelta are dominant over the loose
glumes of other wheats except in hybrids with T. turgidum, in which,
according to Kajanus, the close-fitting character is recessive.

LONG AND SHORT GLUMES. — For inheritance of the long glume of
T. polonicum see pp. 394-399.

KEELED AND ROUNDED GLUMES. — The keeled character of the glumes
of T. durum, T. turgidum, and T. Spelta is dominant over the rounded
glume character of T. vulgare and T. compactum.

BROAD AND NARROW GLUMES. — The broad truncated glumes of T.
Spelta are dominant over the narrower-pointed glumes of other wheats.

FORM AND SIZE OF GRAIN. — Several of the different races of wheat
possess grains of characteristic shape and size which can be recognised
without much trouble, although they are difficult to describe.

The grains of hybrids between the several races usually show the
characters of both parents, but little accurate research into the features of
hybrid grains has been made except in regard to the grain-length of crosses
of T. polonicum with T. durum which was investigated by Engledow.
The mean length of the grain of the T. polonicum used was about 10-2
mm., that of the durum parent 7-7 mm.

The Fj grain was intermediate. In respect of size of grain three types
were found in F2 with the following mean grain-lengths : 8-84, 8-67, and

372 THE WHEAT PLANT

8-33 mm., representing the extracted polonicums, heterozygote inter-
mediates, and extracted durums respectively.

The frequencies of the three types agreed with those found for the
glume-lengths of the F2 of the same cross (p. 396), which suggests a i : 2 : i
ratio.

No grains were found as long as those of the grand-parental polonicum
and none as short as those of the durum ancestor, an inward " shift "
towards the mean of the two grandparents having occurred similar to that
described for glume-length (p. 396).

The length of the grains of the extracted homozygote polonicum
of F2 and F3 was 13-14 per cent less than that of the original polonicum
parent, the F3 durum forms being about 8 per cent greater than those of
the original durum.

GRAIN COLOUR. — In respect of colour, wheat grains are usually divided
into the two classes red and white. In the former a reddish or brownish
substance is present in the testa of the seed and absent from the seed-coat
of white grains.

One or two Abyssinian varieties (pp. 195, 196) of wheat have purple-
coloured grains, but in these the colour is due to an anthocyan pigment
in the " chlorophyll " or " cross layer " of the pericarp.

i. White x White. — The usual result of crossing two white-grained
wheats is the production of white-grained descendants in Fx and all
subsequent generations.

Vilmorin, however, obtained red-grained plants among tlje F2 progeny
of a hybrid between two white-grained forms of T. polonicum and T.
turgidum (p. 397).

Pitsch also records the appearance of a few yellow- or reddish-grained
plants in the sixth generation from the hybrid " Squarehead " x" Chal-
lenge," which he states were two white-grained wheats.

ii. Red x Red. — The progeny from this cross is generally red in all
generations.

Tschermak, however, states that in some instances white-grained
individuals have appeared among the descendants of hybrids between
different red-grained varieties.

Nilsson-Ehle also records the occurrence of white-grained plants in
F2 of the hybrid " Bore " wheat x " Extra Squarehead," both red-grained
parents.

He suggests that two independent and different factors for red (Rj
and R2) are concerned, the respective uniting gametes being RjT2 and ^Rg.

From such a cross, white individuals (rjr2) would appear in F2, in the
proportion of i white to 15 red.

iii. Red x White. — Very numerous investigations have been made
regarding the hybridisation of red- and white-grained wheats.

HYBRIDISATION AND WHEAT HYBRIDS 373

Red dominates white, the Fx generation being red-grained, but the
grains are a paler tint than those of the red parent.

The segregation in F2 depends on the character of the red parent.
In a large number of instances the F2 generation consists of red- and
white-grained descendants in the proportion of 3 red : i white, the red
tint being due to a single Mendelian factor.

Nilsson-Ehle discovered that the trihybrid ratio of 63 red : i white
occurred in the F2 generation when certain red wheats were crossed with
white.

From the hybrid Svalof " Pudel " 9 (a white velvet-chaffed, white-
grained Squarehead) x Swedish " Sammet " <$ (a velvet-chaffed form
with dark red grain) he obtained in Fx plants with pale red grains. All
the grains of the F2 generation were also red, but of several shades. On
raising the F3 generation from one of these hybrids, he found that some of
the plants segregated approximately in the ratio 3 red : i white ; others
gave a progeny of about 15 red : i white, while a smaller number exhibited
the trihybrid segregation of 63 red : i white.

An analysis of the results led him to the conclusion that the red grain
colour of the particular parent employed was dependent upon the presence
of three dominant and independent factors Rx, R2, and R3, each of which
by itself is capable of producing redness, the varying shades of grain tint
among the progeny of the cross with a white-grained parent (rjr2r3) in
which these factors are absent, being brought about by the additive or
cumulative action of the several colour factors. For example, where two
or three factors are present in the zygote the red tint of the grain is twice
or thrice the depth of colour of those of a plant into which only one has
entered. Only when all the factors are present is the original dark red-
tinted grain of the parent reproduced.

On this hypothesis the gametic constitution of the red parent is
RXRX R2R2 R3R3, that of the contrasted white parent r^ r2r2 r3r3. The
sixty-four possible individuals and twenty-seven genotypes of the F2
generation obtained from the cross RjR-jRg x r1r2r3 may be arranged as
shown in table on top of next page.

The number of red-producing factors entering into the various geno-
types of the F2 generation is indicated at the base of each column of the
diagram ; the frequency distribution of the numbers of individuals of
the several tints of the population is that of a symmetrical polygon of
fluctuating variation.

374

THE WHEAT PLANT

No. of red factors
in Zygote

No. of Zygotes .

1 fe3 : i

Do.

Do.

Do. '

Do.

' allied

Do.

r,r1R2r2R3r3
15:1

Do.

Do.

Do.

Do.

Do.

Do.

Do.

1 alf red3

Do.

R,r1R!r2RaR,
all red

Do.

1 i5":"i

Do.

1 'alf red ' "

Do.

Do.

Do.

Do.

Do.

R1R1r2r2R3r3
all red

Do.

1 aH red " '

Do.

IS : i

RlrjR2R2R3R3
all red

Do.

R1r,R0R2r3r3
all red

Do.

1 Y:2i

Do.

Do.

Do.

Do.

Do.

R1R1R2rzR3R3
all red

Do.

R1r1r2r2R3R3

all red

Do.

3 : i

Do.

ririR2R2R8R3
all red

Do.

r,r1r,!r2R3R3
all red

Do.

RlR1R2R2R3r3
all red

R1Rirlir2R3R3
all red

R1R,R2r!,r3r3
all red

r1rJR2R2r3r3
all red

R1r1r2r2r3r3
3 : i

R R R R R R

Do

R R R R

Do

Do

all red

all red

all red

all white

6

5

4

3

2

i

o

i

6

15

20

15

6

i

In F3 the segregation of the sixty-four individuals of the F2 generation
should occur as follows :

No. of Heterozygous Factors present.

4 or more.

3-

2.

I.

No Red Factors.

Segregation .
No. of individuals

Constant red

37

63 :i

8

I5:i

12

3--I

6

Constant white,
i

Segregation of an F2 generation tested by Nilsson-Ehle gave in F3 the
following proportional numbers :

Constant red I 63 : i

37

12-3

6-6

White,
i

which agree very closely with the expected figures.

HYBRIDISATION AND WHEAT HYBRIDS 375

In addition to " Pudel " wheat, Nilsson-Ehle found that the grain
colour of Svaldf " Grenadier " wheat is a three-factor red.

Investigations showed that the grain tints of Extra Squarehead II.,
" Bore " wheat, and a pure line of Swedish T. compactum are two-factor
reds, hybrids of these with white-grained wheats segregating in F2 in the
ratio 15 red : i white.

A. and G. Howard obtained similar evidence of the existence of double
factors for red grain colour in some Indian forms of T. vulgare and of a
three-factor red in T. compactum, var. erinaceum (" American Club ").

iv. Purple x White. — Caporn studied the inheritance of the purple
tint in a cross made originally by Biffen between T. polonicum with white
translucent grain and the purple-grained Abyssinian wheat " T. Eloboni "
( = T. dicoccum, var. Arraseita (p. 195)).

The Fx grains were all uniformly purple. In F2 were found
(i) fully tinted or " flushed " grains of light and dark shades, (2)
grains with longitudinal streaks of purple, and (3) uncoloured grains
the proportions approximating to the ratio 3 flushed : i streaked : 12
uncoloured.

Some of the ".streaked " plants bred true while others were hetero-
zygous " streakeds " whose descendants exhibited the simple 3 : i mono-
hybrid segregation.

The " uncoloured " plants appeared to be of two kinds, namely (a)
those throwing " uncoloured " plants only, and (b) others giving " un-
coloured " and " streaked " progeny, the approximate ratio of a : b being
15 : i.

Segregation similar to that occurring in F2 was not found in F3 ; other
complicated results requiring further study for elucidation were met with
in the F3 generation.

FLINTY AND MEALY ENDOSPERM. — The inheritance of these characters
is complicated by the fact that their appearance is largely dependent upon
external conditions of soil and climate.

The flinty character is stated by Biffen and Tschermak to be dominant,
but further investigation of the matter is desirable.

SUSCEPTIBILITY AND RESISTANCE TO THE ATTACKS OF RUST-FUNGI
(Puccinia sp.). — The variable incidence of the attacks of Yellow Rust
(Puccinia glumarum, Er. et Henn.) is strikingly evident almost every year
on examining the collection of wheats at Reading in spring.

Upon T. aegilopoides and T. monococcum the rust is never found, and
several forms of T. dicoccoides, T. dicoccum, T. durum, T. turgidum, and
T. vulgare are usually nearly free from the parasite. A larger number
of forms of the same races are slightly attacked, while many forms of
T. vulgare, especially those from India, Persia, and Central Asia, are so

376 THE WHEAT PLANT

severely infected that the upper parts of the young leaves are covered
almost entirely with the bright yellow sori of the fungus, and the surface
of the soil around the plants is bright yellow with the fallen uredospores.

Although the extent of the attack on the different wheats varies from
year to year, the classification into resistant and susceptible forms is easy,
and infection or freedom from attack are found to be constant hereditary
characters.

It was Biffen who first investigated the inheritance of resistance and
susceptibility to the attacks of P. glumarum, and discovered that these
characters form a Mendelian allelomorphic pair. On crossing resistant
forms of T. turgidum (Rivet), T. compactum (American Club), and T.
vulgare (Hungarian Red) with very susceptible forms of T. vulgare
(Michigan Bronze, Preston, and Hungarian White) susceptibility was
found to be a dominant character, the hybrid Fj generation being as
easily attacked as the susceptible parents.

In F2 immune and susceptible segregates were obtained in the pro-
portion of 3 susceptible to i resistant, which bred true in the F3 genera-
tion, immunity proving a recessive character.

While the immune or resistant forms were not completely free from
attacks of the rust, they could be distinguished without much difficulty
from the susceptible plants.

Among the infected segregates there appeared to be different degrees
of susceptibility, and the plants in the F3 bred more or less true to the
variable character. Later Nilsson-Ehle also examined the inheritance of
the same species of rust in hybrids between more or less susceptible and
resistant forms of T. vulgare. He confirmed Biffen's results, but observed
that well-marked dominance of susceptibility was rarely found, the Fx
being intermediate.

In the F2 generation segregation was evident, but the establishment
of definite numerical ratios was not possible, the segregates being finely
graded between those which might be classed as resistant and the most
susceptible plants. Moreover, in some of the crosses transgressive segre-
gation occurred in F2, plants more susceptible and more resistant than
either of the grandparents being found in this and subsequent generations.

Biffen also observed plants more susceptible than either of the grand-
parents in the F2 generation of the hybrid Rivet x Red Fife.

Nilsson-Ehle attributes these results to the existence of multiple
factors for the susceptible and resistant characters (cf. inheritance of
grain colour, pp. 373-375).

The researches of Pole-Evans in South Africa regarding the inherit-
ance of susceptibility and resistance of wheats to the attacks of Black
Stem-rust (Puccinia graminis, Pers.) show that susceptibility is the
dominant, resistance the recessive character as in P. glumarum. In the

FIG. 219.

i. T, aegilopoides, var. Larionowi. 2. Hybrid (F;) of 3 ? x i $ . 3. T. dicoccum, var.farrum.

HYBRIDISATION AND WHEAT HYBRIDS 377

cross Bobs x Wol Koren the hybrid Fl proved to be considerably more
susceptible than the susceptible Wol Koren parent.

Hayes, Parker, and Kurtzweil also investigated the susceptibility to
P. graminis of hybrids of wheats belonging to the races T. vulgare, T.
durum, and T. dicoccum.

The susceptible vulgare wheats used were Preston, Marquis, and
Pioneer, the durums Kubanka and three other resistant forms, in addition
to a form of T. dicoccum (White Spring Emmer), being used as the second
parents.

In the Fj of the vulgare x durum crosses susceptibility was dominant,
and in some of the F2 and F3 generations transgressive segregation occurred,
a few plants being more susceptible or more resistant than either of the
grandparents.

In the vulgare x dicoccum hybrids susceptibility was recessive. There
was also evidence of linkage of the durum and dicoccum characters and
resistance to rust.

I. HYBRIDS OF THE WILD WHEATS Triticum aegilopoides AND Triticum
dicoccoides WITH EACH OTHER AND WITH SPECIES OF Aegilops AND RYE

1. T. aegilopoides x T. dicoccoides. — The wild wheats T. aegilopoides and
T. dicoccoides are very frequently found growing near each other in various
parts of Syria and Palestine, and transitional forms between these were
observed by Aaronsohn. Whether these intermediates were the products
of natural hybridisation or varietal forms of the wild wheats was not
determined.

From a study of the glume characters Schulz considers that some of
the specimens sent by Aaronsohn to Germany as T. dicoccoides were hybrids
of T. aegilopoides, var. Thaoudar, and T. dicoccoides.

I am inclined also to the view that some of the wild wheats which I
received from the eastern side of Mount Hermon were hybrids, the empty
glumes having the characteristic lateral tooth of T. aegilopoides, while the
size and general features of the ear as well as the vegetative characters of
the plants were those of T. dicoccoides.

2. T. dicoccoides x Aegilops ovata. — Cook refers to the discovery in
Syria of a supposed hybrid between the wild dicoccoides and Aegilops
ovata, the plant exhibiting intermediate characters of ear, rachis, and
awns. Aaronsohn considered it a hybrid between the two species, and the
figure of the spikelets given by Cook is suggestive of the cross.

Tschermak, in his papers on " Rare Cereal Hybrids," states that he
crossed T. dicoccoides with Aegilops ovata and also with Aegilops cylindrica,
using the wild wheat as the maternal parent. The hybrids were sterile.

3. T. dicoccoides x Secale cereale. — Tschermak states that Jesenko
crossed the wild T. dicoccoides with rye and found the hybrid sterile.

378 THE WHEAT PLANT

II. HYBRIDS OF THE WILD WHEATS T. aegilopoides AND
T. dicoccoides WITH CULTIVATED WHEATS

1. T. aegilopoides x T. dicoccum.

(a) T. aegilopoides $ x T. dicoccum $ . — Beijerinck records the cross-
ing of these two species, the female parent being a winter black-chaffed
variety of T. aegilopoides (T. nigrescens of Pantcic or T. aegilopoides, var.
Pancicf), the male being the strong-growing form of T. dicoccum. var.
farrum (p. 197).

The Fj_ plants were quite sterile and had ears 18 cm. long each with
38-40 spikelets. The rachis was hairy and very brittle, like that of the
female parent, the form of the empty glumes resembling that of the
male.

(Z>) T. dicoccum, var. farrum ? x T. aegilopoides, var. Larionowi <$ (Fig.
219). — I successfully crossed these two wheats in 1917. The hybrid
(Fx) plant was completely sterile and exhibited characters of both parents.
In length and arrangement the hairs on the upper surface of the young
leaves resembled those of the mother. The ears were fragile, intermediate
in breadth and thickness between the two parents, with reddish empty
glumes like those of the pollen parent, but similar in form and terminating
in an incurved apical tooth like those of T. dicoccum.

The base of the spikelets and edges of the rachilla were as hairy as
those of T. aegilopoides, and the flowering glumes terminated in long
awns like those of the latter plant.

2. T. dicoccoides x with Cultivated Wheats. — In his paper on "Rare
Cereal Hybrids " Tschermak states that T. dicoccoides crosses with T. mono-
coccum, T. dicoccum, T. durum, and T. Spelta, but gives no descriptions.

The wild T. dicoccoides crosses readily with T. vulgar e and T. durum.
Natural hybrids of these occur in Palestine and Syria and are frequently
obtained when the cultivated races are grown in the garden in proximity
to the wild species ; in fact, it is difficult to preserve a pure line of T.
dicoccoides where other wheats are grown.

In the cross made by Tschermak between T. dicoccoides and T. vulgare
the Fj_ plant closely resembled the wild species. In F2 appeared T.
dicoccoides and a mixture of T. dicoccum, T. durum, T. vulgare, and T.
Spelta.

I have also frequently observed the similar production of a variety of
forms of T. dicoccum, T. durum, T. Spelta, and T. vulgare in the F2 genera-
tion from natural hybrid Yl plants found in pedigree rows of T. dicoccoides.

The following is an example of the great diversity exhibited among
the segregates of these sporadic hybrids (see Figs. 220, 221).

A single ear of T. dicoccoides, var. Aaronsohni, gave a natural hybrid
from which were obtained the following forms in the F2 generation :

Jo

o

^ 4>

s a-B

Q fa *•"

M S3 rt

& JJ

| 11

^ II'

I tM^

o o

c 5

0

FIG. 221. — NATURAL HYBRID (T. dicoccoides ? x T. vulgar e (?)).

2, 3,4) 5- Segregates of the Fj generation. 4 and 5 almost sterile.

HYBRIDISATION AND WHEAT HYBRIDS 379

1. Typical dicoccoides.

2. A dicoccoides-\ike form with short awns.

3. A beardless form resembling T. Spelta.

4. A long-eared T. vulgar e.

5. A form with characters of T. dicoccum with short awns.

6. A dense-eared durum-like form.

The F3 progeny of these were :
From plant i, only typical dicoccoides.
,, ,, 2. (a) dicoccoides (red-chaffed).

(b) semi-bearded forms approximating to cultivated T. dicoccum

with non-hirsute rachis, some with red chaff, others with
white chaff.

(c) sterile, beardless T. vulgar e, dark brown chaff.
,, ,, 3. (a) beardless Spelta-like forms, white-chaffed.

(b) sterile, beardless, red-chaffed vulgare.

(c) fertile, beardless, white-chaffed vulgare.

(d) fertile, beardless, red-chaffed vulgare.

„ ,, 4. (a) beardless, lax-eared, red-chaffed vulgare.

(b) beardless, dense-eared, white-chaffed vulgare.
,, „ 5. (a) T. dicoccoides (red-chaffed).

(b) beardless, dicoccum-like forms, white- and red-chaffed,

with smooth or hirsute rachis.

(c) beardless, red-chaffed and white-chaffed sterile vulgare.

(d) beardless, dense-eared, sterile, durum-\ike forms, some of

them " freaks."
,, ,, 6. (a) black-chaffed and red-chaffed dicoccoides.

(b) a long-awned sterile durum.

(c) beardless dicoccum-like forms, some with hirsute, others

with glabrous rachis.

(d) several beardless, dense-eared durum and vulgare " freaks."

Most of them sterile.

In morphological characters the glumes are very variable, and among
the segregates resembling the cultivated wheats there is an almost con-
tinuous series of intermediates between completely sterile forms and those
with all the spikelets fertile. While the majority possess easily disar-
ticulated ears, there are many with various degrees of brittleness of the
rachis and similar diversity in the habit of growth and time of ripening of
the several forms.

In general characters these segregates resemble those obtained by
Tschermak, who states that the F2 generation of a hybrid between T.
dicoccoides and T. vulgare gave a mixture of T. Spelta, T. dicoccum , T.
vulgare, and T. durum.

Some of the progeny including the dense-eared " freaks " are singu-

380 THE WHEAT PLANT

larly like the descendants of a cross between a " mutation from Black
Emmer " (T. dicoccum, var. atratum) and a beardless mutation from the
bearded Turkey red wheat (T. vulgar e, var. ferrugineum) recorded and
figured by Buffum.

Although all the four races of cultivated wheats mentioned are gener-
ally seen among the descendants of these natural hybrids, the relative
proportion of the several races is not always the same ; in the one described
there is a preponderance of fertile vulgar e forms, but in others fertile durum
forms predominate ; the former are presumably crossed with T. vulgare,
the latter with T. durum.

Love and Craig observed the occurrence of forms very closely resem-
bling Wild Emmer in brittleness and hairiness of rachis, form of spikelets
and glumes, and shape and size of grain among the segregates in F2 and
F3 of a cross between Early Red Chief (T. vulgar e. var. milturum] and
Marouani (T. durum, var. leucomelari).

III. HYBRIDS OF CULTIVATED WHEATS WITH WILD GRASSES

1. WHEAT x COUCH GRASS.

Triticum vulgare $ x Agropyrum repens (Beauv.) $ . — Hillman figures
a hybrid between these species and the resulting progeny obtained by
Fr. Strube, but no descriptions are given. He also refers to a similar
hybrid exhibited by G. Bestehorn. Further evidence is necessary before
the hybridisation of these plants can be accepted.

2. WHEAT x ITALIAN RYE GRASS.

Triticum vulgare $ x Lolium italicum (A. Br.) $ . — Schliephacke
mentions the production of a sterile hybrid from the crossing of wheat
with Italian Rye Grass, and gives a figure of the ear of the hybrid plant
without any description. The figure resembles that of a wheat x rye
hybrid, which is also illustrated in the same communication. The evidence
for the crossing of these species is inconclusive.

3. WHEAT x Various Species of Aegilops.

(a) (i.) Aegilops ovata ? x Triticum vulgare $ = Aegilops triticoides
(Req.), Bertoloni. — This hybrid is of much interest since it was the first
wild hybrid grass whose origin was determined by experiment, and its
discovery incidentally led to a great pre-Darwinian discussion among
botanists in Europe and America regarding the fixity of species and the
origin of cultivated wheats. Requien first collected the plant in 1821-24
around Avignon and Nimes in the south of France, and on account of its
resemblance to cultivated wheat he named it Aegilops triticoides. He
published no account of it, but Bertoloni, acknowledging Requien's name,
described the plant in his Flora Italica (vol. i. p. 788), and on the authority

HYBRIDISATION AND WHEAT HYBRIDS 381

of Gussone and Tenore recorded its occurrence in Sicily. It has since
been found in Algeria.

Aegilops triticoides (Figs. 222, 223) is a rare sporadic plant, generally
quite sterile, with tall erect culms 50-90 cm. high, bearded or semi-
bearded, cylindrical ears 8-10 cm. long, consisting of 6-10 or more spikelets,
each possessing two flowers. The empty glumes are flatter than those of
other species of Aegilops and distinctly keeled, with only one or two awns
and rudiments of others. When ripe the ear disarticulates near its base
and falls to the ground (Fig. 223).

Aegilops ovata is a common species
in all the countries bordering on the
Mediterranean. It has somewhat decum-
bent stems 20-30 cm. long and short ears
about 4 cm. long. The latter usually
possess two fertile spikelets, each contain-
ing a pair of grains when ripe, the base
and apex of the inflorescence being com-
posed of a few degenerate spikelets or
spikelets with staminate flowers only.

The empty glumes which almost cover
the rest of the spikelet are ventricose,
rounded on the back, without any distinct
keel, but possessing a number of prominent
nerves ; at the apex are 4 awns (Fig. 223).

When the ear is ripe it disarticulates
below the first fertile spikelet, and after
falling to the ground becomes buried in
the soil, where its grains, tightly enclosed
in the glumes, germinate and give rise to
a tuft of three or four closely knitted but
independent plants.

In 1838 Esprit Fab re, a gardener living
at Agde, near Montpellier, observed that
although the grains of A. ovata usually
give rise to plants like the parent, in some rare instances plants of Aegilops
ovata and Aegilops triticoides are found growing from grains of the same
ear. The fact of two strikingly different plants springing from separate
grains of the same inflorescence attracted the attention of botanists, and
its bearing on the fixity of species was widely discussed.

Fabre cultivated A. triticoides for about twenty years, at first in an
enclosed garden, and subsequently in open fields, and found that the
plants in successive seasons became more and more like Bread Wheat
(Triticum vulgar e).

FIG. 222. — Ear of Aegilops triticoides
and empty glume (nat. size).

382 THE WHEAT PLANT

At first almost sterile and sparing in yield of grain during the first
three or four years, they became by degrees more fertile, the ears lost
much of their brittle character, increased in length, and the number of
flowers in the spikelets rose from 2 to 4 or 5.

The empty glumes lost the broad, symmetrical, convex form of those
of the original mother plant, becoming longer, carinate, and unsymmetrical
like those of wheat, and the number of awns decreased from 4 to i with
mere rudiments of others (3, Fig. 223).

Aegilops ovata never appeared among the descendants, and the latter
in seven or eight years were indistinguishable from ordinary cultivated
wheat of the district, having tall stiff straw and ears with 8-12 spikelets,
each bearing 2-3 fertile grains.

From his observations and experiments Fabre concluded that Aegilops
triticoides was a sport of A. ovata, which under cultivation was gradually
transformed into ordinary wheat, and that all wheats had their origin in
Aegilops ovata.

Fabre's facts were confirmed by others, but his conclusions were the
subject of dispute for a long time.

Regel appears to have suggested that A. triticoides was possibly a
hybrid, but it was Godron who first experimentally established the hybrid
nature of Requien's species. This botanist, in 1853, scattered the pollen
of a beardless form of wheat (Touzelle) on six ears of A. ovata about the
time of anthesis ; from five of these ears only typical plants of A. ovata
were raised, but one grain of the sixth inflorescence produced a short-
awned form of A. triticoides. Later (in 1859) he sowed wheat in the
Botanic Garden at Nancy, in rows 50 cm. apart, and planted ears of A.
ovata between them. In 1860 he collected 872 ears from the Aegilops
plants ; these he put in the soil in February 1861, and from them were
produced twelve plants of A. triticoides in July. He obtained similar
natural hybrids in his own garden in 1870 and 1871 from A. ovata exposed
to the pollen of wheat plants growing near.

The artificial production of A. triticoides was carried out by Godron
in 1853. Two flowers of an ear of A. ovata, after careful emasculation,
were pollinated with pollen from a bearded vulgare wheat ; from these
A. triticoides was raised, the remaining close-pollinated flowers of the
same ear giving rise to plants of A. ovata.

Regel, Greenland, and Planchon also produced the hybrid by care-
fully controlled artificial crossing of A. ovata with beardless forms of
T. vulgare.

A. triticoides in a wild state is nearly always sterile, never abundant,
and always sporadically distributed in the neighbourhood of wheat fields,
having long awns in the districts where bearded wheats are cultivated
(Fig. 222), and short awns where beardless wheats are grown (2, Fig. 223).

It

FIG. 223.

i. Aegilops ovata, L., with empty glume and grains. 2. Aegilops triticoides (Req.), Bertol.

3. Aegilops speltaeformis , Jord., with spikelet, empty glume, and grain.

HYBRIDISATION AND WHEAT HYBRIDS 383

These facts, together with the experimental results of Godron and
others, make it clear that the so-called wild " sports " or varieties of
A. ovata from which Fabre began his cultivations \vere natural hybrids,
the result of chance fertilisations of A. ovata by pollen of wheat from
neighbouring fields.

Godron found that the first cross is sterile, due to the imperfection of
the stamens, but, as discussed later, it yields secondary hybrids with
pollen of its male parent (T. vulgare).

Bally, in 1916 and 1917, made the cross T. vulgar e $ x Aegilops ovata $
and obtained 3 Fx plants from some 40 pollinations. He succeeded also
with the reciprocal cross Aegilops ovata $ x T. vulgare $ , obtaining 2
Fj plants from about 200 pollinations.

In both cases the hybrids were alike and sterile, and resembled those
obtained by Godron and others. The pollen grains varied considerably
in size, fluctuating between 20 and 53 ^ in diameter ; they were more
translucent than those of either parent, and devoid of the fine starch grains
seen in the normal pollen grains of wheat and Aegilops. Bally also studied
the cytology of the two parents and the hybrid, and found that the haploid
number of chromosomes in wheat is 8, that of Aegilops 16. The hybrid

presumably should have 12 ( = J as the haploid number. In some

cases he found this number present, but in others the number was greater
than 12, for during the reduction division some of the Aegilops chromo-
somes remained unpaired and divided longitudinally as in the ordinary
homotypic mitosis. The wheat chromosomes are short and thick, those
of Aegilops much longer and thinner. In the meiotic division of the
hybrid the two parental chromosomes which can be recognised with
certainty travel to the poles of the spindle at different rates, the wheat
chromosomes reaching the poles while those of the Aegilops parent are
still in metaphase.

In addition to the irregular meiotic division, abnormalities occur in
the formation of the pollen tetrads, which result in the production of
multinucleate pollen grains, containing from 2 to 4 nuclei of very variable
size and chromosome constitution.

(ii.) (Aegilops ovata ? x Triticum vulgare $ ) x Triticum vulgare <$ =
Aegilops triticoides ? x Triticum vulgare <$ = Aegilops speltaeformis ,
Jordan. — A fierce controversy arose among botanists regarding Aegilops
triticoides and its transformation into ordinary wheat, the polemic being
most actively maintained for several years by Jordan and Godron.

Jordan with his metaphysical and inflexible views on the immutability
of species at first disbelieved the facts described by Fabre, Godron, and
others. He wras, however, ultimately constrained to admit that A. triti-
coides is a sterile sport or " deformation [of A. ovata] due to hybridisation."

384 THE WHEAT PLANT

In regard to the fertile wheat-like plants obtained and cultivated by
Fabre on an extensive scale he held a novel opinion. Their fertility and
morphological stability over twenty or more generations led him to con-
clude that these were not of hybrid origin, but the descendants of a new,
rare, and unrecorded species of Aegilops, distinct from A. triticoides, but
easily confused with it.

To this imaginary species he gave the name A. speltaeformis , and
suggested that it had found its way unobserved into the south of France
from the East along the Mediterranean.

The plant is annual, with broad, flat, green, or glaucescent leaves,
similar to those of ordinary wheat. The straw is from 60 to 100 cm. high,
with an erect rigid ear, which when ripe falls off as a whole. The ears
are about 10 cm. long (3, Fig. 223), consisting of 10-12 somewhat inflated
flowered spikelets, 2 or 3 of the flowers frequently fertile. The empty
glume is keeled, truncate, with an awn 3-4 times as long as the glume
and usually 2 lateral teeth of variable length. The flowering glume is
obtuse, truncate, with an awn 4 times as long as the glume and 2 short
lateral teeth. The grain is oblong, oval, brownish, and angular with a
deep furrow.

A. speltaeformis has rarely, if ever, been found wild either in France or
in the East, and Godron's view that it is a derivative hybrid produced by
the pollination of the sterile hybrid A. triticoides with wind-borne pollen
of wheat has been generally accepted. Its non-establishment in a wild
state Godron explains by the fact that its fallen ears and spikelets have
not the power of burying themselves in the ground like those of A. ovata
and A. triticoides, and soon perish after germination unless artificially
covered with soil.

Godron states that Fabre obtained grains from ears of A. triti-
coides, which produced the very different and more robust A. speltae-
formis. Greenland also grew the latter plant from a grain of an ear
of A. triticoides gathered on the edge of a field near Agde by Dr.
Thevenau.

The derivation of A. speltaeformis by artificial crossing of A. triticoides
with wheat was proved by Godron in 1858, and again in 1860, but the
cross is apparently very difficult, for no other botanist has been successful
with it. failures being recorded by Regel, Tschermak, and Bally.

Some of the hybrids are completely sterile, others ripen a few grains,
from which more fertile plants arise ; these in turn ripen more grain in
succeeding generations, until plants are obtained completely fertile and
indistinguishable from ordinary wheat.

The increasing fertility and closer resemblance to wheat in succeeding
generations observed by Fabre and by Durieu de Maisonneuve, who con-
tinued its cultivation without interruption for twenty-two years, has been

HYBRIDISATION AND WHEAT HYBRIDS 385

presumed hitherto to be due to successive annual fertilisation by wind-
borne pollen.

It is possible, however, that the later generations are self-fertile, and
that their increasing resemblance to ordinary wheat is brought about by
the elimination of the Aegilops chromosomes in the abnormal heterotype
divisions which Bally found in the primary hybrid A. triticoides.

In these irregular divisions the wheat chromosomes sometimes collect
at the poles of the spindle before the Aegilops chromosomes leave the
equatorial position, and in some of the pollen-grains nuclei may form which
contain only wheat chromosomes, the Aegilops chromosomes or fragments
of them appearing distributed within the cytoplasm of the cells, where
they probably degenerate or enter into the formation of dwarf nuclei,
which take no part in the fertilisation process when such grains are trans-
ferred to the stigmas of wheat flowers.

(b) Aegilops triaristata, Willd. $ xTriticum vulgare $ (=A. triti-
coides, Req.). — Planchon crossed these two species, and Fabre states that
examples of A. triticoides arise from individual grains of fallen ears of
A. triaristata as well as from A. ovata ; thus the name A. triticoides is
used for two different hybrids.

Inflorescences of the hybrid from ovata have the glaucous character
of the mother with somewhat closely packed spikelets ; those from
triaristata are non-glaucous like the mother plant, and yellowish some-
times becoming dark brown when ripe, with the spikelets more distant
from each other on the rachis.

(c) A. ovata 9 x T. Spelta $ (Bearded Form). — Godron records
the production of two examples of this hybrid, but gives no descrip-
tion of them. Greenland also obtained the same hybrid which was
sterile.

(d) A. ovata $ x T. turgidum $ . — Greenland obtained five samples
of this hybrid ; they produced a few grains in the first and second years,
but became quite sterile and died out in the third season.

(e) A. triaristata $ x T. durum " barbatum " <3 , — A long -bearded
hybrid was produced but not described by Godron.

(/) A. ovata $ x by all the different races of wheat.

A. cylindrica $ x by all the different races of wheat.

T. Spelta ? x A. ovata 3 .

T. Spelta 9 x A. cylindrica <$ .

These hybrids were obtained by Tschermak : all were sterile, and
attempts to cross them with the wheat or Aegilops parents failed.

(g) A. triuncialis ? x T. vulgare $ . — Loret discovered this hybrid
wild in the south of France.

(h) A. ventricosa $ x T. vulgare $ . — This hybrid was produced by
Vilmorin and Greenland : it is quite sterile. Henslow also obtained a

2 C

386 THE WHEAT PLANT

sterile hybrid among his cultivations of A, ventricosa which he thought
was due to crossing with T. turgidum.

IV. HYBRIDS OF CULTIVATED WHEATS WITH BARLEY AND RYE

1 . WHEAT x BARLEY. — It is frequently stated that the Australian
wheat " Bobs " (p. 290) is a descendant of a hybrid between a sport from
Blount's Lambrigg wheat (a form of T. vulgare, var. albidum) pollinated
by William Farrer in 1896 with Nepal Barley or Bald Skinless barley, a
six-rowed form with nude grains.

Farrer operated on a dozen flowers of the wheat, taking the usual
precautions to prevent self- fertilisation. A single shrivelled seed was
obtained which gave rise to an Fx with somewhat light green foliage and
weak straw suggestive of barley. The F2 generation showed no evidence
of F2 parentage, but small differences were visible among the several
plants from which " Bobs " appears to have been selected about four
years after the original cross.

Mr. J. T. Pridham attempted the cross Fife wheat ? x Nepal barley $
but without success. From the reciprocal cross (Nepal barley 9 x Fife
wheat cJ), however, he obtained 7 grains; the Fj plants from the latter
when young had narrow, somewhat dark green, wheat-like leaves, but
these characters disappeared as the plants grew, the adult plant having
the broad light green leaves and ears exactly like the barley mother.

Mr. Pridham also tells me that Federation wheat was crossed with a
2-rowed naked barley. The Fx of the Wheat ? x Barley £ cross had
pale green foliage and very weak straw, like the barley parent, the re-
ciprocal Ft having narrower, darker, more wheat-like leaves. " In both
cases," he says, " the F2 progeny showed variations, such as the crossing
of two not very distinct varieties would show in the variable generation,
but no resemblance to barley that could be seen." There appears to be
some justification for the belief in the greater than normal variability
in these " crosses," and it is possible that the stimulus of the foreign
pollen may be connected with its manifestation, but the evidence for the
hybridisation of wheat and barley is inconclusive, and Farrer himself
gives reasons for uncertainty in regard to the matter.

2. WHEAT x RYE.

i. T. monococcum x Secale cereale.

ii. T. polonicum x Secale cereale.

Tschermak states that he crossed these two species of wheat with rye,
the resulting hybrids being sterile. No descriptions are given in his
paper.

iii. T. vulgare x Secale cereale (Fig. 224).

The artificial hybridisation of wheat and rye appears to have been

FIG. 224. — HYBRIDS OF WHEAT (T. vulgare) 9 x RYE (Secale cereale)

i, 2, 3. Natural hybrids. 4. Artificial hybrid. (All from C. E. Leighty.)

HYBRIDISATION AND WHEAT HYBRIDS 387

first accomplished by A. S. Wilson, who recorded in 1875 tne production
of a sterile hybrid from the pollination of wheat with rye. Since that
date similar successful crosses have been described or figured by Carman,
Bernard, Rimpau, Signa, Schliephacke, Miczynski, Tschermak, McFadden,
Ito, Jesenko, Leichty, and Backhouse. Nilsson-Ehle has twice observed
natural hybrids between winter wheat and winter rye.

With the exception of Jesenko's hybrids with the wild Triticum
dicoccoides and Emmer (Triticum dicoccum) all the work has been carried
out with Bread Wheat (T. vulgar e) and rye.

The cross only succeeds when wheat is made the mother plant, and
then only with difficulty.

Jesenko pollinated more than 3500 flowers of rye with wheat pollen
without the production of a single fertile grain ; the reverse cross, how-
ever, he found effective on an average of about six times per 1000 pollina-
tions as indicated in the following results of trials in 1909-1912.

In 1909, 580 flowers on 18 ears yielded i hybrid grain.
In 1910, 339 flowers on 12 ears yielded o hybrid grain.
In 1911, 3060 flowers on 102 ears yielded 29 hybrid grains.

(i ear yielded 12.)
In 1912, 2150 flowers on 70 ears yielded 5 hybrid grains.

Similarly, Leighty obtained no result after 80 pollinations of rye with
wheat, and only 3 grains from 172 pollinations of wheat with rye pollen.
Backhouse also found the crossing of wheat and rye difficult except in the
case of a Chinese beardless variety of T. vulgare, which he states set 32
out of 40 flowers pollinated with rye.

Jesenko discovered that the rye pollen germinates on the stigma and
penetrates the style of the wheat gynaecium, probably reaching the
micropyle of the ovum in the majority of cases, but the union of the sperm
with the ovum is prevented, or the cross - fertilised ovum ceases to
develop.

In some instances grains containing a small amount of endosperm
were obtained apparently as the result of fertilisation, but these refused
to germinate, the embryo being missing or imperfectly formed.

The hybrid plant exhibits the blended inheritance of both parents,
with the rye-like characters perhaps more conspicuous than those of
wheat.

When young, the Ft plants resemble wheat in having a green coleop-
tile and well-developed auricles on the leaves, but the culms and leaves
of the fully developed plants are greyish green, or dark green like those of
rye, and the upper internode, especially of the first and second straws, are
more or less pubescent like those of most forms of the latter plant. The
Hybrids tiller extensively, and continue to produce shoots late in the season

388 THE WHEAT PLANT

in the manner suggestive of a perennial grass. In height and thickness
the culms are intermediate between the two parents.

The ears are long and narrow, frequently longer and having more
spikelets than either parent. The empty glumes are elongated, inter-
mediate in width, strongly keeled to the base, and toothed or scabrid
along the keel like those of rye, but they possess more than one nerve,
and the spikelets are many flowered, like those of wheat.

The absence of beards and the pubescence of the glumes are dominant
characters in these hybrids.

In Tschermak's experiments Squarehead, Theiss, and Banat wheats
were used as the mother plants. Jesenko employed Loosdorf Bearded,
Bokhara, Velvet Squarehead, Red Galician, Epp, and other kinds of
wheat.

Although in general features all the hybrids closely resemble each
other, the specific characters of the particular form of mother plant from
which they are descended are clearly recognisable ; but in cases where
different forms of rye are used as the male parent, their influence is not
visible.

At the time of flowering the glumes open widely, and usually remain
separated from six to eight days. The anthers do not dehisce, and con-
tain mostly shrivelled and imperfectly formed pollen grains of variable
size, with occasional transparent round grains smaller than those of
either parent.

Jesenko and Nakao found that the formation of the tetrads from the
pollen mother-cells was irregular, each of the latter being sometimes
divided into 3, 5, 6, or more degenerate cells.

According to Nakao the larger pollen grains of the hybrid have many
more than the normal haploid number (8) of chromosomes of the parents,
the smaller ones less.

In regard to the fertility of wheat x rye hybrids, Tschermak and
Nakao found the F1 quite sterile, and Jesenko obtained negative results
from more than 3000 flowers pollinated with the hybrid pollen. Carman,
however, harvested 19 grains from 10 ears of one of his Fl hybrid plants,
presumably self-pollinated, and from these grains obtained 14 F2 plants
with 107 very variable ears. Among the plants of the F3 generation were
some bearing ears 7 inches long, others with ears only 2 inches long or
less. The form of wheat Rural New Yorker No. 6, still grown in the
United States, is said to be a descendant of one of these wheat x rye
hybrids.

Rimpau, in 1888, harvested fifteen grains from the Fx plant of the cross
Red Saxon " Landwheat " x Schlanstedt rye grown under a net near a
hybrid wheat : it is not known whether they were the result of self- or
cross-fertilisation. The F2 generation, like that obtained by Carman,

HYBRIDISATION AND WHEAT HYBRIDS 389

was very variable, consisting of plants with glabrous stems and ears of
" Squarehead " form, as well as others with more slender lax ears : some
of the plants were quite sterile, while others yielded viable grain in varying
amounts.

The descendants of Rimpau's hybrid were still grown in various
localities in Germany in 1912, more than twenty generations after the
original cross. Such plants are remarkably constant, and exhibit char-
acters intermediate between the two parents. The ears are long and
narrow, some bearded, others beardless, with strongly keeled scabrid,
rye-like glumes, which open widely and remain separated some consider-
able time at the period of anthesis. Much of the pollen is deformed
and sterile, and the ears set a few grains only, even when pollinated with
wheat pollen. Miczynski also obtained a naturally fertilised F2 generation.

More recently Love and Craig produced hybrids of wheat and rye,
most of which were sterile in Fl and in morphological characters resembled
those obtained by other workers. Two of the hybrids, however, were
fertile. One of them, resulting from the crossing of Dawson's Golden
Chaff wheat (a beardless red-chaffed white-grained variety) with common
rye, had brown glumes intermediate in colour between those of the two
parents, and ciliate keels like those of rye.

The hybrid was not quite sterile. A single F2 plant raised from it had
ears with short awns and sharply keeled ciliated glumes like the rye parent.

Only one seed was obtained from the F2. This gave rise to a more
wheat-like plant (F3) with much increased fertility. The succeeding
generation (F4) had wheat-like ears, some of them fully fertile, others
almost sterile. The grains varied in form ; and in respect of length of
awn, colour of chaff and grain, there was much variation with apparent
segregation in the simple Mendelian 3 : i ratio.

With the exception of the Fx plants, none of the later generations showed
any of the rye-like pubescence of the upper internode. The plants of the
F5 generation showed an increasing resemblance to wheat, although some
of them exhibited traces of rye characters.

The true fertility of the Fj wheat x rye hybrids obtained by Carman,
Rimpau, Love and Craig cannot be decided with certainty, for it is
possible that in these cases the generations after F1 were the result of
chance pollinations from neighbouring wheat plants ; nevertheless, the
negative evidence of Tschermak, Nakao, and Jesenko cannot settle the
matter. It is probable that the fertility may be dependent upon the
varieties of wheat and rye used in the cross.

Crossing the Fx hybrid with one of the parents has been carried out
by Carman and Jesenko. The former pollinated an ear of a Fx wheat x
rye hybrid with rye. From this he obtained 17 grains, from which 14
F2 plants were raised, most of them sterile ; the fertile plants crossed

390 THE WHEAT PLANT

again with rye gave an entirely sterile F3 generation. Jesenko obtained
only a single fertile grain from over 4800 pollinations of Fx flowers with
rye, but he was more successful when wheat pollen was used ; in the
latter case about 3 per 1000 pollinations produced viable grain.

Although these derivative hybrids (Fx plants pollinated with wheat)
more closely resembled wheat than rye, no two individuals were morpho-
logically alike ; some of them were quite sterile, while among the rest
there was a great range of fertility, those most resembling wheat being
most fertile,

Jesenko points out that such results must be due to differences in the
genetic constitution of the egg-cells, since the pollen used was always
the same. He suggests that the carriers of the hereditary characters of
the parents are not equally distributed in the eggs of the Fj plant, some
receiving more of the wheat " plasma," some less, and in the fertilisation
of an egg, containing much wheat " plasma " with the gamete of a wheat
pollen grain, the resulting hybrid would more closely resemble wheat
than would be the case where union takes place between a similar wheat
gamete and an egg containing a smaller proportion of wheat " plasma."
In the latter case the hybrid would be less like wheat and more inclined
to be sterile like the original Fx hybrid, which is half wheat and half rye.

V. HYBRIDS OF THE DIFFERENT CULTIVATED RACES OF WHEAT

All the cultivated races of wheat have been successfully crossed with
each other. Those in which T. monococcum is a parent are completely
sterile ; the rest give fertile progeny, although the degree of fertility is
very variable.

In the hybridisation of T. durum, T. turgidum, T. vulgar e, and T.
Spelta, Vilmorin observed forms of all four races among the descendants
of hybrids between either of the two former and one of the two latter,
from which he concluded that all these wheats had a common ancestor.

Tschermak obtained similar results in his experiments. Kezer and
Boyack found T. Spelta among the F2 segregates of the cross T. dicoccum
x T. vulgare (p. 393), and T. durum appeared in the F2 of the hybrid
T. dicoccum x T. sphaerococcum which I produced at Reading (p. 392).

i. HYBRIDS OF T. monococcum WITH OTHER WHEATS. — Hybrids of
T. monococcum with T. dicoccum, T. vulgare, T. durum, T. turgidum,
and T. polonicum have been obtained by various workers. Tschermak
states that he successfully crossed T. monococcum with all cultivated
wheats.

The hybrids are sterile and have the flat brittle ear, keeled glumes,
and marked lateral nerve and tooth of T. monococcum, but show the
influence of the other wheats in the larger ears, clearly developed terminal

HYBRIDISATION AND WHEAT HYBRIDS 391

spikelet, and the presence of two awns on each spikelet : crossing of the
F! with the parent forms does not succeed.

a. (i.) T. monococcum, var. flavescens 9 (Engrain double) x T. dicoccum,
var. farrum $ (Amidonnier blanc). — This hybrid was obtained by Bei-
jerinck in 1882 and its reciprocal a little later. Both were similar to each
other and quite sterile ; the gynaecium was apparently perfect, but the
anthers did not open or shed pollen. The leaves of the hybrid were
yellowish-green, and the nodes of the stem hairy like those of T. mono-
coccum.

In general appearance the ears were intermediate between the two
parents ; the empty glume had a curved apical tooth, somewhat similar
to that of the male parent, the characteristic lateral tooth of T. monococcum
being very much reduced or absent.

(ii.) T. dicoccum, var.pycnurum $ x T. monococcum, var.flavescens <$ . —
This hybrid was obtained by Vavilov, from whom I received specimens
of the parents and the ¥l plant in 1914. The hybrid was sterile with
ears intermediate in size between those of the two parents. In the
glabrous polished surface and reddish colour of the glumes, as well as
in the velvety surface of the leaves, it resembled the mother plant.

b. T. monococcum x T. durum. — Blaringhem obtained the hybrid T.
monococcum, var. vulgar e x T. durum (an Algerian form). Three hybrid
grains were produced by the same ear.

The Fj plants grew vigorously, their ears resembling those of T.
dicoccum. One of the plants was completely sterile, the other two were
semi-fertile.

The F2 and F3 generations of the latter consisted of plants with the
vegetative characters and late-ripening habit of T. monococcum. Most of
the ears were sterile and resembled those of T. dicoccum : the fertile ears
had fewer spikelets and were more like those of T. durum.

Tschermak also obtained a hybrid between T. monococcum and T.
durum.

c. T. monococcum x T. polonicum.

T. monococcum, var. flavescens $ x T. polonicum, var. compactum,
Link <$ .

Blaringhem succeeded with this cross in 1913. From one cross-
pollinated ear of T. monococcum seven shrivelled grains incapable of
germination were obtained, while from another ear of a different plant of
the same pure line growing near and pollinated at the same time seven
viable well-developed grains were produced. In both cases the pollen
parents belonged to the same pure line, but were different individuals.

Tschermak also obtained a hybrid between T. monococcum and T.
polonicum.

d. T. monococcum x T. turgidum. — The hybrid T. turgidum $ x T.

392 THE WHEAT PLANT

monococcum $ was produced by Biffen in 1903. The Fl plant was
damaged by Puccinia graminis and did not flower. Tschermak obtained
a similar hybrid.

e. T. monococcum x T. vulgare. — Cross pollinations of these two wheats
were made without result by Vilmorin in 1880.

Tschermak records the .production of a few grains from a similar cross
in 1906 : the Fx plants, however, do not seem to have been described.

In 1911, after some failures, Vavilov obtained a single shrivelled grain
from the cross T. vulgare, var. erythrospermum ? x T. monococcum, var.
flavescens $ . The mother plant was an early, white, smooth-chaffed,
bearded wheat with red grain, the pollen parent being the variety of T.
monococcum known as " Engrain double."

The hybrid (Fx) was a well- developed plant with five stems about
80 cm. high, ripening about two months later than either of the parents.
The ears were flattened and fragile, and the empty glumes strongly
keeled with conspicuous lateral nerves as in T. monococcum, while the
straw was hollow and the nodes glabrous as in T. vulgare. The plant was
sterile. Its glumes separated and remained apart several days at the
flowering period. The gynaecium and lodicules were apparently normal,
but the stamens were small with non- dehiscing anthers containing very
small, wrinkled, and transparent pollen-grains. T. monococcum is immune
to rust fungi, but the hybrid was readily inoculated with Brown Rust
(Puccinia triticind) and showed the dominant susceptibility of the maternal
parent.

2. HYBRIDS OF T. dicoccum WITH OTHER WHEATS.

a. T. dicoccum x T. monococcum (see p. 391).

b. T. dicoccum x T. durum. I have no records of this cross.

c. T. dicoccum x T. polonicum (see p. 394).

d. T. dicoccum ? , var. atratum (Black Emmer) x T. sphaerococcum,
var. globosum $ (Punjab Type 5).

Black Emmer has tall hollow straw, bearded flat ears, with blue-black
pubescent glumes, red grain, and is a late variety ; the Punjab wheat is
very early, and possesses very short, hollow straw, short square beardless
ears, with pubescent glumes and white, flinty grain.

The hybrid which I obtained in 1917 had ears exactly like those of
the dicoccum parent, so much so that it appeared as if no crossing had
occurred. The F2 generation consisted of a very varied series of red-
grained dicoccum forms, some with black glumes like the grandmother
and late, others of intermediate flowering period, with uniform white
glumes, uniform red glumes, white glumes with black margins and red
glumes with black margins respectively. In addition to the dicoccum
forms, there appeared one or two plants of a typical T. durum, with glabrous

HYBRIDISATION AND WHEAT HYBRIDS 393

leaves, red glumes, white grain, solid straw, and the early habit of the
male grandparent.

On account of the irregular development of the glume pigment in
the homozygotes and the intergrading of glume colour among the hetero-
zygotes, accurate classification by eye-inspection alone was impossible.

The F3 generation showed, however, that the F2 generation consisted
of the following segregates :

T, dicoccum forms :

(1) Homozygous black glumes or blackish glumes with darker margins.

(2) Homozygous uniform white glumes.

(3) Heterozygous white glumes with black margins, splitting into i

white : 3 grey or blackish glumes.

(4) Heterozygous uniform red glumes, splitting into i white : 3 red.

(5) Heterozygous red glumes with black margins, splitting into i white :

3 white with black margins : i red : 3 red with black margins.
T. durum form :

(6) Uniform red glumes.

Factors for black, red, and white glumes were present in the F1} those
for black and red being apparently brought in by the black-glumed
Emmer parent, the allelomorphic pairs of the cross being black and red,
and black and white respectively.

The original parental dwarf sphaerococcum type of ear and straw
disappeared altogether in this cross, and was not recovered as a recessive
in the F2 or F3 generations of the hybrid, its place apparently being taken
by the durum form : the white colour of its grain was seen, however, in
the durum, and the peculiar curved apex of its empty glumes was visible
among the majority of the segregates of F2 and F3.

The factor for beardlessness was not dominant in F1 of this cross,
and its influence was only slightly seen in the F2 and F3 among a few of
the heterozygous plants (No. 3 above) which had shorter awns than the
normal dicoccum.

In all these forms pubescence was linked with the black character.

(e) The hybrid T. vulgare, var. lutescens (Fultz Mediterranean) x
T. dicoccum, var. atratum (Black Emmer), obtained by Kezer and Boyack,
had the flattened ears and the dark, adherent, keeled chaff of the Emmer
parent. The ears were " beard-tipped " and non-glaucous, the latter
character being found in the vulgare parent.

The F2 generation was extraordinarily diverse, from sixty- two to sixty-
five distinct forms appearing, among which were examples of T. dicoccum,
T. vulgare, and T. Spelta. In outline, density, and other characters the
ears were equally diverse, many being quite unlike either grandparent.

Numerous grades were found between long and short, and lax and
compact.

394 THE WHEAT PLANT

The colour of the glumes was also very varied, some had white chaff,
others black, but few or none were as black as those of the Emmer grand-
parent.

As in the previously described Black Emmer hybrid a number had
red or brown glumes, and pubescence was linked with the black character.

Many plants were sterile or semi-sterile.

3. HYBRIDS OF T. durum WITH OTHER WHEATS.

a. T. durum x T. monococcum (see p. 391).

b. T. durum x T. polonicum (see p. 395).

c. T. durum x T. vulgar e.

(i.) T. vulgare, var. albidum (Chidham d'automne) x T. durum
(Ismae'l Velvet Chaff). — The Fj of this cross produced by Vilmorin
in 1879 had square beardless ears with a few short apical awns.

The F2 consisted of an extraordinary series of forms of T. durum and
T. vulgare, some with long awns, others beardless : both lax- and dense-
eared vulgar es were obtained with examples of T. compactum, T. turgidum,
and T. Spelta. None were like the original parents of the cross.

(ii.) T. vulgare, var. milturum (Early .Red Chief) x T. durum, var.
leucomelan (Marouani). — The ¥l of this cross made by Love and Craig
was a vulgare-like form, " beardless " with short, apical, black awns and
red grain.

In the F2 generation, segregation in regard to both chaff colour and
grain colour occurred in the proportion of 15 red : i white, the ratio of
the " beardless " to fully bearded being 3:1. Among the 113 descendants
were forms of T. durum and T. vulgar e, and in addition, two typical plants
of wild T. dicoccoides with flat ears, brittle rachis, long hairs at the base of
the spikelet, and narrow, elongated grain : the proportion 113 : 2 (wild
forms) suggests the trihybrid ratio 64 : i. Some of the plants of this
generation had ears with " reversed " awns, the empty glumes bearing
very long beards, while the awns of the flowering glumes were quite
short.

4. HYBRIDS OF T. polonicum WITH OTHER WHEATS. — Artificial hybrids
have been obtained between T. polonicum and most of the other cultivated
races of wheat. Several natural hybrids have also been observed.

With the probable exception of the cross T. monococcum x T. polonicum ,
the hybrids are fertile or semi-fertile.

a. T. monococcum 9 xT. polonicum <$ (p. 391).

b. T. polonicum ? x T, dicoccum $. — (i.) In 1916 I discovered a
single hybrid ear (i, Fig. 225) among the progeny of a pedigree row of
T. polonicum, growing in close proximity to pure lines of T. dicoccum.
The polonicum parent had white papery chaff, 30-33 mm. long, slightly
hairy on the nerves, and white flinty grain 10-11 mm. long. The

FIG. 225. — NATURAL HYBRID (T. polonicum ? x T. dicoccum

2, 3, and 4. Segregates of the F2 generation.

HYBRIDISATION AND WHEAT HYBRIDS 395

dicoccum parent, I have no doubt, was a white-chaffed European form,
with empty glumes about 10 mm. long, terminating in a curved apical
tooth, and grain red and flinty, about 9 mm. long.

The Fj ear was narrow, about 10 cm. long, with brittle rachis, white
awns, and empty glumes about 15 mm. long, clothed with fine, adpressed
hairs ; the grains were dark red, flinty, and 9-5 mm. long.

The F2 from this gave three clearly differentiated types, viz. :

(1) T. polonicum, almost typical in form and size of ear, with some-
what brittle rachis, thin, quite glabrous, empty glumes, 25 mm. long, and
dark red flinty grain 10 mm. long.

(2) A form closely resembling the white-chaffed European T. dicoccum,
with fragile rachis, stout empty glumes, finely pubescent, about n mm.
long, with a strong, incurved, apical tooth, and dark-red flinty grain :
some of the ears had fine black awns.

(3) An intermediate type with brittle ears, nearer in character to the
dicoccum than the polonicum parent, the empty glumes quite glabrous,
intermediate in texture, and 15-17 mm. long, ending in an intermediate
incurved tooth, the grain 10 mm. long, dark red and flinty.

The types (i) and (2) bred true in the F3 and F4 generations : type (3)
behaved as the Fj plant. The young leaves of all the plants were pubescent
like those of the dicoccum parent ; none were glabrous like polonicum.
Brittleness of ear and red grain were dominant, and the white grain of
the polonicum parent did not reappear in the F2 or F3 generations.

(ii.) T. polonicum x T. dicoccum, var. Arraseita (" T. Eloboni"). — This
hybrid obtained by Biffen in 1913 was investigated by Caporn later.

The average glume-length of the polonicum parent was 29-23 mm.,
the glume-length of the dicoccum parent ranging between 8 mm. and
13 mm.

The Fx plants had intermediate ears with glumes intermediate in size
and form between those of the parents.

The grain was also intermediate in size, but purple like that of the
dicoccum parent.

The F2 generation showed indistinct 1:2:1 segregation, with the
glumes of the homozygous " long-glumed " plants — 24-14 mm. long —
somewhat shorter than those of the polonicum grandparent : those of the
homozygous " short-glumed " segregates being longer than those of the
corresponding dicoccum grandparent.

c. T. polonicum x T. durum (Kubanka). — A cross between these wheats
was made in 1912 by Backhouse.

The average glume-length of the polonicum parent was 29 mm., that
of the durum parent 12 mm. The Fx was intermediate, with glume-
length 18-19 mm.

In F2 wrere long-glumed and intermediate-glumed plants, forming a

396 THE WHEAT PLANT

group inseparable by eye into two classes, but distinct from the short-
glumed segregates. The numbers of each were :

Long and intermediate, 172 ; short, 55 ; or a 3 : i ratio.

The Polish wheat used in the cross had faintly pubescent glumes,
Kubanka being a glabrous-chaffed wheat. The Fj was more pubescent
than the polonicum parent.

In F2 both smooth and hairy-glumed plants appeared. Among the
pure short-glumed plants were three felted to one with smooth glumes :
the proportion of pubescent to smooth in the heterozygotes was 85 : 3 1
(15 strongly felted), while all the pure long-glumed plants had glabrous
chaff, the factor for long glume apparently inhibiting pubescence.

Results of a cross between the Polish and Kubanka wheats (T. durum)
were also investigated by Engledow.

The F! was intermediate. The glume-length of the F2 progeny was
determined and compared with the glume-length of the grandparents,
the measurements being made of a glume from one of the four median
spikelets of each ear.

The mean glume-length of the Polish wheat used in the cross was
30-84 mm., that of the durum being 11-7 mm.

In F2 three glume-length types appeared, the measurements of the
glumes yielding a tri-modal frequency curve, the respective modes being
11-5 mm., 16-5 mm., and 25-5 mm. The calculated frequencies of the
F2 segregates were :

23-29 per cent (Extracted Kubanka), 57-76 per cent (Intermediate),
18-94 Per cent (Extracted Polish).

To determine more accurately the classification of the F2, the F3 genera-
tion was raised and measured : the percentages of the three types were :

23-65 (Kubanka), 55-39 (Intermediate), 20-95 (Polish).

The figures suggest a i : 2 : i ratio, but deviate considerably from it.

A comparison of the measurements of the grandparental forms and
the extracted polonicums and durums in F2 showed that the true polonicum
and durum types are produced in very small numbers if at all in F2, the
extracted polonicum having a mean glume-length from 20 to 25 per cent
less than that of the original polonicum, and the extracted durums some-
what longer glumes than the grandparental Kubanka.

The reduction or " shift " towards the mean glume-length of the two
grandparents is most clearly established in the long-glumed or " poloni-
cum " segregates of F2, and the " shifted " form breeds true in F3, no
evidence of additional " shift " or a reappearance of the pure polonicum
being obtained.

Engledow found that in regard to the length of the grain the Fx

HYBRIDISATION AND WHEAT HYBRIDS 397

was intermediate between the parents, and segregation into long-,
intermediate-, and short-grained plants took place in F2.

The extracted polonicums of F2 showed the same phenomenon of
" shift " in the length of the grain as in the glume-length, but the reduc-
tion was less, the grains being only about 12-5 per cent shorter than those
of the grandparental polonicum.

Engledow found an increase in pubescence in some of the short-
glumed F2 plants similar to that observed by Backhouse.

d. T. polonicum x T. turgidum. — These wheats have been crossed by
Vilmorin, Tschermak, Biffen, Backhouse, and others, and natural hybrids
between them have been recorded by Jordan.

(i.) T. polonicum (Bearded) x T. turgidum (Petianelle Blanche, Bearded).
—This hybrid was raised in 1881 by Vilmorin. The Fx plant (1882) had
very long, lax, beardless ears, with empty glumes intermediate in length
between the two parents.

In the F2 generation (1883) were examples of almost pure T. polonicum ,
together with forms of T. vulgar e and T. durum. Although both original
parents had white grain, some of the segregates had red grain.

(ii.) T. polonicum ? x T. turgidum $ (" Rivet "). — Biffen records the
crossing of these wheats, the polonicum parent being the common lax-
eared form with rachis internodes 6-6 mm. long, pale yellow glumes,
almost or entirely glabrous, averaging about 28 mm. long, and
white grains : the turgidum plant was the dense-eared cone type most
commonly cultivated in this country, with rachis internodes about 3-6 mm.
long, greyish pubescent glumes about 9 mm. long, and reddish grain.

The Fj had lax intermediate ears, rachis internodes 5-8 mm. long,
with pale greyish or dirty-yellow pubescent glumes, intermediate between
those of the parents, measuring 17 mm. in length. The grains of the
F! plants were red, about 9 mm. long, those of the parents averaging
10-1 mm. (Polish) and 7-2 mm. (Rivet) respectively.

Glume-lengths of 595 plants of the F2 generation were measured, and
the frequency distribution of the several lengths plotted ; the curve
obtained was sharply divided into three portions, segregation into 149
long-glumed : 304 intermediate : 142 short-glumed plants, suggesting the
monohybrid ratio i : 2 : i .

The plants of this generation bore long, intermediate, and short
grains, and segregation into white- and red-grained plants also occurred.
Biffen found that the chaff colour of the hybrid and its descendants
resembled that of the pale polonicum parent. The blue-grey colour of
the Rivet ancestor, which in crosses with some other wheats is dominant,
was suppressed in this case and did not reappear in any of the plants up
to the sixth generation.

A Polish wheat with white and practically glabrous glumes was crossed

398 THE WHEAT PLANT

by Backhouse with a dark pubescent glumed variety of T. turgidum.
The average glume-length of the polonicum parent was 28-29 mm-> that
of the turgidum 1 1 mm.

The Fx was intermediate in length of chaff, varying from 14 to 17 mm.
The F2 generation consisted of a series of plants impossible to separate
by eye into long-, intermediate-, and short-glumed classes. The frequency
distribution of measurements of the glume-lengths, however, when
plotted appeared to give a ratio of 514 long and intermediate : 178 short.

The Fj generation had felted white or faintly coloured glumes.

In F2 were pubescent and smooth-glumed segregates, and white,
ringed, and deeply coloured ears, the most highly felted and deepest
colour being associated with short glumes, the factor for long glume
apparently inhibiting colour and pubescence.

e. T. polonicum x T. vulgare. — The F1 of this cross obtained by
Tschermak had empty glumes intermediate in length between those of
the two parents.

In the F2 generation polonicums, intermediates and vulgare forms with
glumes somewhat longer than those of the vulgare parent were obtained,
and in addition a small number (about 6 in 100) of new durum forms
appeared.

Neither the polonicum nor vulgare forms of this nor subsequent genera-
tions were exactly like the parents of the cross.

The proportion of the several forms were : vulgar es, i ; durum, 3 ;
intermediates , 8 ; polonicums , 4 ; the ratio of the vulgare forms to the
rest being i : 15, suggesting the presence of two pairs of factors.

Some of the durum forms were found to be constant, others segregated
in the ratio 3 durum : i vulgare.

From the foregoing records of the hybridisation of T. polonicum with
other wheats it is seen that the Fx is always intermediate between the
two parents in length of ear, form, size, and texture of glume, and form
and size of grain.

In the majority of cases segregation into polonicum, intermediate and
the second grandparental types occurs in the F2 generation, the ratio
being i : 2 : i .

In the natural cross between T. polonicum and T. dicoccum I found
the homozygotes readily distinguishable from the intermediate hetero-
zygotes, but where the second parent is a durum or turgidum it is practi-
cally impossible to classify the F2 generation into three groups by eye
inspection alone, since the ears obviously long-glumed and short-glumed
are always connected by a closely graduated series of intermediates.

In the latter cases, however, the curve obtained by plotting the fre-
quency distributions of the measurements of the glumes is tri-modal,
corresponding to the three types of segregates.

HYBRIDISATION AND WHEAT HYBRIDS 399

The mean glume-length of the extracted F2 polonicums is usually
25 per cent less than that of the grandparental polonicum, the glumes of
the other extracted homozygotes being somewhat longer than those of the
corresponding grandparents.

The means of the curves of the glume-lengths of the homozygotes
are nearer to the mean of the intermediates than are the means of the
original parents. This condensation effect is permanent, the changes in
the average glume-length of the extracted homozygotes being hereditary
and transmitted without further apparent alteration to the subsequent
generation.

It is possible that the observed deviation of the extracted homozygotes
from the pure parental forms in the crosses of polonicum with other
wheats may be due to irregular reduction division in the Fl generation.

In normal reduction division, a certain number of the gametes pro-
duced in the Fx generation contain only polonicum chromosomes, and a
certain number only durum chromosomes. In irregular reduction
division, however, the gametes may never receive chromosomes derived
entirely from either parent, and the zygotes from the fusion of such
gametes would therefore always exhibit the characters of both parents in
greater or lesser degree.

5. HYBRIDS OF T. turgidum WITH OTHER WHEATS.

a. T. turgidum x T. monococcum (see p. 391).

b. T. turgidum x T. polonicum (see p. 397).

c. T. turgidum x T. Spelta,

T. turgidum (Rivet), var. dinurum x T. Spelta, var. Duhamelianum (Red
Tyrol Winter Spelt. Beardless Red Chaff). — This hybrid obtained by
H. Stoll gave an extraordinary number of forms in F2, among which were
several " Squareheads " with and without awns and a beardless compactum.

The glumes were white, red, or bluish and pubescent, like the Rivet
parent. One of the beardless Spelta forms with red chaff and large,
plump spikelets and grain was selected and remained constant after 1904,
when it was placed on the market as Stoll 's Early Giant Spelt.

The Fj in the Spelta-vulgare and Spelta-turgidum crosses is inter-
mediate in ear characters, with a more or less brittle rachis and tightly
fitting glumes, like the Spelta parent. So far as can be gathered from the
published accounts, segregation occurs in the F2 generation in the expected
Mendelian ratios, and no forms appear of races other than those of the
two grandparents.

d. T. turgidum x T. vulgare.

i. T. turgidum (Rivet), var. iodurum $ x T. vulgare (Red Schlan-
stedt) S- — Rimpau crossed these wheats in 1875. The Fx resembled
the male parent. In the F2 were many intermediate forms between
completely bearded and beardless.

400 THE WHEAT PLANT

The characteristic four-sided ear of turgidum was absent.

The glumes were chiefly red with some of bluish tint suggestive of
the turgidum grandparent ; a few forms with white chaff appeared.

He also crossed Rivet with a " Squarehead " white-glumed wheat,
obtaining a beardless ¥l with pubescent chaff of several shades of bluish
black.

The F2 exhibited a large variety of forms, some of which were quite
sterile.

ii. T. vulgare, var. albidum 9 (Chidham d'automne) x T. turgidum,
var. dinurum (Roux velu de Beauce).

iii. T. vulgare, var. pyrothrix ? (B16 seigle) x T. turgidum <3 (Poulard
Ble Buisson). — Vilmorin made these crosses in 1879. The F1 in both
cases was " beardless " like the vulgare parents, with short awns at the
tip of the ear, the glumes slightly downy and somewhat inflated like those
of T. turgidum.

In the F2 generation appeared an extraordinary series of vulgare and
turgidum forms — tall, dwarf, bearded and beardless, some with lax ears,
others with very dense ears. In addition, from the hybrid (ii.) was obtained
a plant with the pith, glumes, and long, pointed, flinty grain of T. durum,
while among the progeny of hybrid (iii.) were forms of T. Spelta, some of
them with branched ears.

6. HYBRIDS OF T. vulgare WITH OTHER WHEATS.

a. T. vulgare x T. monococcum (see p. 392).

b. T. vulgare x T. durum (see p. 394).

c. T. vulgare x T. polonicum (see p. 398).

d. T. vulgare x T. turgidum (see p. 399).

e. T. vulgare x T. compactum (see p. 403).
/. T. vulgare x T. Spelta.

The above crosses give fertile reciprocal hybrids, but according to
Stoll success is most easily secured when T. vulgare is the mother.

i. T. Spelta, var. album $ (White Beardless Spelt) x T. vulgare, var.
ferrugineum $ (Bearded Red Chaff). — In 1876 Rimpau made this cross,
the Fx bearing almost beardless Spelt-like ears with slightly reddish chaff.

The F2 in 1878 gave all the eight possible combination forms of the
three pairs of allelomorphic characters (bearded and beardless, red chaff
and white chaff, tightly fitting and loose chaff) except white beardless
T. vulgare.

In the F3 all eight forms were found, together with a white, dense-
eared compactum form.

ii. T. Spelta, var. Duhamelianum (Rose imberbe), Beardless, Red
Chaff x T. vulgare, var. anglicum (Blanc k duvet), Bearded, White Velvet
Chaff. — Vilmorin records the crossing of these in 1878.

The Fj was intermediate, the ears reddish, pubescent, with somewhat

HYBRIDISATION AND WHEAT HYBRIDS 401

fragile axis and tightly fitting glumes, though less so than the Spelt
parent.

In F2 many forms were found, some Spelt-like with red and white
chaff, others like ordinary wheat with white and reddish chaff, both
glabrous and pubescent : none were completely like either of the parents
first crossed.

iii. T. vulgare, var. lutescens (Main's Stand-up), Beardless, White
Chaff x T. Spelta, var. Duhamelianum (Beardless Red-chaffed Winter
Spelt). — Two constant Spelt wheats were obtained by H. Stoll from this
hybrid, one with white and the other with red chaff. The red-chaffed
form, placed on the market in 1902, has beardless ears, wide spikelets,
with plump, flinty grain and strong straw, and is highly resistant to frost
and rust.

The following successful hybrids of T. Spelta and T. vulgare were also
made by H. Stoll :

Bordier x Red Tyrol winter Spelt.

Heine's Squarehead x Red Tyrol winter Spelt and its reciprocal.

iv. T. Spelta, var. Duhamelianum (Beardless, Red Chaff) x T. vulgare
(Squarehead, Beardless, White Chaff). — The Fx of this cross made by
Miczynski had lax, beardless ears and wide spikelets. Four different
segregates were found in F2, viz. : 4 Spelta forms, 8 like F1 ; 3 Square-
heads ; i compactum form or i Spelta ; 3 non- Spelta forms.

7. HYBRIDS OF T. compactum WITH OTHER WHEATS.

Triticum compactum x T. vulgare. — Many hybrids have been obtained
between these races of wheats, the chief results being described below :

i. T. compactum, var. Humboldtii (Little Club) x T. vulgare.

ii. T. compactum, var. rufulum (Red Chaff Club) x T. vulgare.

In 1899 Spillman crossed " Little Club " and " Red Chaff," two
varieties of T. compactum, with several sorts of long-eared varieties of
T. vulgare.

The Fj in the majority of cases was intermediate in length and density
of ear between the parents. In F2 there appeared long-eared, short-
eared, and intermediate plants : in some instances the numbers of each
approximated to the ratio i long-eared : 2 intermediate : i short-eared,
while in others every gradation was found between the grandparents, and
in addition, some examples with both longer and shorter ears than
these.

Where " Farquhar " — a long-eared, beardless, brown velvet-chaffed
T. vulgare — was crossed with the beardless little Club, the F2 in some
cases consisted chiefly of long-eared individuals with few or no inter-
mediates or compactums.

iii. T. vulgare ? (Red King, Beardless) x T. compactum $ (Rood
Koren, Bearded). — In 1902 Wilson crossed Red King, a long-eared,

2 D

402 THE WHEAT PLANT

somewhat dense, beardless form of T. vulgare (said to be a hybrid), with
Rood Koren, a bearded T. compactum from South Africa.

The Fj plants had squarish compact ears of intermediate length with
a few awns about i inch long at the tip, the compactum parent being fully
bearded with awns 2^-3! inches long.

In F2 four forms appeared, viz., short-eared, long-eared, awnless, and
awned, the numbers of each being :

664 awnless, 207 awned ; 643 short-eared, 229 long-eared, the ratios
approximating to the monohybrid 3 : i type.

Little variation in length and density of ear was observable among
the long-eared segregates, but the short-eared forms were less uniform,
and a few plants had exceptionally compact ears.

The long-eared, beardless plants of this F2 generation gave in F3
almost entirely long-eared, beardless progeny, the bearded F2 plants
giving a similar preponderance of awned, long-eared descendants.

The short-eared, bearded plants of F2 gave mostly bearded progeny
in F3, the length of ear showing considerable variation : the corresponding
short-eared, beardless forms proved to be similar in constitution to the
F! hybrid, and, like it, gave four types in F3.

iv. T. compactum^ var. Humboldtii ? x T. vulgare $ (Epp Wheat). —
Riimker crossed these wheats, the Fx being a typical " Squarehead "
intermediate in ear-density between the two parents. In F2 segregation
was of the monohybrid 1:2:1 type, this generation consisting of 21 Epp :
37 Squarehead : 20 compactum.

A Squarehead of F2 split similarly in F3, giving 18 Epp : 49 Square-
heads : 21 compactum.

v. T. vulgare ? (White Frankenstein) x T. compactum <$ (White Velvet
I gel). — This cross obtained by Rimpau gave more complex results.
The Fj generation consisted of four plants, one, a typical beardless
Squarehead, the others produced both beardless long and lax ears and
Squarehead-like ears on the same plants.

In F2 were obtained five forms, viz. :

a. 21 beardless Squareheads.

b. 12 beardless, dense-eared, not typical Squareheads.

c. 5 bearded Squareheads.

d. 8 lax-eared, beardless plants of the Frankenstein type.

e. i lax-eared, bearded plant.

38 dense : 9 lax (4 : i).

41 beardless : 6 bearded (7 : i).

Two of the beardless Squarehead plants (a) gave all bearded descendants
in F3.

FIG. 226. — SEGREGATES RAISED FROM THE INDIVIDUAL GRAINS OF EACH
SPIKELET OF AN EAR OF A NATURAL HYBRID ( T. -vulgar e x T. com-
pactum), arranged in order from the base (i) to the apex (2) of
the ear.

HYBRIDISATION AND WHEAT HYBRIDS 403

From one of the bearded Squareheads (c) were obtained in F3 u
bearded glabrous and 35 beardless glabrous Squareheads

= Bearded, i : Beardless, 3.

Another gave 15 bearded Squareheads, 58 beardless Squareheads, and
2 lax-eared, beardless plants.

Bearded, i : Beardless, 4.

vi. T. vulgare 9 (Velvet Kolben, Beardless) x T. compactum $
(Velvet Igel, Bearded). — The Fx of this cross obtained by Riimker was
of intermediate ear-density with white velvet chaff and beardless ears.

In F2 appeared plants with bearded and beardless ears, some long
and lax, others of intermediate length and denser, though none were of
the " Squarehead " type. Although both grandparents were velvet-
chaffed, glabrous-glumed forms appeared in this generation.

Ratios : — Velvet-chaffed, 3 : Glabrous, i. Bearded, 5 : Beardless, 101.

Biffen crossed a long-eared vulgare (Devon) with T. compactum
(Hedgehog) and found the Fx intermediate. Strampelli also obtained an
intermediate Fx hybrid from T. compactum (Herisson) x a lax, long-eared
T. vulgare (Rieti), the F2 giving compactums, intermediates, and long-
eared forms in the ratio 1:2:1.

vii. T. compactum (Swedish Binkel) x T. vulgare, — The crossing of
T. compactum with T. vulgare was extensively studied by Nilsson-Ehle,
who crossed Swedish Binkel, an old beardless form of T. compactum, with
three long, lax-eared forms of T. vulgare.

The Fj plants had ears somewhat longer and laxer than those of the
compactum parent; resembling an ordinary " Squarehead." In F2
appeared long, lax-eared, homozygous, recessive forms sharply separated
from a group of " Squarehead " heterozygotes and homozygous com-
pactums, the two latter not always clearly differentiated from each other,
but whose genetic constitution was revealed in the F3 generation.

The ratios of the F2 were of the ordinary monohybrid type, viz. :

i compactum : 2 Squareheads : i Long-eared.

The same variety of T. compactum was also crossed with " Pudel "
and " Bore " wheats — comparatively dense-eared " Squarehead " forms
of T. vulgare.

The F! of these crosses was dense-eared, closely resembling the
compactum parent. Segregation in F2 was complex, no sharp distinction
being visible between the compactums and non-compactums.

In one example of the cross Binkel x Pudel (Squarehead), the F2
consisted of 61 compactums, 5 of them denser than the " Binkel " grand-
parent ; 5 compactum-\\ke plants and 28 non-compactums, chiefly very
lax-eared, only two being as dense as the Pudel grandparent.

404 THE WHEAT PLANT

The progeny of these in F3 is indicated below :

F2 generation. F3 generation.

( 5 very dense compactums . . 5 constant compactums.

j'i8 constant compactums.
56 compactums .... - 38 segregating into compactums and non-

compactums.
5 compactum-\ike ... 5 segregating into compactums and non-

compactums.

28 not compactums . . . Only non-compactum progeny, but of

variable ear length and density.

The F3 results show that the F2 generation consisted of 23 homozygous
compactums, 43 heterozygotes, and 28 homozygous non-compactums ,
suggestive of the simple monohybrid 1:2:1 ratio.

Similar gradation of density of ear between the compact and laxer-
eared segregates occurred in the F2 generation of the cross " Binkel " x
" Bore " wheat, but the nature of the plants of the F2 generation was
cleared up in the F3 progeny :

F2 generation. F3 generation.

f 25 constant compactums.
46 compactums . . . . •: 21 splitting into compactums and non-

compactums.
31 less dense compactums . . Splitting into compactums and non-

compactums.

1 1 Binkel " Squareheads " . . Do.

i " Squarehead "... Do.

37 plants with lax ears . . . Descendants all non-compactums, but

exhibiting considerable variation
in ear-density.

The results of the F3 generation show that the F2 consisted of 25
homozygous compactums, 64 heterozygotes, and 37 homozygous non-
compactums, approximating as in the previous cross to the 1:2:1 ratio.

The results obtained by Nilsson-Ehle led him to believe that in the
long lax-eared " Land wheats " two independent factors Lx and L2 are
present, each of which, by itself, increases the length of the internodes of
the ear ; when both factors are present their effects are cumulative.

The dense ear of T. compactum he considers due to a factor C which
shortens the rachis-internodes and is epistatic to Lx and L2.

On this hypothesis plants of the genetic constitution CL^L^, CL^,
C^Lo, as well as those containing the C factor alone (Cljlg), would have
short compactum ears, though not all of equal ear-density, since the L
factors are not completely masked by the presence of C.

From analysis of his results he concluded that the Swedish Binkel is a

HYBRIDISATION AND WHEAT HYBRIDS 405

CLjL2 compaction, and that the " Squareheads," Bore and Pudel, are of
the constitution cl^, neither the factors C nor Lx or L2 being present in
these wheats.

In the crossing of T. compaction CLjL2 with T. vulgare, where the
vulgar e parent is of the long lax-eared form cLjLg, the compactum is
generally dominant but not completely so, the Fx often resembling the
Squarehead " wheats. In F2 segregation occurs into long-eared plants,
intermediate and compactums in the ratio i : 2 : i, but exact measurements
of the internodal lengths of the rachis are often necessary to establish the
ratios. The heterozygotes, though found to be somewhat less dense than
the homozygous compactums, form with them a group sharply differ-
entiated from the long lax-eared plants.

New combinations of the factors in the cross between a compactum of
the constitution Cljlg and a lax-eared " Landwheat " cLxL2 would result
in the production of " Squareheads " (cl^) in F2 : these were found by
Nilsson-Ehle, though their number was not so large as the expected i in 3.
The " Squareheads " of Rimpau's hybrids may have arisen in this way.

In crosses between T. compactum and the denser-eared vulgares or
Squareheads, the Fx very closely resembles the compactum parent. Segre-
gation in F2 is complex and apparently impure, long lax-eared forms
longer than the Squareheads, and compactums denser than the Binkel
appearing with forms resembling the two grandparental wheats.

The cross between the Binkel compactum and Bore " Squarehead "
may be represented thus :

CLXL2 (Binkel) x cl^ (Squarehead).

From the hybrid ¥1 four forms would appear in F2, viz. : Cl^ (very
dense compactum), CLjL2 (Binkel), cl^ (Squarehead), cLjL^ (long lax-
eared forms) ; the first and last being new hereditary types respectively
denser and laxer than the compactum and Squarehead grandparents.

As previously noted, all these were obtained by Nilsson-Ehle, and the
appearance of the very dense and very lax ears in some of Spillman's com-
pactum x vulgare hybrids may be explained in a similar manner.

The C factor acts more strongly in shortening the ear internodes of
the long-eared wheats than those of the Squarehead form, and the com-
pactums CL1L2, CL]12, CljL.2, Cl^ form a graduated group less homo-
geneous than the non-compactums (Squareheads \j\2 and " Landwheats "
LXL2) of the same cross.

8. HYBRIDS OF T. sphaerococcum WITH OTHER WHEATS. — T. sphaero-
coccum x T. dicoccum (see p. 392).

9. HYBRIDS OF T. Spelta WITH OTHER WHEATS. — T. Spelta x
T. vulgare (see p. 400).

406 THE WHEAT PLANT

VI. HYBRIDS OF VARIETIES AND FORMS OF THE SAME RACE

OF WHEAT

By far the greatest number of wheat hybrids have been produced by
the crossing of varieties and forms belonging to the same race, and the
hybridisation of the vulgare forms is assiduously carried on by many
workers in almost all the large wheat-growing countries.

The inheritance of the several differentiating characters is generally in
agreement with the conclusions summarised in pages 363-377.

CHAPTER XXVI

THE IMPROVEMENT AND BREEDING OF WHEAT

THE amelioration or improvement of plants consists in securing and pro-
pagating individuals possessing in greater degree than the unimproved
form those characters which render them useful or attractive to man.

Such improvement is dependent upon three factors, namely : (i)
variation, (2) heredity, and (3) artificial selection.

If all plants were exactly alike no improvement would be possible ;
moreover, in the case of plants, such as wheat, which are propagated from
seeds, unless the desirable variation from the ordinary type is hereditary it
is useless.

It is important to emphasise the fact that the creation of plant char-
acters is beyond human power, and the causes which lead to the produc-
tion of hereditary variations are quite unknown.

Given, however, the improved hereditary variant, all that is necessary
is to select and isolate it from its neighbours and grow it. Such selection
is, in fact, the only means of improving plants. Selection, of course, does
nothing more than select, and in itself produces or creates nothing,
although it is frequently tacitly assumed to be causally responsible for the
modifications which are secured.

The new individuals which are selected may arise spontaneously among
ordinary crops, or they may appear among the progeny of artificial hybrids.

It should, however, be clearly understood that in the improvement of
plants hybridisation does nothing more than manufacture material upon
which selection can be exercised. Without selection, it leads to the pro-
duction of a multiplicity of forms, and consequent confusion.

By crossing, individuals are obtained which differ from the parental
forms, but the differences are due only to a recombination of already
existing characters and not to the creation of any new character.

In the improvement of wheat three methods of selection are practised,
namely : (i) mass selection, (2) selection of spontaneously occurring
individuals, and (3) the selection of plants with desirable characters from
the descendants of artificial hybrids.

407

408 THE WHEAT PLANT

Mass Selection. — The selection of a number of the best ears from an
ordinary crop of wheat, and the grading of the grain by sieves and other
machines in order to obtain the largest for use as seed, has been practised
by the most advanced agriculturists in all ages. Virgil (Georgics, i. 197)
says :

" I've seen the largest seed, tho' viewed with care,
Degenerate, unless the industrious hand
Did yearly cull the largest."

Columella (De re rustica, ii. 10) also refers to the necessity for
sifting out the largest grains for use as seed, and mentions that Celsus
advises, where the crop is small, to pick out all the best ears and store
them for seed ; Varro (De re rustica, i. 52) similarly advises the
selection of the best ears for seed.

The process of mass selection, as generally practised, consists in
choosing from the standing crop 100, 1000, or some other arbitrary
number of ears possessing characters which the plant-breeder considers
it desirable to fix or increase. The choice may be based upon the excep-
tional length or density of the ear, the number of grains in the spikelets,
the height of the straw, or other characters. The grains from these
selected ears are sown, and from their progeny a further selection is made,
and the process repeated annually as long as it is deemed desirable.

When carried out by skilled observers acquainted with the morpho-
logical differences of wheat plants, mass selection rapidly leads to the
establishment of a uniform type of plant from a variable crop, and appar-
ently new or improved forms have been frequently obtained by this
means in various countries. When the characters upon which the selec-
tion is based are correlated with prolificacy of the plants, it results in the
production of a crop whose yielding capacity gradually increases from
year to year as the selection is continued.

Where improvement occurs, it is the custom to assume that the crop
is, in the first instance, a mixture of a number of distinct and independent
forms and the selection does nothing more than isolate one of these from
the rest, just as a white-chaffed form would be established by selecting all
the ears with white chaff from a field mixture of white- and red-chaffed
wheats.

Whatever the explanation, the fact remains that mass selection has
led to an improvement in the yield and uniformity of the crop when
applied to fields of wheat as ordinarily cultivated on the farm, and it is
a method which should not be abandoned nor despised.

Apart from the possibility of obtaining new forms by its means, the
process is invaluable in maintaining the purity of those wheats which have
descended from single ears.

Individual Selection. — The selection of single ears or plants and their

IMPROVEMENT AND BREEDING OF WHEAT 409

subsequent propagation has produced the majority of the world's most
famous wheats.

The practice was initiated by Le Couteur and Shirreff in the early
years of the nineteenth century, and was subsequently adopted in this
country by Hallett, by Vilmorin in France, Nilsson at Svalctf in Sweden,
and Hays in the United States.

All the widely grown English wheats of last century originated in this
manner. Chidham wheat was derived from a plant found in a hedge in
Sussex ; Fenton came from a three-strawed plant discovered in a quarry
in Scotland ; Hunter's White was the progeny of a plant taken from the
roadside, and Browick and Squarehead wheats were descended from single
" rogue " plants growing among crops of another kind.

Similarly, the American Fultz originated from three beardless ears
observed in a field of Lancaster wheat, and the renowned Canadian Fife
was derived from a single plant grown from a mixed sample of imported
European wheat.

The true ancestry of these selected plants can only be surmised. In
some instances it is probable that they are stray plants of kinds already
cultivated elsewhere. Others are doubtless ordinary Mendelian segre-
gates from recently produced natural hybrids. The majority, however,
appear to be hereditary sports or mutations arising out of plants whose
remote ancestors were hybrids. A few may be variants unconnected with
hybridisation and due to causes which are at present unknown.

After the initial selection of a special ear or plant, the time required
to obtain a sufficient quantity of grain to sow in the field is comparatively
short, enough for the ordinary seeding of 15 to 20 acres being easily
secured at the third harvest, without adopting any very special methods
for increasing the crop.

The following is a typical example of the rapidity of multiplication of
a new form of wheat from a single ear, the results being obtained by Mr.
Jonas of Liverpool in 1838-41.

Fifty grains taken from a selected ear were sown (dibbled) in 1838.
Only thirty grew, but the produce from these was 14! oz. From this
quantity he obtained i^ bushels in 1839, which returned 45 bushels in
1840. From the latter amount he harvested 537 bushels in 1841, four
years after the selection of the single ear.

A good ear usually yields from 4-5 to 5 grs. of grain, which when
sown on good land in rows 9 inches apart each way will produce rarely
less than 450 grs. — about i Ib. at the first harvest. This amount sown
thinly yields about i bushel of grain or 60 Ibs. at the second harvest,
which when sown on an acre of good land in clean condition generally
produces 35 to 40 bushels (third harvest).

Shirreff laid special emphasis upon the initial selection, and con-

4io

THE WHEAT PLANT

sidered that after this was made, all that was needed was to grow and
multiply the selected plant.

Hallett, on the other hand, did not specially seek for exceptional
plants, but endeavoured to improve those already in cultivation, by
means of " pedigree " culture, believing that the good qualities of the
plant could be increased by this process, which consists in repeated
selection of the best plants in successive generations, and propagation
from these only.

Beginning with a single plant or ear, he made an annual choice of the
most prolific individual, being convinced that as good effects of continued
selection would be secured among the crops as among the animals of the
farm.

The grains from the chosen ears were sown singly in rows i foot
apart, the grains being deposited in holes i foot asunder in the row. At
harvest the individual plants were examined, and the best in respect of
their grain production, length of ear, or tillering power was selected for
further propagation. From this elite plant the best ear was isolated and
its grain sown, the plants produced being subjected to the same careful
scrutiny at the following harvest.

By this process he considered that the good qualities of the original
selected plant were not only maintained but increased from year to year.

The following are the results which he records after four years' selec-
tion of Red Nursery wheat :

Length.

No. of
Grains.

No. of Ears
on the Best Plants.

inches.

1857. Original ear .

4£

47

1858. Finest ear

6i

78

10

1859. Finest ear

7f

91

22

1860. Ears imperfect owing to

wet season .

. .

39

1 86 1. Finest ear

8|

123

52

" Thus," he states, " by means of repeated selection alone the length
of the ears has been doubled, their contents nearly trebled, and the tillering
power of the seed increased five-fold."

These figures only indicate an increase in the length and number of
grains of the finest ears in successive generations, but give no evidence of
any improvement in the averages of these characters. They may indeed
be taken to indicate nothing more than wider fluctuating variability in
carefully cultivated progeny of selected ears.

Nevertheless, Hallett, who was an advocate of thin seeding, secured
some extraordinary crops of grain from his pedigree seed, in one instance

IMPROVEMENT AND BREEDING OF WHEAT 411

obtaining if bushels on 698 square feet (108 bushels per acre) of un-
manured land, from pedigree Nursery Red wheat seed sown singly in
rows 9 inches apart each way. Further, there was in his lifetime a wide-
spread belief in the superior quality of his pedigree seed.

In recent times there has been much scepticism regarding the possi-
bility of obtaining modifications of particular characters by repeated or
" pedigree " selection applied to the progeny of a single self- fertilised
plant such as wheat.

The doubt has arisen in consequence of the experimental results of
Johannsen, who failed to secure any significant alteration in the average
size of Dwarf beans (Phaseolus vulgaris) by selecting either the largest or
the smallest seeds in successive generations of self-fertilised descendants
which originally sprang from a single individual bean.

To the progeny of a single homozygous organism propagated by self-
fertilisation, Johannsen applied the term pure line, and his evidence and
that of others has led to the conclusion that selection within such a line
has no effect.

All that can be legitimately concluded, however, from these experi-
ments is that variations of the fluctuating type (p. 346) are not inherited ;
but to assume that so-called pure lines are incapable of heritable variation
is unwarranted, for hereditary sports or mutations, though somewhat
rare, do occur among the descendants of wheats raised from single ears.

Commencing with a single ear of average length, repeated " pedigree "
selection will lead with certainty to the production of a line with an
increased average length if hereditary sports with longer ears arise during
the time selection is continued.

Since mutations are erratic, often small, and comparatively rare, the
time taken to obtain improvements by selecting within lines descended
from single individuals will necessarily be subject to much variation.
Nevertheless, in spite of the irregularity of the results, the method should
not be discontinued or cast aside as useless.

Selection of Hybrids and their Descendants. — The observed variability
of hybrids and their offspring soon led to the employment of hybridisation
as a means of producing desirable varieties from which improved forms
could be obtained by selection. In the formula of the plant-breeders of
last century, crossing of stable varieties was practised in order to " break
the types " or induce them to sport, after which the fixation of desired
forms by repeated selection was undertaken.

All the early wheat hybrids were produced with this object. Among
the most prominent and successful breeders who adopted this method
of wheat improvement were Shirreff in this country, Vilmorin in France,
Rimpau in Germany, Pringle and Blount in the United States, and
Saunders in Canada.

412 THE WHEAT PLANT

At first the crossing was of a simple character between two kinds of
wheat, the grains of the hybrid plant being sown and selection made from
the progeny. Later, Farrer in Australia, Jones in America, and the
Garton Brothers in England resorted to multiple crossing, the first hybrid
being immediately crossed with a third form, and the plant produced
crossed with another wheat, successive hybridisations with different
forms being continued for several seasons.

The following is an example of such multiple crossing, from which
the Australian wheat Bunyip was derived :

Blount's Lambrigg x Hornblende

Improved Fife x Purple straw An unnamed hybrid x King's Jubilee

Rymer 9 Maffra <$

Bunyip.

In the majority of cases there was little or no system in the choice of
parents of these simple multiple hybrids, the chief object being to obtain
immediate variation, and wheats chosen at random frequently gave satis-
factory results from this point of view.

Farrer and others, however, first obtained a clear conception of the
desired improvements, and selected the plants to be crossed only after a
careful study of their morphological and physiological characters, in some
instances subjecting the parents to pedigree culture before hybridisation.

Although the methods of the early hybridists were of a somewhat
haphazard nature, they led to the production of many valuable wheats,
among which may be mentioned Vilmorin's Bon Fermier and Dattel,
Pringle's Defiance, Jones's Winter Fife and Early Red Clawson, Saunders's
Marquis and Preston, and the Australian wheats Federation and Come-
back, produced by Farrer.

Before 1900 the result of hybridisation was a matter of chance and its
nature could not be foreseen, since no reliable generalisations were avail-
able regarding the transmission of the characters of the parents to their
offspring in the case of sexually produced plants.

About the date mentioned the rediscovery of the Mendelian laws of
inheritance removed some of the uncertainty of hybridisation, and the
plant-breeder is now able to predict with a high degree of assurance the
character of the offspring derived from the crossing of parents possessing
certain morphological and physiological features.

Many of the characters of the wheat plant, such as the presence or
absence of awns, the colour of the chaff and grain, the hairiness and
smoothness of the chaff, and the solid or hollow condition of the straw are
dependent upon single Mendelian factors, which are transmitted from
parent to offspring independently of each other, and the plant-breeder is

IMPROVEMENT AND BREEDING OF WHEAT 413

able to treat these as separate hereditary units, which can be added or
removed at will.

Moreover, he is able to combine in one plant many of the characters
possessed by two or more different varieties, and from a knowledge of the
established Mendelian ratios the selection of desirable forms which will
breed true is simplified and rendered more certain. For example, from
a red-chaffed, red-grained wheat and a white-chaffed, white-grained
variety it is easy to obtain both white-chaffed red-grained and red-chaffed
white-grained individuals, and plants constant to these new combinations
of characters can be selected with certainty from the second generation of
the hybrid's descendants.

By crossing any of the forms just mentioned with plants having
glabrous or pubescent glumes, these latter characters can be introduced
into the progeny of the hybrids, and, similarly, the bearded or beardless
characters can be added to plants in which they are absent.

In the chapter on hybridisation (Chap. XXV.) the mode of inheritance
of many of the different characters of the wheat plant is indicated so far
as it has been determined, and the knowledge can be utilised for the pro-
duction and breeding of plants in which are present any combination of
the separate characters there discussed.

Unfortunately, colour of chaff and grain, presence and absence of awns,
and most of the characters whose inheritance has been clearly established
are of no economic importance.

The character which above all others it is desirable to control and
utilise is the grain-yielding capacity of the plant, but this, like almost all
quantitative characters, either does not Mendelise or is beyond present
Mendelian analysis, and its manifestation is so greatly influenced by so
many external conditions that its inheritance is obscured.

Other characters which it is desirable to improve on account of their
bearing upon yield are length of growing-period, resistance to drought,
frost, and lodging, and immunity to diseases, all of which are of a highly
complex nature, and their inheritance in terms of Mendelian factors still
uncertain.

Whatever methods are adopted for the improvement of wheats, it is
essential that a clear conception should be obtained of the ideal towards
which the plant-breeders' efforts are directed. This is especially neces-
sary since the aims of the breeder vary in different countries ; the wheat
which best suits the English farmer is useless in Central Europe, India, or
Australia, and vice versa.

Moreover, real progress can only be made by those who are thoroughly
familiar with the morphological details and variability of the wheat plant,
and the raising of improved kinds adapted to the requirements of the
grower to be successful must be carried on in the country in which the

4H THE WHEAT PLANT

varieties are to be cultivated and under the conditions which allow of the
full development of the desired characters.

Where high grain production is desired, selection of forms possessing
a naturally long growing period is essential. Nevertheless, the climatic
conditions of the country in which the improved wheats are to be grown
must be considered, for, although the long-lived English wheats give
excellent yields in this country, they are unable to complete their life-
history in India, and are useless there, being easily surpassed in grain
production by the rapid-growing short-lived endemic forms.

On the other hand, Indian varieties are valueless here on account of
their short growing period and consequent low yield.

The breeding of drought-resistant forms is not likely to be successful
when prosecuted in a humid climate, nor can rust-resistant varieties be
secured or isolated in the absence of rust fungi.

In all wheat-growing regions the primary object of the cultivator is to
obtain the highest financial return from the land, and in the majority of
cases this result is only secured by the growth of wheats which give the
greatest yield of grain. High yield is, in fact, the ultimate aim of the
plant-breeder when he directs his attention towards the production of
varieties which are resistant to drought, frost, and diseases.

So-called " strength " of grain is important, but wheats of the highest
quality in this respect invariably give small yields, and the consumer or
his agents rarely pay enough for the superior quality to cover the loss due
to diminished yield. It usually pays the producer to grow wheat of
inferior milling quality, and this has been specifically recognised and
adopted as a sound policy by the most successful wheat growers during
the last two hundred years in this country.

Throughout Western Europe wheats of the highest quality have been
abandoned as unprofitable, and in other regions where the cost of cultiva-
tion is increasing, wheats of high quality and low yield are being replaced
by better yielding sorts of inferior quality.

The growth of wheat of inferior milling quality is sometimes con-
demned, but as Cobb has well said : " Farmers do not grow wheat for
philanthropic reasons ; they grow it to make money, and it will be a long
time before they grow it for any other reason." ..." Give the grower
a new wheat that will bring him more money for his outlay, and he will
grow it whether the consumers starve on it or grow fat."

CHAPTER XXVII

YIELD

THE world's annual harvest of wheat at the present time reaches the
enormous total of more than 3500 millions of bushels, and efforts to
increase it are continually being made.

Its cultivation is being extended into new country, and in districts
where its growth is already established there is unceasing endeavour to
increase the yield by improvement in the methods of cultivation, the
application of fertilisers, the selection of new varieties better adapted to
the climatic and soil conditions, and the use of superior seed.

In the table below are given approximate figures for the total amount
raised annually in the various wheat-growing countries, together with the
average yield per acre.

1908-1913.

1908-1913.

Average
Annual Yield.

Average
Yield
per Acre.

Average
Annual Yield.

Average
Yield
per Acre.

millions of bushels.

bushels.

millions of bushels.

bushels.

Denmark

4i

?

Chili .

21

*4

Belgium

15

37

Spain .

130

13

Holland

5-

35

Australia

90

12

Switzerland

3i

32

India . ...

351

12

United Kingdom

60

32

Asiatic Russia

120

12

Germany .

153

32

European Russia 666

II

Sweden

3i

Argentina .

147

II

New Zealand

7

(3i)

Greece

(8)

II

France

31?

20

Mexico

(5)

IO

Austria

60

2O

Uruguay

7

IO

Japan . .

24

20

Persia

(16)

IO

Canada

204

19

Turkey in Europe

(22)

II

Hungary

169

19

Turkey in Asia .

(33)

10

Roumania .

88

19

Algeria

35

IO

Italy

• 183

16

Portugal

8

9

Egypt . .

34

16

Union of S . Africa

6

9

Serbia .

14

15

Tunis .

6

6

Bulgaria

46

15

—

United States .

685

14

The world's average . -13

At the head of the list in point of total yield stand the United States

415

4i6 THE WHEAT PLANT

and European Russia, each with an annual harvest of over 600 millions of
bushels. Occupying the third and fourth places, with yields of over 300
millions, are France and India, followed by Canada with some 200 millions.
Other countries whose harvests yield from 150 to 180 millions of bushels
are Italy, Hungary, Germany, and Argentina.

While all countries which grow wheat use it for bread-making, some
of them produce more than they need and export it to others whose crop
is insufficient for their requirements.

From 500 to 600 millions of bushels are needed by the importing
countries, about four-fifths being supplied by the United States, Canada,
Russia, and Argentina. Of all countries the United Kingdom imports
by far the greatest amount, some 200 millions being required annually.
Holland and Belgium absorb about 60 millions each, while Italy takes
between 40 to 50 million bushels.

While the total yield of grain which a country produces is of national
interest, the yield per acre is of primary importance to the farmer, and
all his efforts are directed towards improving it.

In a class by themselves may be placed the countries in which the
average yield is over 30 bushels per acre, and with the exception of New
Zealand all are found in Western Europe. The conditions governing
these high yields are the rainfall during the growing period and the
climatic conditions which allow of the cultivation of winter wheats with a
long growing period, together with the intensive cultivation and manuring
of the soil which is practised in these countries.

In contrast with these countries are the large numbers in which the
average yield is below 20 bushels per acre. On account of the severity
of the winters, the low rainfall, or the very brief growing season, only
spring wheats of short growing period can be grown, and the difficulties
connected with the cost of better cultivation and manuring of the land
militate against higher yields.

The average yields of wheat in this country have risen gradually from
the earliest times down to the present day. Rogers states that 8 bushels or
less was the average yield during the first half of the fourteenth century,
and records of 3 to 6 bushels are not infrequent at that period, the amount
sown being about 2 bushels per acre.

In the thirteenth century Walter of Henley gives 10 bushels per acre
as a fair return, and incidentally remarks that if the yield is only three
times the amount of the seed sown (i.e. 6 bushels per acre) a farmer will
gain nothing unless the price of grain is abnormally high.

In the work on " Hosebonderie." written in the early part of the four-
teenth century, it is said that " wheat ought by right to yield to the fifth
grain," i.e. 10 bushels per acre.

From this period up to the middle of the sixteenth century, yields of

YIELD 417

8 or 10 bushels were considered normal. There was no rotation of crops,
and little or no manure was applied to the soil, which was worn out and
poorly cultivated.

When the soil became exhausted the land was fallowed and allowed
to lie idle for a whole year, during which period it was ploughed several
times. After this treatment returns of 8 to 10 times the amount of seed
sown (i.e. 16-20 bushels) were not infrequently obtained. The ground
was thereafter cropped annually, the yields diminishing until no more
than a three- or four-fold return was secured, when the fallowing process
was again repeated.

With the introduction of stock-raising in the early part of the seven-
teenth century a greater amount of manure became available, and the
land was better cultivated, average yields rising to 12 or 14 bushels per
acre.

Improvement continued slowly during the seventeenth century, and
at its close average yields of 1 8 to 20 bushels were estimated by contem-
porary writers.

In the eighteenth century further progress was made, and the average
of 23 or 24 bushels per acre is given for the wheat crop by the authors of
the county agricultural surveys carried out for the whole of Great Britain
during the last few years of the eighteenth and the first ten years of the
nineteenth centuries.

In the middle of the nineteenth century Pusey, Caird, and others
estimated the yield at 26 or 27 bushels ; at the end it had risen to 30
bushels.

The average yield per acre during the first twenty years of the twentieth
century has been 31 bushels per acre.

It can be safely assumed that the increase in the yield from an average
of 8 bushels in the fourteenth century to one of 27 bushels in the middle
of the nineteenth century was a real improvement in the average yield per
acre, due chiefly to an extended application of manures and more efficient
cultivation of the soil.

Regarding the more recent advance of 3 or 4 bushels in the yield per
acre during the last sixty or seventy years, there is less certainty in respect
of its real character and cause. It is probable that the increase is not due
to improvement in cultivation or general management, but to the fact
that land which gave poor return has been utilised for other purposes,
only that specially suited to wheat being retained for this crop. Under
such circumstances the average yield per acre would rise even if the
manuring, cultivation, and seasons remained unchanged or were worse
than before.

Real progress is only made when the increase in the yield per acre is
obtained from the same area, and it is doubtful if any material improve-

2E

4i 8 THE WHEAT PLANT

ment has been made in this sense during the last half century or more in
this country. There is much evidence that most of the good land was
highly farmed seventy to a hundred years ago, and rarely gave less than
40 bushels per acre.

Moreover, in Great Britain the higher and lower limits of yields per
acre during the last hundred and fifty years or more appear to have been
much the same, records of 50 and 60 bushels per acre are frequent as far
back as the middle of the eighteenth century, and Bradley in 1757 states
that " the largest produce of an acre of wheat land, so far as has yet been
experienced, seems to be 10 quarters or 80 bushels per acre," a yield which
is rarely equalled at the present day.

Norden in 1607 observes that in Somersetshire on well-tilled good
land " they have sometimes in some places 4, 5, 6, 8, yea, 10 quarters in
an ordinary acre."

The largest yield per acre hitherto recorded in any part of the world
upon a field of more than i acre is 117-2 bushels, which is given in the
United States Monthly Crop Report for July 1918. The crop was grown
in 1895 on a field of 18 acres in Island County, Washington, U.S.A., the
soil being described as a black sandy loam overlying a clay subsoil. The
variety of wheat was Australian Club. No manure had ever been applied
to the land, and in the three previous seasons potatoes were grown on the
field.

In 1918, Mr. Alfred Amos of Wye, Kent, obtained a yield of 96
bushels (63 Ibs.) per acre of " Yeoman " wheat on an area of about 3^ acres.
The seed was sown on November 20, 1917. The soil of the field is a
deep rich loam overlying 6 feet of brick earth. No fertiliser was applied
to the wheat crop, but the previous crop of beetroot was well manured.

Morton (Journ. Royal Agric, Society, 1859) reports a yield of 90 bushels
per acre obtained in 1844 on a field measuring 5^ acres in the parish of
Haisborough in Norfolk.

The wheat grown was " Spalding," sown in the autumn of 1843 at
the rate of 3 bushels per acre.

Land almost adjoining gave 80 bushels per acre in the same season.

The lower limits of the yield of wheat as ordinarily grown in Great
Britain are 10 or 12 bushels per acre, returns of this amount being not
infrequent during the last hundred and fifty years.

The yield per acre is dependent upon a very large number of factors.
Some of them cannot be altered by the farmer, while others are within
his power to modify and control.

To the former class belong climate, season, and original physical and
chemical condition of the soil ; among the latter are drainage, cultivation,
and manuring of the soil, the yield per ear and the number of ears grown
per acre.

YIELD

419

Climate. — That an adequate temperature, a certain light intensity, and
a suitable supply of water are essential conditions for the growth of all
crops is well understood, but the influence of variations in each of these
factors upon the yield of the wheat crop has not been exactly investi-
gated.

The highest yields of wheat are only obtained in districts where the
mean winter temperature is comparatively high, the rainfall 20 to 30
inches during the growing season, and bright warm weather at the time
of the development and ripening of the grain.

It is well known among the agriculturists of this country that an ex-
cessive rainfall during the winter is disastrous to the yield of autumn-
sown wheat, and especially so where the sowing has been delayed. The
root-system and the young shoots of the plants are seriously checked in
their development, doubtless by the want of adequate aeration of the
water-logged soil and the loss of soluble food constituents in the drainage.

It has been remarked by many observant farmers of the last two
centuries that upon the same farm in one season the yield per acre has
sometimes been 30 bushels, while in other seasons upon the same fields
with similar cultivation, manuring, and general treatment of the crop, the
return has been less than 20 bushels.

In Great Britain the best season of last century was 1863, when the
wheat crops throughout the country were extraordinarily good : the worst
was 1879, when the yield was estimated at less than half the average.

The difference between these seasons is strikingly illustrated by the
returns, given below, of the wheat crop upon several plots at Rothamsted
which have received the same manures and general cultivation annually
for over sixty years.

YIELD OF GRAIN PER ACRE

Best

Worst

Average

Plot.

Manures.

Season

Season

Difference.

6 1 Years

(1863).

(1879).

(1852-1912).

bushels .

bushels.

bushels.

bushels.

3

No manure

i7i

4f

iai

12-6

2

Farmyard manure

44

16

28

35'2

5

Mixed mineral manures

19*

51

14

H'5

6

Do. NH4 salts (43 Ibs. N)

39f

loi

29i

23-2

7

Do. NH4 salts (86 Ibs. N)

53f

rtj

37l

32-1

9

Do. nitrate of soda (86

551

22

33l

Ibs. N)

8

Do. NH4 salts (129 Ibs. N)

55f

20|

35*

36-6

In 1863 the winter and early spring were extremely mild, with a rain-
fall below the average. In the growing season of the early summer there
was abundant rain, followed by dry weather in June and July up to harvest.

420

THE WHEAT PLANT

In the season of 1878-9, from the 22nd of October to the first week
of December, the weather was cold and wet. The five months ending
March 31 were cold with much snow and rain, and from April to July
the days were cold and wet with little sunshine. Lawes states that it was
the worst season experienced in Great Britain since 1816.

Examination of the annual yields shows that even on the exhausted
plot which has carried wheat for sixty years without manure the impress
of a favourable season is very marked by a distinct increment, which in
1863 amounted to more than 4 bushels per acre. Perhaps more remark-
able is the fact that in the best season, plot 7 gave 53 bushels per acre,
while the application of the same amount of manure to this plot in 1879
resulted in the production of only 16 bushels.

Thus it is evident that season is of far greater import than any other
factor in determining yield : cultivation and manuring are of little avail
when the climatic conditions are unfavourable.

So far as this country is concerned the average annual temperature is
usually too low for the best development of the wheat plant, but reduced
yields of the grain are attributable to the high rainfall rather than to the
low temperature.

The relation between the average yields in England during the twenty-
one successive years 1884-1904, and the total rainfall for the three
autumnal months, October, November, and December, was investigated
by Shaw, who found that a close approximation to the actual returns at
harvest is given by the formula :

Yield per acre =39-5 bushels -f of the rainfall (expressed in inches)
of the previous autumn. In years of high yield the autumnal rainfall is
considerably below the average, in bad seasons it is excessive.

The importance of rainfall is seen in the following Rothamsted returns :

Average of Ten
Wettest Winters.

Average of Ten
Driest Winters.

Rainfall (November-February in-
clusive)

inches.
13-01

inches.

579

Average yield per acre
6, 7, and 8 .

of plots

bushels.
26-2

bushels.
34'9

Soil. — Although wheat will grow upon almost all kinds of soil from
the heaviest clays to the sands and gravels, the highest yields are only
obtained upon rich, deep, well-drained loamy clays, the physical character
of which is fairly uniform down to a depth of 2 or 3 feet.

Upon clays, especially if imperfectly drained, the yields are con-
stantly smaller, as they are also upon the light sands, gravels, and peaty

YIELD 421

soils, which require the application of much manure and careful cultiva-
tion before they will give remunerative returns of this cereal.

Cultivation and Manuring. — The influence of cultivation of the soil
and the manuring cannot be discussed here : they are nevertheless im-
portant factors in determining yield. Early ploughing in order to sow
on a stale furrow is a maxim followed by all farmers who obtain the best
yields of wheat.

With this cereal as with all other crops the application of suitable
manures improves the returns.

For the highest yields the soil should be rich in phosphates and potash
and well supplied with available nitrogen.

The two former elements are often obtained from the residues of
fertilisers used for the previous crops rather than from direct application
to the wheat plant, although the addition of superphosphates and kainit
may be profitably employed where the land is in poor condition. The
nitrogen is frequently obtained from the residue left by leguminous crops,
but the application of nitrate of soda or sulphate of ammonia to the plant
in spring is generally practised where heavy crops are obtained.

Amount of Seed. — The amount of seed sown has a large influence upon
the yield per acre, for upon it depends, in measure, the number of ears
which are produced, and it is these and the grains which they bear which
determine the yield.

In countries where the rainfall is low, the growing season short, and
the yield necessarily limited, the amount sown is often less than i bushel.
In Australia 40-60 Ibs. per acre is general, and in many parts of the
United States 45-60 Ibs. are often sown.

In Western Europe, with its higher rainfall, better cultivation, and
expected yield of 30 bushels or more per acre, very much larger quantities
are sown, the amount varying between 2 and 4 bushels (126-252 Ibs.)
per acre.

About 2 bushels per acre were recommended by the ancient Roman
agriculturists, Varro and Columella, and this quantity has been in general
use in Great Britain, apparently from the time of the Roman occupation
to the present day.

Since the seventeenth century it has been the practice to increase
the seed sown, 2 bushels being found insufficient in many instances to
provide the best number of plants or ears for a good crop of grain.

On land in a high state of fertility in a warm district, 2 bushels is
sufficient to produce a thick enough " plant " for the production of a
maximum crop. Where the soil is infertile, dry or cold, and in districts
in which adverse climatic conditions prevail, the amount is increased with
advantage up to 3 or 4 bushels per acre.

When fixing the amount of seed to sow, it is necessary also to take into

422 THE WHEAT PLANT

consideration the variety of wheat, the time and method of sowing, and
the average size of the seed.

Variety. — There are very considerable differences among wheats in
regard to their grain production under the same soil and climatic condi-
tions. This is chiefly due to the difference in the length of their growing
period. Many of the rapid-growing short-lived spring varieties complete
their life-history in 95-120 days. On the other hand, some of the winter
wheats require 180-200 days for their growth. The latter produce more
stems, leaves, and ears, and usually yield not less than 25 per cent more
grain than spring varieties on the same land.

Even among spring and winter varieties certain forms are found to be
more efficient than others when grown under the same conditions. For
example, it is generally believed that Marquis wheat produces more grain
than many other spring forms on most of the spring wheat-growing areas
of the American continent, and among winter wheats in Western Europe
the Squarehead types almost invariably give higher yields than any of the
lax-eared forms.

Time of Sowing. — There are two periods at which wheat is generally
sown, namely (i) autumn and (2) in spring. On account of the longer
growing period allowed by autumn sowing, the yield is invariably higher
for wheats sown at that time than after winter. The autumn-sown plants
are already established and possess several shoots before the later-sown
plants make a start, and this advantage is never lost.

The loss varies with the kind of wheat, the deficiency being least in
the case of the spring wheats, whose growing period is naturally short, and
greatest in the winter wheats, which require a long period for their highest
development.

Where the sowing of autumn wheats is delayed until February or
March there is generally a loss in yield of not less than 15-20 per
cent, and on land in poor condition it may amount to as much as 50
per cent.

At the College Farm, Reading, forty kinds were sown on October 21,
1910; and again on February 9, 1911. In all the varieties there was a loss
of yield of grain through sowing at the later date : the percentage de-
ficiency of a few showing the greatest and least variation are given below :

Loss per cent.

Red Dantzig ... 55

Chidham .... 49

Noe 49

Partridge .... 46

Red Nursery ... 44

Golden Drop 37

Swan . . . . 37

Loss per cent.

Sicilian .... 30

Hungarian ... 28

Japanese . . . . 26

Talavera .... 17

March .... 14

Fern .... n

Californian . 10

YIELD 423

The average yield of grain per ear of plants from seed sown in autumn
was i -45 gr., the ears of the spring-sown plants averaging -99 gr. of grain.

Many of the spring varieties tillered better when sown in February
than they did when sown in October.

Not only is there a difference between autumn and spring sowing, but
there is a progressive decrease in the yield with every week's delay in
sowing from October to December.

The advantage of early sowing is seen in the following results of trials
at the Harper Adams Agricultural College with Browick ( = Squarehead)
wheat.

Yield per Acre.

Time of Sowing.

Grain.

Straw.

bushels.

cwts.

September 30, 1916

34-00

43

October 13, 1916 ....

30-16

39

November 6, 1916

23-96

36

The results obtained from wheat (Swan) sown on the ist and
of each month throughout the year 1915 and first three months of 1916
are given on page 424 (Fig. 227).

The " tillering " or number of ears decreases more or less regularly
the later the grain is sown from May onwards to March of the following
year.

The average weight and length of the ears and the number of spikelets
are highest in plants obtained from grain sown in October and November ;
in earlier and later-sown plants these diminish.

The maximum average height of the straw is found in plants from
grain sown in September.

Although the number of ears per plant is greater when the grain is
sown before September, the ears are larger and contain a greater amount
of grain when the seed is sown in September, October, and November,
just those months in which it has been the practice of the farmers of this
country to sow their autumn wheats from the earliest times.

Which of the three months just mentioned gives the best returns in
particular cases depends upon a number of circumstances, such as the
physical and manurial state of the soil, aspect, altitude, and other con-
ditions. Whatever the time, it is important that the seed should be sown
in dry weather.

Generally speaking, the earlier the sowing the better the yield, especi-
ally in late districts and where the soil is comparatively poor. The

424

THE WHEAT PLANT

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YIELD 425

greatest area is sown in October, but in the warmer southern parts of
Great Britain, on well-drained soils in high condition, sowing in the first
half of November is often practised and gives good results. In such
districts if the grain is sown earlier the plants are liable to become " winter
proud," or too luxuriant in spring, a condition which frequently leads to
the " lodging " of the crop before harvest.

Methods of Sowing. — Three methods of sowing are in use, namely,
(i) broadcasting, (2) drilling, and (3) dibbling. The system of scattering
the seed broadcast by hand is the most ancient method, and is still prac-
tised wherever wheat is grown. It is a rapid method but involves the use
of more seed than the other methods.

Drilling in rows, first introduced by Jethro Tull towards the middle
of the eighteenth century, has now almost entirely superseded the broad-
cast system. Less seed is needed, and all the seed is deposited at a fairly
uniform depth in the soil when this method is adopted. Moreover, it
allows of the hoeing and cleaning of the crop during growth, and on this
account leads to a larger yield per acre than the broadcast system in which
the weeds remain unchecked.

Where the land is in a clean state and in a high state of fertility both
methods of sowing give similar results.

Dibbling, or depositing the seed by hand in holes made in the soil by
a blunt-pointed tool, had its advocates in former times. The holes were
made in rows 5-9 inches apart, and 3-5 inches asunder in the rows,
and two or three grains were dropped into each hole. Six to seven pecks
of seed were used, the work being carried out by the labourer and his
children.

Size of Grain. — The influence of the size and weight of the in-
dividual grains upon the size of the plants which they produce and
the amount of grain which these yield has often been the subject of
research.

The average weight of 100 grains of Bread Wheat varies considerably.
In some forms it is not more than 3-5 grams, with a maximum near 4-7
grams and a minimum of 1-6 grams ; others, especially those commonly
grown in this country at the present time, have an average weight of about
4-5 grams per 100 grains, with a maximum over 5 grams.

Peters in 1721 states that, in his day, in an ounce of good average wheat
there were about 700 grains, the best numbering 600, the smaller samples
800 per ounce : these figures correspond with a weight of 4-05 grams per
100 grains of average wheat, 4-72 grams for the best, and 3-54 grams for
the smaller grains.

In 1911 I selected ten large and ten small grains from a single ear of
several kinds of wheat, the average weight of each large grain being -05
gram, that of the small grains less than half or -025 gram. The average

426

THE WHEAT PLANT

yield of grain of the plants produced by these selected grains was obtained,
the experiment being repeated for five successive years.

The plants were grown in rows 6 inches apart and 3 inches asunder in
the row. The results are given below :

Ears

Average

Weight of

Average No. of

Average Yield of

per Plant.

Length of Ear
(cm.).

Grains per Ear
(grams).

Grains
in 10 Grams.

Grain per Plant
(grams).

From

From

From

From

From

Large.

Small.

Large.

Small.

Large.

Small.

Large.

Small.

Large. | Small.

PARTRIDGE WHEAT

1911-12

4-0

3-0

i*5 >

1-54 227

216

6-0

4-6

1912-13

12-3

II-O

3'44

3-11

148

190

42-5

34'7

1913-14

II-I

4'7

9-0

9-1

2-30

2-00

241

241

25-6

9-2

1914—15

10-5

9'4

10-2

10-2

2-02

1-72

275

297

21-2

16-2

1915-16

13-1

11-3

9'7

9-1

I-I3

260

279

I4-8

15-5

Average

10-2

7-9

9-6

9'5

2-08

i-9S

230

245

22-0

16-0

Percentage in- \

crease of large t

29'I

1-05

6-6

. .

37'S

. .

over small J

FOX WHEAT

1911-12

6-0

3-0

2-20

1-46

218

249

I3-2

4'4

1912-13

14-9

10-4

1-97

2-13

200

212

29-3

22-1

1913-14

10-2

7-6

9-7

9'3

1-85

1-70

255

265

I9-O

I3-I

1914-15

I2'I

8-3

n-7

11-3

1-63

1-96

270

265

I9-8

l6'3

1915-16

I3-I

n-8

10-4

10-1

1-07

•84

265

276

I4-O

9'9

Average

11-3

8-2

10-6

IO'2

1-74

1-62

242

253

I9-I

13-2

Percentage in- }

crease of large r

37-8

3'9

7-4

44'7

over small J

SHIRREFF'S BEARDED WHEAT

1911-12

9-0

s-o

1-78

2-18

200

191

16-0

I0'9

1912-13

10-8

9-0

1-74

1-82

218

189

18-7

16-4

1913-14

5'4

4-0

7'4

7'3

1-83

i-55

240

289

10-0

6-2

1914-15

9-2

5'4

9'9

9'4

1-64

1-15

324

355

15-2

6-2

1915-16

8-6

5-8

8-3

8-2

•73

•78

266

297

6-3

4-5

Average

8-6

5-8

8-5

8-3

i-54

1-50

250

264

13-2

8-8

Percentage in- ]

crease of large j-

48-3

2'4

2-7

50

over small 1

From these results it is seen that the average lengths of the ear and the
average weights of the grains per ear of plants produced from the large
grains are but slightly greater than those of the progeny of small grains,
and the difference may not be significant.

The number of ears per plant and the average yield per plant is, how-
ever, very much greater every year from the large seed than from the
small, and the difference in vigour was manifest throughout the whole
life of the plants.

YIELD

427

Zavitz carried out experiments with large and small wheat grains for
six years with winter wheats and eight years with spring wheats, and
found an average annual increase of 6-5 bushels per acre from large as
compared with small grains in the winter wheats, and 3-7 bushels increase
in the spring wheats.

Similar pre-eminence of large over small seed grain has been observed
by others.

The plants from small grain are weak, and many succumb to the
adverse soil and climatic conditions of autumn and winter.

In a few exceptional instances, where the land has been in a specially
high condition, small seed grain has given remunerative results almost
equal to those in which larger seed has been sown ; nevertheless the
inferiority of small grain as seed is always strikingly evident when its
yields are compared with those from large grain upon soil in compara-
tively poor condition.

Number of Grains sown.-^- The number of grains in a bushel (63 Ibs.)
of Bread Wheat varies between 500,000 and 800,000.

In the former case the grains are large, 100 weighing 5-71 grams, 100
of the latter weighing 3-57 grams. The number of grains in a bushel of
wheat of the type usually grown in this country is about 650,000, 100
weighing 4-4 grams.

From the following table it is seen that when one bushel of an average
sample is used there are sown about 134 grains per square yard, the rate
when 3 bushels are sown being 392 per square yard.

NUMBER OF GRAINS SOWN PER ACRE
(100 grains = 4-4 grams).

Bushels.

Number
per Acre.

Number
per Square Yard.

I

650,000

134

II

975,000

2OI

2

1,300,000

268

**

1,625,000

336

3

1,950,000

392

Even under favourable conditions the number of plants which appear
above ground after sowing these amounts of grain are generally 5-10 per
cent less than the figures quoted, since the germinating capacity of the
seed wheat is usually not more than 90-95 per cent.

Not infrequently 15-30 per cent of the plants which are seen soon
after the seed germinates disappear later. This decrease is due to de-
struction by birds, insects, and fungi, as well as by frost and adverse con-
ditions of the soil. Some of the loss is also brought about by competition

428 THE WHEAT PLANT

among the young plants and the deleterious effect which one plant has
upon another even of the same species when grown together. In Mont-
gomery's experiments the death-rate among wheat plants, grown in rows
8 inches apart and J inch asunder in the row, was 23-38 per cent, the loss
being 12-25 Per cent among the less crowded plants growing i| inches
asunder in the rows.

The number of plants which survive until harvest under ordinary
field culture has rarely been determined, but the number of ripe ear-
bearing straws has been frequently counted, the number per acre ranging
from about 150 to 400 per square yard, where 2^-3 bushels of grain have
been sown. In many instances there are fewer straws with ears than
grains sown, owing to the high death-rate among the plants.

It is found that a good crop is rarely obtained in this country where
the number of ears is less than about 300 per square yard.

Not only is the number of ears per acre significant, but the number and
weight of the grains in each ear is of the greatest importance, and efforts
to increase the yield per acre are usually directed towards improvement in
this respect.

The total yield of an ear is dependent upon the number of its spikelets
and the number of grains developed in each spikelet.

The average number of spikelets in an ear of Bread Wheat (T. vulgar e)
is about 20, but it differs, though not very widely, in different varieties and
forms, some having an average of 18, others 22 or 23 per ear.

In Rivet wheats (T. turgidum) the average is higher, being about 25.

The number of flowers in a spikelet varies from 3 to a possible 8 or 9,
but many of the upper flowers prove abortive, the number of grains which
develop varying from i to 7, the latter number being only met with in
the spikelets of ears borne upon plants which have been exceptionally
manured (Fig. 228).

Well-grown ears of the field often bear 5 or 6 grains in each of the 3
or 4 median spikelets, the numbers falling gradually to i or 2 in the basal
and apical spikelets.

Many of the spikelets at the base of smaller ears are abortive, the
number of these increasing with adverse climatic conditions, poverty of
the soil, and excessive tillering of plants sown earlier than September.

The largest ears of an ordinary field crop in this country frequently
possess 40 to 50 grains, weighing from 1-8 to 2-25 grams, the smallest
not more than 12 or 14 grains, weighing -54- -63 grams, an average ear
having from 25 to 30 grains, weighing 1-25 to 1-35 grams.

In the following table are given the calculated yields per acre, when the
number of straws varies between 50 and 400, and the yield per ear between
•5 gram (12 grains) and 3 grams (72 grains).

FIG. 228. — EARS OF " SWAN " WHEAT.

i. Normal ears. 2. Ears of plants excessively manured with nitrates, phosphates, and potash.

YIELD

TABLE OF YIELD PER ACRE IN BUSHELS (63 Ibs.) WITH VARYING
EAR WEIGHT AND NUMBER OF EARS PER SQUARE YARD

(100 grains = 4- 1 66 grams).

429

No. of

Yield per Ear.

Ears per
Square Yard.

•5 Gram

(12 grains).

i-o Gram
(24 grains).

1-5 Grams
(36 grains).

2-0 Grams
(48 grains).

3-0 Grams
(72 grains).

bushels.

bushels. bushels.

bushels.

bushels.

5°

4-23

8-46

12-69

16-92

25-38

IOO

8-46

16-92

25-38

33-84

50-76

15°

12-69 25-38

38-07

50-76

76-14

200

16-92

33-84

50-76

67-68

IOI-52

250

21-15

42-30

63-45

84-60

126-90

300

25-38

50-76

76-14

101-52

152-28

35°

29-61

59-22

88-83

118-44

177-66

400

33^4

67-68

101-52

J35'36

203-04

The amount of seed which should be sown in order to obtain the most
productive return has been settled by experience long ago in most of the
wheat-growing regions of the world.

In this country, as already mentioned, the amount lies between 2 and
4 bushels per acre.

In almost every generation, however, during the last two centuries
there have arisen thin-seeding advocates, who recommend the use of very
much smaller quantities of seed. Some of them refer to the sowing of
2^-3 bushels as " a barbarous system," and give evidence of records of
40 bushels per acre after sowing one peck (15-16 Ibs.) or less per acre in
drills 8-9 inches apart, the grain in the rows being on an average 4-5 inches
asunder.

From an examination of such cases it is seen that the land is especially
suited to the growth of wheat, and is in a high state of fertility and free
from weeds, conditions which lead to extensive tillering of the plants.

Experience has shown that on most of the wheat land in England
there is great risk in sowing less than 2 bushels per acre, and although
the success of these exceptional examples may lure the farmer to give thin-
seeding a trial, the practice is swiftly abandoned, for upon unsuitable soil
the experiment is a disastrous and total failure.

The extraordinary prolificacy of the wheat plant attracted attention in
early times.

Everard in 1692 (Houghton's Husbandry) states that he obtained from
single grains sown 10 inches apart, plants which produced 60-80 ears,
the largest of which contained 40-60 grains, the best plants yielding over
4000 grains.

Tull in 1731 refers to plants with 40 ears. In 1870 C. H. Shirreff

43°

THE WHEAT PLANT

found in his garden a single plant bearing 80 ears, which yielded 4524
grains, and I have had plants bearing 60-70 ears and over 2500 grains.

Many other examples might be given of the enormous number of
grains which can be obtained from the sowing of one.

In all these examples the plants were grown on soils in a high state of
fertility and had unrestricted space for their development.

The most important limiting factors in these high returns of single
plants is the space upon which they are grown, and the extent of the soil
through which the roots can freely penetrate without competition from
other plants.

These factors are frequently overlooked in discussing methods of
improving the yield per acre by the growth of exceptional plants.

The necessary space required to secure these highly productive plants
is not known, but it is certain that it is vastly greater than 10 square inches,
the space which is allotted to each grain when one bushel (650,000 grains)
is sown on an acre.

In the table below, from a single grain sown on the most widely
spaced plot 354 grains were produced, and from i bushel of similar grains
354 bushels would be obtained ; nevertheless, since each plant was allotted
4 square feet, to secure 354 bushels from such plants an area of nearly 60
acres would be needed. In other words, the yield is less than 6 bushels
per acre, although the statement that the return was 354 grains or bushels
for one sown appears exceptionally large.

Similarly the statements made by some thin-seeding enthusiasts that
they had experienced " a return of 500 for i on an acre," and " 804 for i
on a plot of 700 square feet," are no indications either of good or bad
yields per acre, for the statements apply equally to i grain, i pound, or
i bushel sown per acre or plot.

With increasing space the number of ears per plant and the number of
grains in each ear rises rapidly.

Below are given the results of the growth of plants of Swan wheat ( J1.
vulgare) produced from grains sown at varying distances apart, the smallest
space allotted to each plant being 6 square inches, the largest 576 square

inches.

TABLE I

Space
allotted.

No. of
Plants
per square
yard.

No. of Ears
per Plant.

No. of
Grains
per Ear.

No. of
Grains per
spikelet.

No. of
Grains per
Plant.

Weight of
Grains per
Ear.

Total
weight of
Grain per
Plant.

Bushels
(63 Ibs.)
per acre.

inches.

grams.

grams.

6xi

216

1-61

27-1

1-44

43

I-I4

1-18

43-15

6x3

72

1-52

32-6

1-69

50

1-48

2-25

27H3

6x6

36

1-82

34-6

1-66

63

1-58

2-87

17-49

12X6

18

2-92

43-3

1-89

126

1-88

5'49

l6'73

12 X 12

9

4-04

48-1

2-10

194

2-08

8-40

I 2-80

24 x 24

2-25

7-03

50-3

2-16

354

2-14

15-04

5-73

YIELD

Montgomery gives the following results of experiments upon the rate
of seeding and yield in an American winter wheat sown in rows 8 inches
apart and J inch, T, inch, i, and 2 inches asunder in the rows.

TABLE II

Area allotted to

No. of Plants

Yield of Grain

Total Yield

Yield

each Plant.

per Plot.

per Plant.

per Plot.

per Acre.

inches.

grams.

grams.

bushels (63 Ibs.).

8xJ

238

•91

259

45'3

8xi

217

1-36

159

41-2

8xi

204

2-58

79

38-7

8x2

I65

4-10

40

31-6

The relationship between tillering and yield per acre is of a compli-
cated nature. So far as the yield of the individual plant is concerned, the
more it tillers the greater the return in grain and straw. Moreover, when
sown at the normal time in autumn or spring the more highly tillered the
plants the greater the average weight of the ears, although when the tiller-
ing is excessive through sowing abnormally early the average weight of
the ears decreases. These conclusions are evident from the table above,
and from the results given on page 424 (Fig. 227).

The number of straws or ears and the average weight of the ears are
the factors controlling the yield per acre.

With thick sowing the number of ears per acre increases, but the
weight of the individual ears decreases. A thinly sown crop in which the
same number of straws are present as in one thickly sown gives a higher
yield on account of the fact that tillered plants have a higher average
weight of ear.

Nevertheless, if the number of grains is much less than that ordinarily
sown the increased number of straws per plant and the greater average
weight of each ear do not compensate for the loss of plants which thin
sowing entails.

Thus tillering may result either in an increase or a decrease in the
yield per acre when compared with a crop in which each plant has
produced but a single ear on account of being thickly sown.

While increased tillering leads to the production of more straw per
plant, the number of straws per acre decreases with tillering, a paradoxical
statement which depends for its truth upon the fact, that the smaller the
number of plants the greater the tillering ; at the same time this does not
compensate for the loss of plants incurred by thin-sowing.

In Table I. there is seen a rise of the yield per acre as the plants
become closer from 24 inches x 24 inches down to 6 inches x i inch.

432 THE WHKAT PLANT

In spite of the fact that under special circumstances thin-sowing may
succeed, in practice it is found to be less hazardous to attempt to obtain
an adequate number of ears per acre by thick-sowing rather than by thin-
sowing and its concomitant tillering, especially when the amount of seed
sown is far removed from that ordinarily sown.

Proportion of Grain to Straw and Proportion of Grain to Chaff. -The
ratio between the weight of the grain and that of the straw of the wheat
plant varies very much with the variety, the space allotted to the plants,
and their nutrition.

As ordinarily grown in the field the average proportion is usually
32-36 per cent of grain to 68-64 Per cent °f straw, but large amounts of
farmyard manure or nitrogenous fertilisers stimulate the production of
leaves and stems and reduce the proportion of grain to straw.

Plants grown at wide intervals show a different ratio.

Of seventeen common forms of Bread Wheat grown in rows i foot
apart, the plants 6 inches asunder in the rows, the highest proportion
was 47 per cent of grain to 53 per cent of straw, the lowest 33 of grain
to 67 of straw.

The proportion of the weight of the grain to that of the rest of the ear
(chaff and rachis) in a beardless wheat whose ears (500) were selected at
random from the field was 66-6 per cent of grain to 33-3 of chaff and
rachis, or 2 of grain to i of chaff.

In ears of plants grown 6 inches apart the proportion of grain was
higher, reaching in a number of cases a ratio of 75-78 per cent of grain to
25-22 per cent of chaff, i.e. between 3 or 4 of grain to i of chaff.

LIST OF WI1HATS

di. — /'. tltcoccunt.

du. ••• T. ifurum.

t. — T. turniilum.

v. «• T. vulgare,

iph. - /'. ipkatrocoecum.

o.^T. orientate.

po. " T. polonicutn.

py. — T. pyramidale.

c.—>T. compactutn.

Sp. - T. Spilta.

Country of

Country of

Name.

R

M . .111.1 Variety. Origin or where

Name.

K.I.I' .in.l V.nii ly. 0

r-ni "1 wlii-l''

chiefly grown.

chiefly grown.

Ablakh

v

Hostianum

IVrsia

Aurora

V. Hostianum

Canada

Abundance

T

ill/mi iihrum

Canada

Australian White

v. albidum

U.S.A.

Abyssinian

Azulejo

. du. obscurum

Spain

I'm ple = Tukur

Sim If

. .

. .

Badger . v. erythrospermum England

Admiral, Red

7'

lutescens

England

Baladi .

. du. leucurum

Asia Minor

Agh Bogda

V

graecum

IVrsia

Banat .

v. erythrospermum

Hungary

Akabozu .

V

lutescens

Japan

Bansi .

. du. apulicum

India

Aka yenidashi .

7!

. ferrugineum

,

t

Barbarusa .

v. ferrugineum

Spain

Ak Bashak .

t.

gentile

Turkey

Barbella

. v. erythrospermum

Portugal

tt

7)

ini'i iilininil,'

it

Barbu

a gros

Alaska

/.

pseudocervinum

U.S.A.

grain

. .

it

„

Fiani <•

Alrpli

V

lutescens

1 ranee

Barletta

. v. ferrugineum

Argentina

Alexandre .

du. africanum

Portugal

Baroota Wonder

v. albidum

Australia

Alfaro, de .

du afjine

Spain

Bascunano .

du. erythromelan

Spain

Alonso lanfanon

du. leucome.lan

n

,, . . du. alexandrinum

a

Altkirch rouge .

v

milturum

France

Bayah

. v. alborubrum

Australia

Amaiclla barba

Beech wood

Branca

tin. n If mi'

Portugal

Hybrid

v. milturum

U.S.A.

Am.uella barba

Beloglena .

v. erythrospermum

a

breta

du. melanopus „

BeloKlas .a /.aj-am

du. Valenciae

Bulgaria

Am her, Egyptian

v

erythrospermum

U.S.A.

Belokoloska

v. erythrospermum

Russia

„ Imperial

V

ferrugineum

M

„ . v. lutescens

it

,, Martin's

T

albidum

,, . du. hordeiforme

a

Ami-i n an Banner

1)

ullini nhrum

If

Belotourka . ,,

M

Amiiioiinier

d

France

Benefactor .

v. leucospermum

England

Analil .

du. melanopus

Portugal

Berbcrisco .

du. erythromelan

Spain

du. africanum ,,

Bianchette .

v. alborubrum

Italy

Andaluze .

du. melanopus

a

Bianco

. du. leucurum

it

Andriolo rosso .

t.

dinurum

Italy

Biancone .

t. gentile

,,

Anti-fly

t.

IIIK i ali'

England

Bishop

. v. albidum

( 'anaila

April Kern

v

.ferrugineum

tt

Blanc a paille

Arctic Junior .

V

.ferrugineum

U.S.A.

raide .

. v. albidum

France

Arnautka .

tin. Imrdeiforme

Russia

Blanc vcloutc .

v. leucospermum „

Arnovka

Blanco vcrdeal .

du. leucurum

Spain

Asa de Corvo .

du. lihycum

Portugal

Blanquillo .

n

tt

433

27

434

THE WHEAT PLANT

Country of

Country of

Name.

Race and Variety. Origin or where

Name.

Race and Variety. Origin or where

chiefly grown.

chiefly grown.

Blaue Dame

v. lutescens Germany

Challenge .

v. albidum England

Bl£ a dpi carre

Chamorro .

v . alborubrum Spain

= Squarehead .

France

Champion .

v. lutescens Sweden

Blood Red .

v. milturum Scotland

C h a m p 1 a i n

Blue Cone .

t. iodurum England

(Pringle's)

v. erythrospermum U.S.A.

Blue Ridge

v . ferrugineum U.S.A.

Champlan .

v. lutescens France

Blue Stem,

Chidham .

v. albidum England

Haynes

v. velutinum ,,

,, de Mars

,, France

Blue Stem,

Chile de fideos .

po. martinari Argentina

Palouse

v. albidum ,,

Chilian

v. albidum

Blue Velvet Chaff

v. cyanothrix Germany

»

c. Humboldtii

Bobs .

v. albidum Australia

Chul .

v. graecum U.S.A.

Bobs Red .

v. lutescens Canada

Claro .

du. leucurum Spain

Bomen

,, Australia

Clawson, White.

v. albidum U.S.A.

Bon Fermier

,, France

Clawson, Early

Bordeaux .

v. milturum ,,

Red .

v. milturum ,,

Bordier

v. albidum ,,

Clock .

t. dinurum England

Bore .

v. lutescens Sweden

Clover's Red

v. milturum ,,

Branco

t. lusitanicum Portugal

Clubhead .

v. albidum Australia

Briquet Jaune .

v. lutescens France

Club, Little

c. Humboldtii U.S.A.

Brodie's White

College Eclipse .

v. alborubrum Australia

= Oxford Prize

. .

Colorado .

du. erythromelan Spain

Browick

v. milturum England

Comeback .

v. albidum Australia

Browick White

Comet

S. Africa

— Squarehead .

• • »

Cone Rivet=

Buda Pesth

v. erythrospermum U.S.A.

Anti-fly .

Budd's Early

v. alborubrum Australia

Crepi, de .

v . lutescens France

Buisson

t. nigrobarbatum France

Criewener .

,, ,,

Bunge

v. albidum Australia

Crimean

v. erythrospermum U.S.A.

Bunyip

» »

Cumberland

v. albidum Australia

Burgoyne's Fife •

v. albidum England

Currawa

,, ,,

Burwell Red

v. milturum „

Currel's Prolific.

v . milturum U.S.A.

Bushak

v. Delfii India

Dantzig, Jersey •

v. albidum England

Caledon Baard .

v. erythrospermum S. Africa

Dantzig Red

v. alborubrum ,,

Caliph

v. albidum Australia

Darling

S. Africa

Canberra .

v. alborubrum ,,

Dart's Imperial .

v . albidum Australia

Candeal

v. graecum Spain

Daruma

v . ferrugineum Japan

Candial

t. lusitanicum Portugal

Dattel

v. alborubrum France

Cannu altu

du. leucomelan Spain

Dawson's Golden

Canoco

t. megalopolitanum Portugal

Chaff

,, Canada

Canuto

c. erinaceum Spain

Defiance

v. albidum U.S.A.

Carbillo

c. splendens Chili

,, Paine's .

t. dinurum Germany

Carman

v . ferrugineum

,, Pringle's

v. albidum U.S.A.

Carosello .

v. lutescens Italy

„ Webb's

v. lutescens England

Carosellone del

Deitz .

v. erythrospermum U.S.A.

Molise

t. melanatherum ,,

Del Frailte

v. erythroleucon Spain

Cascalvo

du. affine Portugal

Diamond Grit .

v. erythrospermum U.S.A.

Catanzaro duro .

du. leucurum Italy

Dickkop

v. albidum Holland

Cedar .

v. lutescens Australia

Dicklow

U.S.A.

Ceres .

v. albidum England

Dinkel

Sp.

Chacon

du. leucomelan Spain

Diozeg Banat

v. erythrospermum Hungary

LIST OF WHEATS

435

Country of

Country of

Name.

Race and Variety.

Origin or where

Name.

Race and Variety.

Origin or where

chiefly grown.

chiefly grown.

Dividenden

7;. milturum

Germany

Fern = April Fern

, .

England

Don, Black

du. obscurum

Field-Marshal .

v. albidum

Australia

„ Velvet

du. africanum

Fife, Bearded .

v. erythrospermum U.S.A.

Downy Kent

v. leucospermum

England

Fife, Bearded

Dreadnought^

Winter

v. meridionale

,,

Bon Fermier •

Fife, Burgoyne's

v. albidum

England

Duckbill .

t. iodurum

England

„ Crail .

v. velutinum

U.S.A.

Dugdale

,,

,,

Fife, Jones'

Durazio mollar .

du. melanopus

Portugal

Winter

,,

,,

,, rejo

,,

,,

Fife, Power's

v. milturum

,,

. Du Toit .

v. albidum

S. Africa

„ Red.

v. lutescens

Canada

„ White

v. albidum

• "

Early Baart

v. graecum

U.S.A.

Filipino

t. mirabile

Spain

„ Ripe .

v. milturum

Firbank

v. albidum

Australia

Ecksteen

v. graecum

S. Africa

Flandres, Blanc.

,,

France

Eclipse = Red

Florence

„

Australia

Admiral .

Focense

v. ferrugineum

Spain

Eggshell

v. albidum

England

»

t. dinurum

,,

Egypcio

v. erythrospermum Azores

Forte nero .

t. iodurum

Italy

Egyptian Cone .

pyr.

Egypt

Fortyfold .

v. alborubrum

Canada

Spelt .

di tricoccum

,,

Fourie

v. graecum

S. Africa

Ein el Bent

pyr. copticum

,,

Fox

v . ferrugineum

Germany

Eley's Giant

v. albidum

England

,, . . .

v. pyrothrix

England

Emmer

di.

Frankenstein

v. albidum

Germany

„ Black .

di. atratum

Germany

Frenisburg

v. lutescens

Switzerland

„ Red

di. rufum

,,

Fretes

v. erythrospermum U.S.A.

„ Slav

di.farrum

Russia

Fucense

v. ferrugineum

Italy

„ White .

,,

Germany

Fultz .

v. lutescens

U.S.A.

Emperor

v. albidum

England

Enano de Lorca

du. melanopus

Spain

Galician

v. lutescens

,, fuerte

.

'

Galland

t. lusitanicum

France

Epautre

Sp.

France

Gallego barbado

v . ferrugineum

Portugal

Epp .

v. albidum

Germany

„ rapado .

v. milturum

,,

Essex Conqueror

Gallego rapado

= Squarehead .

England

branca

v. lutescens

,,

Essex Hybrid .

v. albidum

,,

Garagnon, Lan-

Essex Velvet

guedoc

t. melanatherum

France

Chaff .

v. leucospermum

t

Garagnon, noir .

t. Herrarae

,,

Essex White

v. albidum

Australia

Garnovka .

du. hordeiforme

Russia

Gatinais, White

t. gentile

France

'Fanfarron .

du. leucurum

Spain

Red .

t. dinurum

„

,, bianco

du . fastuosum

tl

Genesee Giant

(Fanfarron,villoso

Early

v. erythroleucon

U.S.A.

t raspinegro

du. africanum

,,

Genoa .

v. albidum

Australia

Fanfarron, villoso

Gentile Bianco .

,,

Italy

'. rubion

du. italicum

Ghirka, Red

v. milturum

Russia

Farmer's Friend

v. albidum

Australia

„ White .

v. lutescens

M

,, Trust .

v . ferrugineum

U.S.A.

Gironde

v. lutescens

France

kFederation

v. alborubrum

Australia

Gluyas

v. alborubrum

Australia

,, hard .

,, Bearded.

v. erythroleucon

,,

[Penman

v. lutescens

England

Gneisendorf

v. lutescens

Austria

IFenton

v. albidum

M

Goldcoin .

v. alborubrum

U.S.A.

436

THE WHEAT PLANT

Country of

Country of

Name.

Race and Variety. Origin or where

Name.

Race and Variety. Origin or where

chiefly grown.

chiefly grown.

Golden Aue

v. velutinum

Hunter's White.

v. albidum England

„ Ball

du. hordeiforme

S. Africa

Huron

v . ferrugineum Canada

,, Cross

v . ferrugineum

U.S.A.

„ Drop Red

v. milturum

England

Imperial Buff .

v. velutinum England

Golden Drop

,, Street's

v. lutescens ,,

White

v. lutescens

Invernanca

v . ferrugineum Spain

Golden Gem

v. Delfii

N. Zealand

Inversable rouge

Goose

du. melanopus

Canada

= Bordeaux

France

du. Valencia

Iron .

v. lutescens Sweden

Goose, Black =

Ironclad

v. erythrospermum U.S.A.

Nicaragua Black

Ismael

du. Valencia France

Goose, Wild .

du. leucurum

S. Africa

Jade .

v. albidum Australia

Granadino •

du. melanopus

Spain

Japhet

v . lutescens France

Granella de Car-

„ White .

v. albidum S. Africa

pegna
Grano foote

v. lutescens
du. Reichenbachii

Italy

Javardo
John Brown

du. libycum Portugal
v. alborubrum Australia.

„ nero

t. iodurum

,,

Jonathan . .

v. albidum ,,

Grenadier .
Grimbeck's Klein.

v. lutescens

Sweden

Jumbuk

v. leucospermum „

Koren

v. erythrospermum

S. Africa

Kadiz Oglou

v. erythroleucon Turkey

Gros bleu .

v. lutescens

France

Kalkori

v. Delfii Persia

Grosse t e1 te,

Kansas Mortgage

Hybride .

„

,,

Lifter

v. erythrospermum U.S.A.

Guisano

v . ferrugineum

Spain

Kara Bashak

du. coerulescens Asia Min

Gypsy

v. erythrospermum

U.S.A.

Kar-i-safid

v. turcicum Persia

Kent, Red .

v. milturum England

Hadmersleben .

v. lutescens

Germany

Kessingland

v. lutescens ,,

Haie, de = Tun-

Keupely

t. pseudocervinum Turkey

stall .

France

Khapli

di. Arras India

Hallett's Pedigree

King's Red

v. erythrospermum Australia

White

v. albidum

England

„ White .

v. graecum „

Hardcastle .

v. milturum

U.S.A.

King William =

Helena Hybrid .

v . ferrugineum

Hickling .

England

Hembrilla .

„

Spain

Kintana

v. erythrospermum Japan

H6risson, Bearded

c. icterinum

France

Kinver Red

v. lutescens England

Herisson, Beard-

Kizildaly .

v. alborubrum Turkey

less .

c. creticum

„

Koffoid

U.S.A.

Hickling

v. lutescens

England

Kostroma .

v. albidum Russia

Hickman .

„

U.S.A.

Krasnokoloska .

v . ferrugineum „

Hindi .

v. graecum

Egypt

Kubanka .

du. hordeiforme Russia

Hoary, Old

v. leucospermum

England

Kubb .

c. Wernerianum Sweden

Hochfeldt .

v. milturum

Switzerland

„ Velvet

c. Wittmackianum ,,

Hogan, White .

v. albidum

Australia

KwangT'ouMai

v. milturum China

Hongrie Rouge

v. milturum

France

Hopetoun .

v. albidum

England

Ladoga

v. ferrugineum U.S.A.

Hosagara .

v . ferrugineum

Japan

Lamed

v. milturum France

Hsu Hsu Mai .

China

Lammas, Red .

,, England

Hudavendigar .

t. dinurum

Turkey

White

v. albidum ,,

Huerta

v . ferrugineum

Spain

Lancaster, Red .

v. erythrospermum U.S.A.

Huguenot .

du. australe

Australia

Leap Prolific

v. lutescens ,,

Hundredfold

v. milturum

England

Little Club

c. Humboldtii England

Hungarian .

„

Argentina

,, Joss .

v. milturum ,,

LIST OF WHEATS

437

Country of

Country of

Name.

Race and Variety. Origin or where

Name.

Race and Variety. Origin or where

chiefly grown.

chiefly grown.

Lobeiro

du. affine

Portugal

Model Red

v. lutescens

England

Lofthouse .

v. lutescens

U.S.A.

Mogul

v. alborubrum

Canada

Loosdorf , Bearded

v . ferrugineum

Austria

Mold's Red Pro-

Loosdorf, Beard-

lific .

v. milturum

Germany

less .

v. lutescens

,,

Monarch = Wil-

Lucerna, de

v . ferrugineum

Spain

helmina .

England

Lulea .

v. velutinum

Sweden

Mongia

du. niloticum

Portugal

Montjuich .

v . ferrugineum

Spain

Macolo

du. leucomelan

Spain

Moro .

v . ferrugineum

,,

»

du. melanopus

,,

Moula Oglou

t. Mertensii

Turkey

Madonna .

du. libycum

Germany

»

t. Dreischianum

. .

Mainstay .

v. Delfii

England

Mourisco ver-

Majestic

v. albidum

Australia

melho

du. erythromelan

Portugal

Major

,,

,,

Mullybrack

v. albidum

England

Malakov

v. erythrospermum

U.S.A.

Mummy

t. mirabile

Mammoth Red .

,,

,,

Mundi Pissi

v. albidum

India

Manchester

v. albidum

Mungoswells

„

Scotland

March

,,

Murwarid .

v. alborubrum

Persia

,, Bearded •

v. erythrospermum

Muzaffarnagarh

v. graecum

India

Mareuil, de

v. albidum

France

Myburgh .

v. erythrospermum

S. Africa

Mark Lane

v. albidum

England

Marouani .

du. erythromelan

Nach Sinde

di. tomentosum

Abyssinia

Marquis

v. lutescens

Canada

Nanchero .

v. milturum

India

Mars rouge, barbu
Mars rouge, sans

v. ferrugineum

France

Naples White .
Negro de Camelas

v. albidum
du. apulicum

Spain

barbes

v. milturifm

New Zealand

Marshall's No. 3

v. albidum

Australia

Long

v . ferrugineum

N. Zealand

Martin

v. milturum

England

Nicaragua Black

du. obscurum

S. Africa

Marvel, Red =

Niekirks

v . ferrugineum

»

Bon Fermier .

Nigger

v. erythrospermum

U.S.A.

Marvel, White .

v. albidum

Canada

Nobbs

v. albidum

S. Africa

Marzuolo .

v . ferrugineum
du. Reichenbachii

Italy

Noe

v. alborubrum
v. lutescens

France

Massengo .
Massy •

v. erythrospermum
v. lutescens

Italy
France

Nonette de
Lausanne

t. dinurum

»

Matador Rood

Nonpareil .

v. alborubrum

England

Kop .

v. alborubrum

Holland

Nursery Red

v. milturum

>»

Matador Wit Kop

v. albidum

,,

Oakley, Extra

Mazzochio

t. speciosum

Italy

Early

v. lutescens

U.S.A.

Medeah

du. erythromelan

Spain

Obispado .

du. affine

Spain

Medeah Bald =

Odessa

v. alborubrum

||

Huguenot

Australia

v. albidum

Mediterranean .

v . ferrugineum

U.S.A.

Onega

v . ferrugineum

Canada

Michigan Amber

v. milturum

,,

Onigera

U.S.A.

,, Bronze

v . ferrugineum

,,

Ontario, Early .

v. albidum

»

Milanaise, Geante

,, Wonder

v. lutescens

de .

t. dinurum

France

Oregon Red

n

n

Million

v. albidum

Holland

Ou-Baard .

v. graecum

S. Africa

Minnesota 169 .

v. velutinum

U.S.A.

Oxford Prize

v. milturum

England

Minuto d'Albona

v. ferrugineum

Italy

Miracle

t. mirabile

Partridge .

v. lutescens

England

Mocho

c. creticum

Madeira

Pearl .

v. leucospermum

N. Zealand

438

THE WHEAT PLANT

Country of

Country of

Name. Race and Variety. Origin or where

Name. Race and Variety. Origin or where

chiefly grown.

chiefly grown.

Pellisier . du. melanopus S. Africa

Raspinegro espiga

Pendulum = Rivet .. England

negra . du. libycum Spain

Penny . . v. albidum Australia

Raspinegra espiga

Pera, La . . v . ferrugineum Spain

pequena . du. leucomelan ,,

Perle de Nuise-

Ratel . . t. dinurum U.S.A.

ment . v. erythroleucon France

Realforte . du. leucomelan Sicily

Persian Black . di. persicum

Recio de Granada du. melanopus Spain

Petianelle blanche t. lusitanicum France

Red Wave . v. milturum U.S.A.

Pe'tianelle noire

Regadio . . v . lutescens Spain

de Nice . t. iodurum ,,

Rentpayer . ,, England

Peton v. alborubrum Canary Isles

Reynold's Dis-

Phillipino . . t. nigrobarbatum U.S.A.

covery . t . iodurum Australia

Phillipolis . v. albidum S. Africa

Ribeiro . . v. erythrospermum Portugal

Pilbeam . . v. milturum England

Richelle de Naples v. albidum France

Pinet . . . du. erythromelan Spain

Rieti . v. erythrospermum Italy

Pirkey . . v. lutescens England

Riet Koorn = Rieti .. S.Africa

Pisanablanca . t. gentile Spain

Rivet . . t . dinurum England

,, francesca. t. iodurum ,,

Rochester Red . v. milturum U.S.A.

Pithiviers . v. milturum France

Roermaker . v. graecum

Plover . v. lutescens England

Romanella . . v . ferrugineum Italy

Pole Rivet . t. iodurum ,,

Rooi Klein Koren ,, S. Africa

Pombinho . t. buccale Portugal

Rooi Wol Koren v. Delfii ,,

Poole . . v. milturum U.S.A.

Roseau . v. albidum France

Footing Bearded v . ferrugineum ,,

Roseworthy . ,, Australia

Portuguez . du. leucurum Portugal

Rosso gentile

Potschefstroom . v. albidum S. Africa

aristata . . v. erythrospermum Italy

Poulard blanc . t. gentile France

Rosso gentile

Poulard

mutico . v. milturum ,,

d'Australie . t. iodurum ,,

Rouge barbu

Poulard de

d'automne . v . ferrugineum France

Gatinais . . t. speciosum ,,

Rouge barbu de

Poulard a six rangs t. gentile ,,

Mars . . ,, ,,

Preston . . v. erythrospermum Canada

Rouge d'Ecosse . v. milturum ,,

Prince Albert . v. milturum France

,, de Mars . ,, ,,

Pringle's Champ-

Rouge prolifique

lain v. erythrospermum U.S.A.

barbu . . v . ferrugineum ,,

Prizetaker . . v. alborubrum Canada

Roumanian . ,, Roumania

Probsteier . v. lutescens Germany

Rousselin . . v. alborubrum

Prolifique barbu v . ferrugineum France

Rubiao . du. hordeiforme Portugal

Propo . . v. erythrospermum U.S.A.

Rudy . . . v. erythrospermum U.S.A.

Prosperity . v. lutescens ,,

Ruio . . du. hordeiforme Spain

Provence, rouge

Rural New Yorker v.Hostianum U.S.A.

de . v. milturum France

Russian Red . v . ferrugineum ,,

Pudel . . v. leucospermum Sweden

Rymer . . v. albidum Australia

Purple Straw . v. albidum Australia

,, . v. lutescens U.S.A.

Saidi . . . py. recognitum Egypt

St. Helena, Giant t. dinurum England

Queen, Fan . v. albidum Australia

St. Laud Rouge • v. milturum France

,, White . ,, England

Salamanca . t. lusitanicum Spain

Salmeron . . t . pseudomirabile ,,

Rapado quadrata c. creticum Madeira

Sammet . v . velutinum Sweden

Raspinegro . du. Reichenbachii Spain

Samogyer Tar . v. lutescens Hungary

LIST OF WHEATS

439

Country of

Country of

Name.

Race and Variety. Origin or where

Name.

Race and Variety. Origin or where

chiefly grown.

chiefly grown

Sandomir .

v. alborubrum

Russia

Suede rouge de

Santa Marta

du. mur dense

Portugal

Mars

v . ferrugineum

France

Saragolla .

du. leucurum

Italy

Sun

v. lutescens

Sweden

Sari Bashak

du. hordeiforme

Asia Minor

Sunagawa .

v . ferrugineum

Japan

Saumu r

Superlative

v. alborubrum

Canada

d' Auto nine

v. lutescens

France

Swan .

v. lutescens

England

Saumur de Mars

j

;)

Saxonka

v. erythrospermum

Russia

Taganrock

t. gentile

Schilf .

v. albidum

Germany

Talavera

v. albidum

England

Schlanstedt Red

v. milturum

,,

Talavera Belle1 Vue

,,

>»

Schneider .

v. albidum

Australia

Tarragon .

,,

Australia

Schdnrader

v. velutinum

Sweden

Tasmanian Red .

v. ferrugineum

Scotch Blood

Taunton Dean .

v. lutescens

England

Red .

v. milturum

Scotland

Temporas de

Seccano

v. erythrospermum

Spain

Coruche .

v . ferrugineum

Portugal

Seigle .

v. pyrothrix

Germany

Teverson .

v. milturum

England

Seneca Chief

v. graecum

U.S.A.

Theiss = Banat .

Hungary

Seven-headed

t. pseudocervinum

,,

Thew

v. albidum

Australia

Shirokawa Soshu

v. erythrospermum

Japan

Thickset Suffolk

Shirokawa

= Hickling

. .

England

Kanagawa

v. lutescens

»

Tom Thumb

v. albidum

N. Zealand

Shirokawa

Touzelle anone .

v. lutescens

France

Yenidashi

v. erythrospermum

,,

Touzelle de Pro-

Shirreff's White .

v. graecum

England

vence

v. milturum

,,

Shutar Dandan .

o. insigne

Khorasan

Touzelle Velvet .

v. leucospermum

,,

Siah Das .

o. notabile

,

Treasure White.

v. albidum

N. Zealand

Sibiey's Golden.

v . ferrugineum

U.S.A.

Tresor

v. lutescens

France

Sicilian

c. creticum

Trimenia .

du. affine

Italy

Siciliani

du. Valenciae

Italy

Triumph

v. albidum

Australia

.»

du. italicum

,,

Trump

v. lutescens

Sicilio

t. buccale

» •

v. albidum

Siebritz

v. erythroleucon

S. Africa

Tiichtiger Land-

Silver Dollar

v. erythrospermum

U.S.A.

wirth

v. lutescens

Germany

,, King

v. velutinum

,,

Tukur Sinde .

di. Arraseita

Abyssinia

„ Sheaf .

v. erythrospermum

u

„ White .

M

Smyrna = Miracle

Tunis .

py. pseudo-com-

Egypt

Snowdrop

v. lutescens

England

pressum

»

v. leucospermum

N. Zealand

Tunstall .

v. leucospermum

Solina d'Ascoli .

v . ferrugineum

Spain

Turkey

t. iodurum

France

Sonora

v. Delfii

U.S.A.

Red

v. erythrospermum

U.S.A.

Sorrentino

du. affine

Germany

—Tuscan Red

v. lutescens

N. Zealand

Spalding's Prolific

v. milturum

England

White .

v. albidum

,,

Squarehead

v. lutescens

,,

Tuscan Solid

Squarehead's

Straw

„

»

Master

v. milturum

)(

Tusela

v. lutescens

Argentina

Standard Red .

,,

M

Tystofte Smaa-

Standup, Main's

v. albidum

,,

hvete

»

Denmark

Red .

v. milturum

,,

Tystofte Stand-

White

v. albidum

,,

hvete

»

»>

Stanley

v. milturum

Canada

Starling

v. albidum

England

Ulka .

v. erythrospermum

Russia

Steinwedel

»

Australia

»

v . ferrugineum

»

440

THE WHEAT PLANT

Country of

Country of

Name.

Race and Variety. Origin or where

Name.

Race and Variety.

Origin or where

chiefly grown.

chiefly grown.

Ulka .

v. lutescens Russia

Wards Prolific .

v. alborubrum

Australia

,, . . .

v. milturum ,,

Warren

v. albidum

Upper Cut .

v. albidum Australia

Wave, Red

v. milturum

U.S.A.

Urtoba

„ Germany

White Straw Red

v. lutescens

England

Usher's Red

v. erythrospermum Australia

Whittington's

Utendorf .

v. milturum Switzerland

White

v. albidum

(>

Wilhelmina

M

Holland

Valley

v. erythrospermum U.S.A.

Willem I. .

,,

11

Veloute", Blanc .

v. leucospermum France

Windsor Early .

v. alborubrum

Canada

Velvet, Blue

v. cyanothrix Persia

Windsor Forest .

v. lutescens

England

Chaff .

v. leucospermum England

Wit Klein Koren

v. graecum

S.Africa

„ Red .

v. barbarossa

WitWol Koren.

v. leucospermum

,,

Verdeal

du. leucurum Spain

Wonder, Red

v. erythrospermum U.S.A.

Vermelejoilo

du. hordeiforme Portugal

Vermelho fino .

du. aegypticum „

Xeixa .

v. alborubrum

Spain

Victor

v. albidum England

Xeixa invernanca

v . ferrugineum

,,

Victoria de Mars

v. erythrospermum France

Xeres .

du. leucurum

France

Red .

v. lutescens ,,

White .

v. albidum England

Yandilla King .

v. albidum

Australia

Victris

!> • •

Yaroslav

v . ferrugineum

U.S.A.

Vroe Baard =

Yeoman

v. lutescens

England

Ecksteen .

Zeeland

v. albidum

Wallace

' v. albidum Australia

Zimmerman

v. lutescens

U.S.A.

Warden

v. lutescens ,,

Zwaart Baard

du. melanopus

S. Africa

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BIBRA, 1861. Die Getreidearten und das Brod. Histor. Skizze. and ed. Niirnberg.
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2 G

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INDEX

Aaronsohn, 167, 178, 183, 184, 377

Ablakh, 275, 280

Abundance, 299

Abyssinian Emmer, 193 (2, Fig. 129)

Purple-grained Emmer (Fig. 130)
Aegilops, 152

Crithodium, 164

cylindrica, 339, 342-346, 385 (Figs. 213,
214)

ovata, 336, 342-345, 380, 381, 383, 385
(Fig. 223) ,

speltaeformis , 383 (Fig. 223)

triaristata, 345. 385

triticoides, 380-383 (Fig. 222, 223)

triuncialis , 385

ventricosa, 385
x wheat, 380-386
x wild wheats, 377
Age of grain and germination, 33, 34
Agga, 193

Agh Bogda, 275, 279
Agropyrum repens, 380
Ajar, 193
Ak Bashak, 250
Ak Poussan, 213
Aka yenidashi, 284
Akerman, 356

Alaska wheat, 253 (i, Fig. 160)
Alefeld, 151
Aleuron grains, 1 1

layer, 7, 31, 130 (Figs. 3, 7, 8)
Alexandre, 220
Alexandretta, 213
Amar, 186

Amarella de barba branca, 215
Amelkorn, 186
Amer, 186
Amidonnier, 186
Amos, A., 418
Anafil, 220

Ancient wheat grains, 32-33, 187 (Fig. 124)
Andriolo rosso pelosa, 258
Anther, in (Figs. 84-87, 96-98)

dehiscence, 123
Anthesis, 122-128 (Fig. 102)
Antipodal cells, 122, 132 (Figs. 101, 105)
Arab boyde, 228
Archesporial cells, 116 (Fig. 96)
Aristobulus, 335
Aristotle, 171

Arnautka, 222
Arnovka, 222
Arraseita, 195
Arsus, 213
Ascherson, 153
Astrakhan wheat, 231
Auricles, 48, 61 (Fig. 42)
Awns, 104, 359

anatomy of, 108 (Figs. 82, 83)

colour of, 105

inheritance of, 365-368
Azima, 285, 292

Backhouse, 387, 395, 397

Badger, 278

Balansa, 167

Bald Medeah, 229

Bally, 119, 383

Banat, 278

Bansi, 341

Barletta, 283

Barley, Nepal, 386

x wheat, 386

Bauhin, 147, 148, 230, 307
Bayle-Barelle, 148
Bearded vulgare wheats, 273, 274

wheats, 104, 105
Beardless vulgare wheats, 273, 288

wheats, 104, 105
Beijerinck, 378, 391
Beloklassa zagani, 217
Berosus, 335
Bertoloni, 380
Bianchetta, 299
Biancone, 249
Bieberstein, 164
Bielotourka, 222
Biffen, 376, 392, 397
Binkel, 308
Blanc a duvet, 297
Blanc a paille raide, 292
Blanc de Flandres, 291
Blaringhem, 354, 391
Ble a duvet, 305
Ble Garagnon, 251, 259
Ble Hybride Galland, 248
Blue Cone, 260

Stem, Haynes', 298

Velvet Chaff, 306
Bobart, 231

•155

456

THE WHEAT PLANT

Bobs, 290, 386

Bon Fermier, 294

Bordeaux, 301

Bordier, Hybrid, 291

Bote, 187

Bourgeau, 167

Bread wheat, 157, 265-306, 341

Breeding of wheat, 407-414

Brenchley, 133

Browick, 303

Greychaff, 296
Brown and Escombe, 19, 31
Bruns, 17

Brush, 7, 8 (Fig. 2)
Buca vera, 255
Bud sports, 356
Budge, 187
Buffum, 355, 358, 380
Bulliform cells, 54 (Fig. 56)
Burgerstein, 34
Buschan, 266, 307

Candolle, A. de, 241

Caporn, 374, 395

Carman, 285, 388

Caron-Eldingen, 359

Carosellone del molise, 251, 252

Carpel, development of, 115 (Figs. 94, 95)

Carruthers, 34

Caryopsis, 7

Cawnpore, No. 2, 218

No. 9, 218

No. 18, 226
Cedar, 294
Celakovsky, 17
Cherler, 230
Chidham, 290
Chili velu, 317
Chlorophyll, layer, 8
Chromosomes, 117-119
Classification, 147-163
Clemente, 149
Clock wheat, 258
Club wheats, 157, 307-320, 346
Clubbed ear, 101 (Figs. 184, 189)
Cluster wheats, 307-320
Cobb, 414
Coleoptile, 14, 17, 44 (Figs. 10, 35)

anatomy of, 5 1

vascular bundles of, 15, 16 (Figs. 15, 61)
Coleorhiza, 14 (Fig. 10)
Colour of awns, 105

of glumes, 102

of grain, 17, 137
Columella, 147, 148, 408
Columnar epithelium, 13, 29, 30 (Figs. 10,

n, 14, 19)
Comeback, 290
Cone, Blue, 260

Egyptian, 262-264

Grey, 261

Long, 261

Red, 259, 261

wheats, 156, 241-261

White, 261
Conqueror, Essex, 295

Cook, 170, 184, 377

Correlations, 349

Cotyledonary sheath, 13, 135 (Fig. 10)

Couch grass, 380

Crease of grain, 7

Crescentius, 325

Criewener Squarehead, 296

Red, 303

Crithodium aeqilopoides , 164
Crossing, multiple, 412
Cross-layer, 8, 9, 138 (Figs. 3, 4)
Cross-pollination, 129
Crystals, 57 (Fig. 58)
Culms, 62

anatomy of, 92-99 (Figs. 70-74)

diameter, 64

internodes of, 63

lengths, 63
Cytase, 31
Cytohydrolysis, 31

Darling, 299

Daruma, 284

Dattel, 300

Dawson's Golden Chaff, 300

Dead-ripe, 141

De Alfaro, 215

De Haie, 298

Deina, 151

polonica, 230
Deininger, 265, 266
Del Frailte, 282
Delisle, 263
Density of ears, 160

inheritance, 365
Desfontaines, 207
Dikkop No. i, 291
Dinkel, 158, 325-344

Schlegel, 333

Tyrol, 334

Vogeles, 333
Diocletian, 325
Diodorus siculus, 335
Dioscorides, 149

Disarticulation of ear, 100 (Fig. 77)
Discontinuous variations, 352
Dodoens, 207, 241
Dreadnought, 294
Duckbill wheat, 261
Dudley, 118
Dugdale wheat, 261
Durazio mollar, 219

rijo, 219

Duro Catanzaro, 213
Duthie, 321
Duval-Jouve, 54
Dwarf cells, 107, 108, 109 (Fig. 81)

wheats, 307-320

Indian, 157, 321-324
Dwarfs, 355

Ear, 100
Ear-density, 160
Earing of wheats, 84-92
Earliness, 161
Early wheats, 85

INDEX

457

Ears, branched, 358

malformed, 358

number per plant, 73, 77

short, 358

twin, 358
Ecksteen, 275
Egyptian Cone wheat, 156, 262-264, 341

grain, ancient, 32, 33, 187 (Fig. 124)
Ein el Bent, 264
Einkorn, 154
Embryo, 7, 12 (Figs. 9, 10)

development of, 133-136 (Figs. 106, 107)
Embryo-sac, 1 14, 120 (Figs. 91, 101, 105)
Emerum, 186
Emmer, 155, 186-203, 338

Abyssinian, 11.3

Black, 202

European, 192, 196-203

Indian, 194

Indo-Abyssinian, 192, 193-196

Large White, 197

Red, 200

Slav, 198

White, 197

Wild, 154, 178-185
Emperor, 291
Empty glumes, 101, 161
Endodermis, 39, 40 (Figs. 26, 27, 30, 33)
Endosperm, 10

development of, 130, 132

inheritance, 375
Endothecium, 112
Engledow, 371, 396
English wheat, 242
Engrain, 154

double, 176

Epiblast, 13, 17 (Figs. 9, 10)
Epithelium of scutellum, 13 (Figs. 10, u,

14)

Epp, 289

Erect habit, 69, 162 (Fig. 65)
Erysiphe graminis, 81
Essex Conqueror, 295

Hybrid, 291

Rough Chaff, 297

Squarehead, 296

Fabre, 340, 381-384
Far, 186, 187, 325
Farrer, 355, 386, 412
Farro, 325
Fayum wheat, 264
Federation, Red, 299
Fellasito, 231
Fenton, 290
Fertilisation, 129
Fife, Red, 293
Flaksberger, 231, 337
Flandres, Blanc de, 291
Flinty grains, 18-22
Flower, 110-143 (Fig. 84)
Flowering, 122-128 (Fig. 102)
Flowering-glumes, 104
Fluctuations, 347
Foliage-leaves, 44-51 (Figs. 36-42)
anatomy of, 53-61 (Figs. 43-57)

Form, 159
Forte nero, 260
Fortyfold, 299
Fox, 305

Frankenstein, 294
French, 301
Frenisburg, 293
Frontal tuft, 100
Frost, resistance to, 162
Fuchs, 241
Fuller, 321
Fusion-nuclei (Fig. 100)

Gaertner, 12

Galen, 149

Gallego rapado, 301

Gametes, male, 119 (Figs. 103, 104)

Gandum-i-Kaiseh, 289, 293

Gandum Kamisheh, 297

Garton, 412

Geldersche, 294

Gentile bianco, 289

Germination of the grain, 24-34

chemical changes, 28-30

of pollen grain, 129 (Fig. 103)

of Spelt grains, 26

old grain, 33, 34

temperatures, 27, 28
Giant Rye, 231

St. Helena, 258
Gigachilon polonicum, 230
Gigante milanese, 258
Gitkova, 124

Glabrous leaves, inheritance of, 364
Gland of scutellum, 13 (Fig. 12)
Glaser, 188
Glume, anatomy of, 107

colour of, 102, 103, 369

empty, 101

flowering, 104

glabrous, 103, 371

hairs on, 103, 371 (Fig. 79)

keeled or rounded back, 101

length, 371
Gluten-cells, n
Gluyas, 299

Early, 299
Gneisendorf, 295
Godron, 45, 382-385
Golden Cross, 285

Drop, 302

Gem, 305
Golinski, 120
Goose, 220
Graebner, 153
Grain, colour of, 17, 372

development of, 129-143

ripening of, 139

twin, 359

vitality of, 32-34
Grains, in bushel, 427

in ear, 428

Granella de Carpegna, 292
Grass-clump plants, 356
Grenadier Svalof, 295
Greenland, 384, 385

THE WHEAT PLANT

Groom. 1 1
Gross, 34
Griiss, 30
Gynaecium, 113

Haberlandt, 29, 31

Habit of plant, 69, 78, 79, 363 (Figs. 65, 66)

Hackel, 152

Hadmersleben, 294

Haie, De, 298

Hairs on glumes, 103 (Fig. 79)

on leaves, 48, 53, 54 (Figs, in, 118, 137

151, 164)
Haller, 231
Hallett, 410

Hallett's Pedigree White, 291
Hard grains, 18-22

wheat, 207
Haussknecht, 188
Hedgehog wheats, 307-320
Heer, 266

Height of straw, 160
Herisson, 308

barbu, 313

sans barbes, 318
Hermann, 230
Hickling's Prolific, 295
Hieronymus, 325
Hindi, 274

selected, 301
Hoary, Old, 297
Hochfeldt, Homer's, 302

Winckler's, 302
Hofmeister, 16, 17
Hongrie rouge, 303
Hosagara, 284
Hosebonderie, 416
Host, 148

Howard, 360, 367, 368, 375
Hrozny, 187
Hsu Hsu Mai, 284
Huguenot, 229
Hyaline layer, 10
Hybrid grains, 364
Hybridisation', 360-406

method of, 362
Hybrids, compactum x other races, 401-

.405

dicoccum x other races, 392-394
durum x other races, 394
monococcum x other races, 390-392
polonicum x other races, 394-398
Spelta x other races, 405
sphaerococcum x other races, 405
turgidum x other races, 399-400
vulgar e x other races, 400-401
wheat, 377-406

x Aegilops sp., 380-386

x barley, 386

x couch grass, 380

x rye, 386-390

x rye grass, 380
wild wheat x Aegilops, 377

x cultivated races, 378-380

x rye, 377
Hypoderm, 92, 93, 98 (Figs. 72, 73)

Igel, 308

Imperial, Street's, 295

Improvement of wheat, 407-414

Indian Dwarf wheat, 157, 321-324, 346

Individual selection, 408

Inflorescence, 100

Inheritance of wheat characters, 363-377

Integuments, ovular, 136 (Fig. 109)

Internodes, bent, 357

diameters of, 64

lengths of, 63

malformed, 357

numbers of, 62
Inversable, Hybrid, 294

Japhet, Red, 295

White, 291
Javardo, 229
Jesenko, 387-390
Johannsen, 411
John Brown, 299
Jonathan, 291
Jones' Winter Fife, 298
Jordan, 283

Kajanus, 360, 371

Kalkori, 304

Kallak, 321

Kara Bashak, 228

Kar-i-safid, 286

Kathias, 221, 340

Keupely, 253

Kezer and Boyack, 393

Khapli, 194

Khorasan wheat, 155, 204, 340

Kintana, 278

Kinver Red, 295

Kizil Bogdai, 222

Kizildaly, 299

Kizil Goon, 275, 280, 286

Klebs, 17

Knight, T. A., 361

Kornicke, 118, 122, 152, 158, 178, 231

Kotschy, 178

Kubanka, 222

Kubb, 317

Velvet, 317

Kwang T'ou Mai, 303, 353
Kyu Shu, 303

Lagasca, 149

Lamarck, 147

Lancaster, 278

Larionow, 166-168

Late wheats, 85

Lateness, 161

Lawes and Gilbert, 81

Leaf arrangement, 45, 46 (Figs. 37, 38)

blade, 47 (Figs. 40, 41)

parenchyma of, 56

sheath, 47, 50, 51

vascular bundles of, 57 (Figs. 53-58)
Leaves, 44

anatomy of, 51-61 (Figs. 43-52)

hairs on, 48

length of, 49

INDEX

459

Leaves, number on culm, 49
Left-handed plants, 45, 46 (Fig. 36)
Leighty and Hutcheson, 124, 129
Leutowitz, 296
Ligule, 48 (Fig. 42)

structure of, 61
Link, 164
Linnaeus, 147
Little Club, 318

Joss, 301
Lobeiro, 215
Lodging, 78-81
Love and Craig, 364, 380, 394

Macaroni wheats 155, 206-229, 340
Madonna, 228
Magnesia, 213
Malakov, 278
Mark Lane, 292
Markischer, 290
Marquis, 293
Marvel, Red, 295
Mass selection, 408
Matador Rood Kop, White, 300
Maund, 361
Mazzochio, 254
Mealy grains, 18, 22
Medea, 224

Menenem Boydesi, 213
Mestome-sheath, 58 (i. Fig. 55)
Mettel's Squarehead, 295
Metzger, 151
Michaux, 335
Michigan Bronze, 285
Miczynski, 387, 389
Mid-season wheats, 85
Milk-ripe, 140
Miller, C., 73
Million, 292

Miracle, 253, 256 (2, Fig. 160)
Mocho de espiga quadrata, 318
Model Red, 295
Mogadur wheat, 231
Mogul, 299
Monarch, White, 292
Montgomery, 431
Morison, 230
Morton, 418
Motor-cells, 54 (Fig. 56)
Moula Oglau, 255, 256
Mourisco vermelho, 224
Multiple crossing, 412
Mummy wheat, 33, 256 (2, Fig. 160)
Mutants, 353
Mutations, 353
speltoid, 355

Nach Sinde, 194-196

Nakao, 119, 388

Napoles, 289

Nemec, 29

Niekirk's, van, 283

Nilsson-Ehle, 355, 360, 365,369,370, 373,

376, 403
Nishkin, 281
Xivieria, 151, 170, 173

Xobbs, 291, 299
Nomenclature of wheats, 163
Nonette de Lausanne, 258
Norden, 418
Norfolk, New Red, 295
North Russian, 302
Nowacki, 63
Nucellus, 10, 130

Oakley, 298

Obermayer, 129

Odessa, 299

Old Hoary, 297

Olivier, 335

Olura, 187, 325

Onega, 285

Ophiobolus, 8 1

Origin of wheats, 335-346

Ovary, 113, 130 (Figs. 89, 91)

Overton, 118

Ovule, 113 (Figs. 91, 99)
development of, 120
integuments, 136 (Fig. 99)

Paighambari, 321
Palea, 104

anatomy of, 109
Pearl, 297

Pedigree selection, 410-411
Pee-Laby, 54
Pendule wheat, 261
Pera, La, 285

Pericarp, 7, 136, 138 (Figs. 89, 109}
Pericycle, 40 (Figs. 26, 29, 30, 33)
Perle de Nuisement, 282
Persian Black, 196
Persoon, 148
Petianelle blanche, 248

noire de Nice, 260
Petiver, 261

Petrie, Flinders, 32, 33, 206
Phillipino, 251
Pierre, 81

Piliferous layer, 39, 43 (Fig. 26)
Pisana blanca, 249

francesca, 260
Pliny, 148, 241
Plot, 261
Plukenet, 230
Plumule, 12, 14 (Figs. 9, 18)

sheath, 14

Polar nuclei, 120, 130 (Fig. 100)
Pole-Evans, 376
Polish wheat, 156, 230-240, 340
Pollard, Grey, 261

Red, 259

wheat, 241
Pollen grain, 112 (Fig. 88)

mother-cells, 117-119 (Fig. 97)
Pollination, 129, 362
Pombinho, 252
Poulard a six rangs, 250

blanc Taganrog, 249

d'Australie, 260

rouge, 255
Pridham, J. T., 386

460

THE WHEAT PLANT

Prizetaker, 299
Prolific, Hickling's, 295
Prophyll, 44, 51, 52 (Fig. 59)
Proportion, grain to chaff, 432

grain to straw, 432
Prostrate habit, 69, 162 (Fig. 66)
Pubescence of leaves, 364
Pudel, 297
Punjab, Type i, 219

Type 2, 221

Type 3, 212

Type 4, 323

Type 5, 324

Type 6, 324

Type 7, 324

Type 9, 281

Type 20, 299

Type 21, 298

Type 22, 301
Pure line, 411
Puriewitsch, 30
Pur os, 335
Purple grain, 17, 195, 196

Rachilla, 101 (Fig. 76)
Rachis, 100 (Fig. 76)

anatomy of, 105, 106

brittle, zoo, 364

tough, 100, 364
Rainfall and growth, 5

and tillering, 75

Rapado de espiga quadrata, 318
Ratel, 258
Ray, 230
Raynbird, 361
Regel, 16, 382
Rent Payer, 296
Requien, 380
Resistance, frost, 162

rust, 375

Reynold's Discovery, 260
Rhizome, 65, 67 (Figs. 59-61)

anatomy of, 105, 106
Richard, 7
Rieti, 277

Right-handed plants, 45, 46 (Fig. 36)
Rimpau, 357, 388,. 399, 402
Rimpau's Schlanstedt, 301
Ripening of grain, 139, 140

inheritance of, 363

stage of grain, 141
Risciola, 214
Rivet wheat, 156, 241-261

Red, 258

White, 259
Roemer, 150
Roermaker, 275
Romanella, 284

Root, anatomy of, 39-43 (Figs. 26-34)
Root-cap, 42, 43, 135 (Figs. 31, 107, 108)
Root-hairs, 35, 39 (Figs. 26, 28)
Root-system of embryo, 14

of plant, 35-38
Roots, adventitious, 35-37 (Figs. 22, 24, 25)

and lodging, 79

origin of lateral, 43 (Fig. 33)

Roots of Spring and Winter wheats, 79

(Fig. 67)
seminal, 35, 36
Rostafinski, 231
Rothamsted, 34, 419, 420
Rouge barbu de Mars, 285
Rough Chaff, Essex, 297
Riimker, 354, 402
Russian, North, 302
Rust-resistance, 375
Rye, Giant, 231
Rye grass x wheat, 380
Rye, Montana, 231
Rye x wheat, 386

Sachs, 17
Saidi, White, 263
Sakamura, 119
Salmon, 129
Sandomir, 300
Santa Marta, 223
Saragolla, 213
Sari Bashak, 222

Bogda, 280, 287

Sakij, 222

Saumur de Mars, 294
Snunders, C. E., 366
Sax, 119, 130
Schenk, i i

Schimper, W., 195, 231, 262
Schlegel Dinkel, 333
Schliemann, 170, 206
Schoute, 68, 72
Schrank, 148
Schiibeler, 84
Schubler, 150
Schultes, 150

Schulz, 183, 184, 187, 336, 377
Schwendener, 58
Scutellum, 13, 16, 17, 134 (Figs. 9, 10)

vascular bundles of (Figs. 13-15)
Seed, 10-17

amount sown, 421

large v. small, 426, 427

size of, 425

variety, 422
Seigle wheat, 305
Selection, individual, 408

mass, 408

of hybrids, 411

pedigree, 410-411
Self-pollination, 129
Seringe, 150, 151, 173, 231, 263
Serum reactions, 338
Seven-headed wheat, 253
Shaw, 420
Sherard, 168
Shirreff, C. H., 429
Shirreff, Patrick, 73, 358, 360, 409
Shirreff, S. D., 296
Shirreff's White Bearded, 276
Shooting of the stem, 81-92
Shutar Dandan, 205
Siah Das, 205
Sicilio, 252
Siebritz, 282

INDEX

461

Silver King, 298

Sinz, 162

Sitopyros, 152

Small Spelt, 154, 170-177, 338

Wild, 154, 164-169, 338
Snowdrop, 297
Solid Straw Tuscan, 290
Sonora, 304
Sorrentino, 215
Soshu Shirokawa, 277
Sowing, time of, 422
Spelda, 325
Spelt, 100, 147, 158, 187, 189, 325-334»

345

Small, 154, 170-177, 338

Wild, 154, 164-169, 338
Spelta, 325
Spelta amylea, 151, 186

vulgaris, 151
Speltoid mutations, 355
Spermoderm, 7
Spikelet, 101, 428

development of, 114 (Figs. 92, 93)

supernumerary, 359

terminal, 100
Sports, 347-352

tera to logical, 357
Spring wheats, 162
Squarehead, 296

Beseler's, 296

Criewener, 296

Essex, 296

group, 272

Mettel's, 295

Strube's, 296

Svalof, 296

Squarehead's Master, 303
Stamen, 1 1 1
Standard Red, 303
Standup, 291

Red, 303
Stapf, 342, 346 '
Starch parenchyma, 1 1
Stem, 62-99
Stems, diameter, 64

number of, 72, 73, 75, 77

shooting of, 81

vascular bundles of, 95 (Figs. 72-74)
Stereome of leaf, 53, 60 (Figs. 46, 47, 55)
Stigmatic branch, 113 (Fig. 90)
Stoll, 399, 401
Stomata, 55 (Figs. 43, 48, 49)

number of, 55
Strabo, 335
Strauss, 167, 178
Straw and grain, 432
Straw, growth of, 82, 83

height of, 1 60

hollow, inheritance of, 364

solid, inheritance of, 364

strength of, 80, 81
Straws, number of, 72, 73, 75, 77, 428

per square yard, 428
Strong wheats, 23, 414
Styles, 113
Suffolk Thickset, 295

Sunagawa, 284
Superlative, 299
Surinam wheat, 231
Susceptibility to rusts, 375

Tapetum, 119 (Figs. 97, 98)
Temperature of germination, 27, 28

of growth, 5

Teratological sports, 357
Testa, 7, 10, 137
Teverson, 303
Thaoudar, 167
Theophrastus, 171
Thew, 291
Thick-seeding, 429
Thickset, Piper's, 361

Suffolk, 295
Thin-seeding, 429
Tillering, 64-78

and yield, 431

depth of node, 67

of Spring and Winter wheats, 75, 76
Tip-bearded, 288
Tiphe, 171, 207
Touzelle rouge, 302
Treasure, White, 291
Trichomes of leaf, 54
Trigo alonso, 216

Andaluze, 219

bascunano, 224, 225

berberisco, 224

bianco Verdeal, 213

blanquillo, 213

candeal, 275

cannu altu, 216

Canuto, 313, 315

carbillo, 312

Chacon, 216

Chile de fideos, 238

claro, 213

Colorado, 224

de espiga azulada, 216-

de espiga larga blanca, 213

de la Vuida, 312

enano de Larco, 220

Fanfarron, 212

Focense, 257

Granadino, 220

Macolo, 216, 220

marzuolo, 217

negro, 220

Obispado, 214

Pinot, 224

raspinegro, 216, 224, 229

Recio, 220

Regadio, 294

Rubio, 254

Salmeron, 256

Santa Salamanca, 248

Siciliani, 217, 225
Trimenia, 214
Triticum, 147, 266

aegilopoides , 154, 164-169, 178, 184

aestivum, 147

alatum, 206

amyleum, 186

462

THE WHEAT PLANT

Triticum Arias, 149

aristatum, 150

Arras, 193

atratum, 201

Bauhini, 149

bicorne, 149

Blat de Caure, 149

boeoticum, 164, 166

candidissimum, 148

cereale, 148

cerulescens, 148

Cevallos, 149, 150

Cienfuegos, 149, 200

Cochleare, 149, 150

compactum, 157, 307-320

compositum, 147, 148

creticum, 148

dicoccoides, 154, 178-186

dicoccum, 155, 186-203

durum, 155, 206-229

Eloboni, 195
farrum, 149, 186, 197
fastuosum, 149, 150

Forskal, 149

Gaertnerianum, 149, 150
glaucum, 230

hermonis, 178

Hisbu, 200

hordeiforme, 148

Horneman, 149

Horstianum, 150

hybernum, 147

Koeleri, 149

levissimum, 230

Linneanum, 149, 150

maximum, 148

monococcum, 154, 170-177

muticum, 150

orientale, 155, 204-205

platystachyon, 150

platystachyum, 149

Poloniae, 231

polonicum, 156, 230-240

pyramidale, 156, 262-264

ramosum, 241

romanorum, 241

sativum, 148, 149

sibiricum, 150

stculum, 150

Spelta, 158, 186, 325-344

speltoides, 337

sphaerococcum, 157, 321-324

spinulosum, 149

sylvestre, 148, 307

Thaoudar, 167

tomentosum, 148

touzelle, 148

tricoccum, 196

tumonia, 149

turgidum, 157, 241-261

typhinum, 241

velutinum, 150

venulosum, 151

villosum, 148

vulgare, 148, 151, 156, 265-306

Ze#, 148, 1 86

Tschermak, 368, 369, 377, 378, 385, 391,

398

Tsirali-ak-sowsam, 213
Tube-cells, 9, 138 (Fig. 5)

rp, AJ- O

1 uedi, 228
Tukur Sinde, 195
Tunis wheat, 264
Turkey Red, 278
Tuscan, Solid Straw Red, 293

Solid Straw White, 290
Tusser, 258, 259
Twin grain, 359
Typhew wheat, 241
Tystofte smaahvete, 295
standhvete, 295

Ulka, 285, 302
Union C., 299
Uses of wheat, 5, 6
Utendorf, 302

Variation, 347

discontinuous, 347, 352

racial, 347

Variegated leaves, 358
Variety, 158
Varro, 148, 408
Vascular system of embryo, 14
Vavilov, 166, 168, 336, 392
Velvet Chaff, 297

Blue, 306

White, 297

Ventral scale, 13 (Fig. 10)
Vermelho fino, 226
Vesen, 174, 176, 327, 330 (Fig. 207)
Victor, 291
Victoria, White, 291
Villars, 148

Vilmorin, 151, 364, 400
Virgil, 408

Vitality of grain, 32-34
Voelcker, 20
Vogeles Dinkel, 333
Vogl, 9, 138

Walla Walla, 313, 316, 319
Walter of Henley, 416
Way, 8 1

Weak wheats, 23
Welsch wheat, 241
Wheat x Aegilops, 380
x barley, 386
x couch grass, 380
x rye, 386
x rye grass, 380
Wheat, origin of races, 335

uses of, 5, 6

where grown, 3, 4, 5
White Straw Red, 295
Wild Emmer, 178-185, 336

Small Spelt, 164-169

wheats, 335
Wilhelmina, 291
Willem I., 291
Wilson, 387, 401
Winckler's Hochfeldt, 302

INDEX

463

Windsor, Early, 299
Winter Fife, Jones', 298

wheats, 167
Wittmack, 206
Wit Wol Koren, 297
Wol Koren, Rooi, 304

Wit, 297
Worlidge, 259
Worsdell, 17 •

Xeixa, 284, 299

Yabroudi Bayadi, 218
Yellow-ripe, 141
Yerli, 217
Yiaf boyde, 214
Yield and amoirit of seed, 421
and climate, 419

Yield and methods of sowing, 425
and number of ears, 43 1
and number of grains sown, 427
and size of grain, 425
and soil, 420
and time of sowing, 422
large, 418
of single ears, 428
of single plants, 73, 426, 429-431
of wheats, 415-432
small, 418

Zade, 338
Zavitz, 427
Zea, 147, 187
Zeeland, 290
Zeia, 187
i Zwaart Baard, 219

THE EN7D

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