the origin of hooded barley

7/26/2019 The Origin of Hooded Barley

1/9

WN ED ND HOODED B RLEY

Frontispiece

Awned (Winter Club), Hooded (Nepal), and a hybrid showing weak awns born on

second flowering glumes of the hood. The Hooded form is very attractive to those who

handle barley because the stiff barbed awns are a serious annoyance, and even a danger.

Nevertheless the hooded varieties are so unproductive that they never s tay long in favor.

Th e change from th e stiff barbed aw n to the soft, much m ore leaf-like,, hood is a s trik -

ing one. W he re and when it originally occ urred is not certain ly kno wn , but a rece nt

mu tation has bridged the gap in a s ingle jum p.

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2/9

THE ORIGIN OF HOODED BARLEY

H A R R Y V . H A R L A N

Office of Cereal Crops and Diseases, Bureau of Plant Industry, U. S.

Depa rtment of Agriculture, Wa shington, D. C.

T

HERE must have been varied

and prolonged Himalayan whoops

of delight when some lucky Nepal

farmer found the first hooded barley.

For some thousands of years barley

harvesters had searched more or less

clothes for stray irritating awns, and

doubtless the vows were many to grow

anythin g else but, another year. The n

this new thing appeared, quite different

from any barley seen heretofore . Th e

long, harsh, stiff awn, with its edging

of countless needle-like teeth, was re-

placed by a three-pronged appendage

at the tip of the gra in. N ot only was

the awn gone, but the new structure

was not at all harsh, being more

leaf-

like in tex ture than the awn. Th e

gods of gossip and sex in the village

market must have been dethroned for

at least a day, and many a flea surely

had a chance to find new pastures in

the crowds that debated over the novel

exhibit, for primitive mountain farmers

are observant, at least until the advent

of schools.

Yet, the story of the hooded barleys

is not so complete as we would like

to have it. Such early referen ces as

we have been able to find, all trace

back to a barley which was found in

Nepal apparently between 1830 and

1837. A t that time exten sive collec-

tions were being accumulated at Sahar-

um pore. It appears that at least two

collectors, Dr. Wallich and Dr. Hamil-

ton, travelled in Nepal between 1810

and 1840 collecting for the Botanical

Ga rden at this place. An yhow , it was

about this time that someone found

Nepal farmers cultivating this variety.

Botanists over the world immediately

scrambled into print to give it a name.

Only a few succeeded in being the

first to do so. Th e Botanica l Gard en

of Saharumpore was under the direc-

tion of J. F. Royle, and in his treatise

on the botany and animal life of the

Himalaya Mountains in 1839 he gives

a drawing of a hooded barley under

the name of

Hordeum aegiceras.

Sch-

lechtendahl, in 1837, had already de-

scribed this barley in Linnaeus as Hor-

deum trifurcatum. Th is description

undoubtedly was made from specimens

sent to Euro pe from Saharum pore. In

the next few years numerous references

appeared in botanical literature . Th ere

were also frequent notes in the agricul-

tural press where it was known as

Nep al. Th e date of the introduction of

this barley into America is not known.

It probably was introduced quite soon

after it reached Eu rop e. It is men-

tioned by name in the

Genesee Farm-er

of 1844, but we have found no com-

plete description given until 1855, when

the editor of the

Country Gentleman

comments on a sample supplied by

I . W. Briggs, W. Macedon, N. Y.

Briggs attributes its origin to Hindus-

tan and the editor identifies it from

the description in Lowe s Practical

Aqriculture.

Th e first farm repo rts in

this country were quite favorable even

as they now are with farmers who try

it for the first time . W ith in two or

three years, however, the growers be-

gan to express dissatisfaction. On e

such expression occurs in the Cultivator

of 1862 in which a correspondent who

signs himself J. L. R. advertises seed

for sale. H e is appare ntly am ong the

first to be afHicated by the tr u th in

advert ising complex. H e say s: I

have grown it 2 years and want to go

back to the old varieties. O ne grea t

objection I have to raising it is, that

it thrashe s so hard . A machine might

separate it from the straw, but it can't

265

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3/9

The Journal of Heredity

be done by the flailat least that was

the case with mine. Th e heads break

very easily from the straw, and the

kernel is so completely enveloped in

the chaff that it is necessary to liter-

ally chop the heads fine to get the

gr ain out. I can see no object in rai s-

ing this grain, unless one is overrun

with rats and mice, in which caseif

this is his only crop of grainhe will

be likely to star ve them out. Th ey

would either have to emigrate or die

fo r they could not shell it fast enough

to keep themselves alive.

The novelty of hoods has always

proved very attractive to farmers.

Hooded sorts are still carried by all

seedsmen although even now we have

no hooded varieties that are equal to

the awned sorts in production, despite

the long and frequen t use of hooded

parents in hybrid combinations.

AH Varieties from a Single Origin

It is difficult t o surm ise w hethe r

all of the hooded barleys of the world

have come from the original sport in

N ep al or not. W hile we are not cer-

tain that the barley of that name

originated in Nepal, it was found there.

It must have originated there or near

by and probably within a reasonable

time prior to its discovery for it had

reached neither western Asia nor

Ja pa n. In the latter country it cer-

tainly would have been of interest to

plant grow ers. Th e Japan ese have al-

ways shown great interest in unusual

plan ts. W e do not know enough about

the western fringe of China to be able

to say what types are found there.

Hooded barleys, however, have been

found in numbers north and west of

Chin a proper. Th e ones I have seen

appear to be very similar to segregates

w hich we have isolated from Nepa l

crosses. Ba rleys indistinguishable from

the Nepal have been received from

China recently but their history is as

yet not clear. Even though a Nepal

type should be found in western China,

the information would add little that

might be used in unravelling the story

of this barley. T he hoods are obviously

a direct mu tation. Such a mu tation

would need to occur but once. N ative

agricultural people exchange seed and

plants with a facility that is surprising.

Moreover, Nepal was once a vassal

state of China and even up until the

time of the overthrow of the Chinese

Em pire , tribute was sent overland from

Nep al to Pek in. A dissemination of

this striking plant along the route

travelled would be less surprising than

its absence.

Structure of the Hood

The exact nature of the hood is

difficult to de term ine . T he tissues al-

tered are probably modified leaf tis-

sues. Th e alterna tion is in the direc-

tion of sexual modifications. Th e cen-

tral one of the three projections is a

flower. Th is flower often bea rs sta-

me ns with fertile pollen. T he two

lateral projections are quite leaf-like in

character and possibly represent outer

glumes. Th e aber rant floret may pro -

ject upward, outward or even down-

wa rd from the tip of the lemma. Th e

tissues immediately beyond the tip of

the lemma in the varieties with sessile

hoods are reverse d. T he small spines

or teeth which occur on the lemmas

and awns of barleys reveal the arrange-

m ent of tissue s. In awned varieties

of barley the teeth which occur on the

ne rve s of the lemmas and on the

awns point upwards in the direction of

grow th. In the hooded sorts, such as

Nepal, the spines on the nerves of the

lemmas point upward as in all other

barleys. Th e teeth on the central nerve

of the hood, however, reverse their

direction at the point where the hood

is attached, whether this be the tip

of the lemma or a point on the awn

wh ere the flower originates. A t the

base of the hood which is hanging free,

there is a cup-like cavity, in which more

or less perfect floral organs are devel-

oped. A s the tissue of the hood turn s

to form the cavity the teeth are again

rever sed. T he tissue beyond the base

of the cavity constitutes a normal

floral glume. Th is glume sometimes

terminates in a thin awn of some length

and may represent a third lemma rather


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H a rl an : Origin of Hooded Barley

7

than a palet, although it is smaller

than the opposite glume.

As has been indicated, 'hoods, in

hybrid populations, are of great in-

teres t. In some cases enorm ous pockets

are developed. Ag ain, the hood dwin-

dles to a me re trace . In the heteroz y-

gous condition involving a strong-

awned parent, combined with a weak-

hooded one, the hood is often repre-

sented by a tiny depression on the

dorsal surface of the awn and a cor-

responding protruberance on the ven-

tral side. Th is is accompanied by a

disturbance of the venation of the awn

in which the veins are for a short dis-

tance diverted from straight lines.

Otherwise the awns are of full length

and width, and normal in structure.

T he hoods of N epal are sessile. In

hybrids they are often elevated on a

short awn. Som etimes this awn may

be one-third or even one-half the length

of a norm al one. Usua lly the longer

the supporting awn, the smaller the

hood. In the produ ction of hooded

varieties by hybridization, difficulty is

experienced in securing fully sessile

strains. This difficulty varies with

the paren ts. H ybr ids involving barleys

of the Manchuria group produce many

strain s with sessile hood s. Th ose in

which varieties of the Coast group are

used as the non-hooded parent, sel-

dom produce segregates that are fully

sessile. In some crosses strains occur

in which the hoods are attached at a

point below the normal tip of the

lemm a. Th is results in a cleft which

exposes the tip of the kernel. Th ere

are also variations in which more than

one ex tra floral center develops. In

hybrids awns are often found showing

two successive centers of floral

stimulus. (S ee Figu res 2 and 3.)

The hoods are not always confined

to the lemmas and the awns. Som e-

HOODS ON OUTER GLUMES

Figure I

Above are shown outer glumes bearing

hoods; below, enlargements of two of them.

Hoods in such an unusual position are oc-

casionally found in hybrid progenies. Th e

existence of hoods on the outer glumes opens

up interesting speculations on the homolo-

gous relation of outer glumes and lemmas.


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5/9

VARIATION IN HOODS

Figure

A, hood of normal Nepal; B , hood on (awnless X N epal) s egr ega te: C, large hood; D ,

subsessile hood; E-G, unusu al hood forms, showing reversal of direction of teeth. Th is

suggests that the hood represents two flowers a'.tached end-to-end.



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T R A C E S O F H O O D S O N A W N S

Figure 3

Awns of plants heterozygous for hoods, showing vestigial hoods, some of them very small

as on the awn at the righ t. No te also the repea ed stimulus to hoo d-form ation on some of

the awns.


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7/9

MUTANT PLANT WITH HOODED INFLORESCENCE

Figure

At left is shown the entire plant and at right the tips of the two sterile culms and a

natu ral-siz e view of the head. Th is plant was a hybrid- between an awned variety from

Nepal, with Man churia, another awned variety. The plant was sterile, so it could not be

tested further. It is significant in throw ing light on the origin of hooded barley that this

mutant appeared in a form that came from near the place of origin of the hooded forms.



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H a r la n : O rig in of Hooded Barley 27 1

times, in hybrid progenies, hoods are

found on the outer glumes. The se

are naturally quite small as nutrition

is limited. T he occurrence of hoods

on the glumes leads one into specula-

t ion regarding the analog}' between

these organs and the lemmas, and into

the general field of homologous tissues.

(Se e Fig ure 1.)

The writer is not attempting to make

a news item out of tissue origins that

were known before he was born. How-

ever, there are certain relationships

evident in barley which are worth not-

ing and there are occasional abnormal

developments which, while not bearing

directly on the question of hoods, do

show variation which, in its fluctua-

tions, might include hoods.

Aw ns and Stygm as Hom ologous?

The vegetative part of a barley plant

is very simple. It consists essentially

of a jointed culm bearing alternate

leaves. Inclo sed in the base of each

leaf sheath is a bud. Flo ral org ans

are modified vegetative parts and in

barley there is not much to modify.

The nature of the modification is most

app are nt wh en it is incomp lete. In

1918 a plant was found at A berd een,

Idaho, which showed a whole series of

half-modified leaves at each node of

the rachis* Sometimes a flower was

enclosed; sometimes the vegetative

stimulus overcame the floral one and

a new branch arose. W hile flowers are

themselves modified branches, it is

modified leaves' tha t con stitute almost

the whole of the floral struc ture. Th e

ovary wall, for instance, is green be-

cause of the leaf chlorophyll layer

which it contains. Th e w riter has felt

for some years that the barbs on the

awns and the hairs on the stigma arise

from th e same basic tissue . If the awn

and the stigma branch are homologous

tissues their correlation is morphologi-

cal. In 1915 a high positive corre la-

See Many-noded Dwarf Barley, Ha rry

V. Harlan and Merritt N. Pope, Office of

Cereal Investigations, U. S. Department of

Agriculture, Washington, D. C. Jour.

ered Vol. XIII , No. 6; Wash., D. C,

June, 1922.

tion was found between the number of

teeth on the awns and the number of

hairs on the stigmas of smooth-awned

baileys. Th e awns of most barleys

are heavily toothed and the stigma

branches covered with abundant hairs.

In the fully smooth-awned barleys the

stigma branches are almost free from

hair s. In hybrid progen ies all deg rees

of smoothness of the awns and hairi-

ness of the stigma are found. Th ese

variations are hard to classify due to

the difficulty of counting either teeth

or hairs, but they are obviously cor-

related.

Whether or not these facts have any

significance here is an open question.

It does seem certain that the hoods

came about by modification of leaf tis-

sue. W he re modified, the changes were

such as to make the new cells com-

parable with those of the rachis nodes

which produce buds. Th is is shown

not only by the formation of stamens

and fairly well-developed ovaries, but

also by the profound effect seen in

tissues adjacent to the disturbance.

A Hooded Mutation

Supposing that this erratic stimulus

occurred but once and that in the

mountains of Nepal, in what barley

did it occur? W e have recently come

upon a little evidence. A m ong the 28

varieties used in the making of the

Composite Crossf there was one from

the high Himalayas of Nepal called

Ev eres t. Th is variety, l ike trifurc a-

tum , is naked but it is awned . In

one of the F

2

hybrid populations of

Everest X Manchuria , a mutat ion was

discovered. Th is plant was nearly leaf-

less, the leaf blades on the upper nodes

being reduced to mere spur s. T he

peduncle never elongated and th e. two

upperm ost leaves did not unroll . Th e

two stalks thus culminated in conical

t A Composite Hybrid Mixture, Ha rry V.

Harlan and Mary L. Martini, Principal

Agronomist in Charge, Barley Investigations,

and Assistant Botanist, Barley Investigations)

respectively, Office of Cereal Crops and

Diseases, Bureau of Plant Industry, U. S.

Department of Agriculture. Jour. Am. Soc.

Agron. Vol. 21, No. 4, Ap ril, 1929.



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7

The Journal of Heredity

tips consisting of rolled leaves. Th e

spikes did not develop beyond the em-

bryonic stage so that the culms were

not at all distended by any internal

gro wth . Special care was given this

plant, for mutations of such propor-

tions are rare in barley nurs eries. W e

have found but three of any magnitude

in 20 ye ars. De spite the special care

devoted to it, the plant did not thrive.

The two culms became yellow and the

plant, to all appearances, was dying.

W e had about given up hope of any

further growth when a third culm did

develop and eventually headed. Much

to our surprise, it was hooded. Hooded

barleys occur in progenies with cer-

tain awnless forms where the awnless-

ness is due in part to hooded ancestry,

but here for the first time hooded

barley appeared under observation from

parents of purely awned constitution.

Fortunately we had convinced ourselves

we were dealing with a mutation be-

fore we realized it carried unusual

significance. W e checked all possibili-

ties carefully. Th ere were no hooded

varieties near the Fi in the previous

season and even if there had been,

crossing could not account for the ap-

pearan ce of such a plant. Th e pro -

genies of Everest with hooded barleys

always behave as any other hooded

cross.

Th e possibility of a field hyb rid

and a mutation occuring in the same

seed are too remote mathematically to

need consideration. U nfo rtun ately the

variation was sterile. Its appearance

indicated that such was the case and

frequent pollinations were made with

no result. (See Fig ure 4.)

This , in brief is the most important

evidence we have found. W e would

have been better satisfied if the plant

had not been sterile. O n the other

hand, if it had been fully normal, the

evidence would have been less conclu-

sive. W hile the mutation did not give

us Nepal, it did give us a glimpse of

variations of the same nature.

The origin of any form is a guess,

but in summation we have these facts.

Hooded barley was found in Nepal at

a time when it had not spread very far

east or west of tha t longitu de. Th ese

barleys differ from the awned by a

single step which is in the form of

an extra floral stimulus resulting in a

second floral center. In a hybrid po pu-

lation involving a naked awned barley

from Nepal, a sterile hooded mutant

appeared. O ur guess is that hooded

barley appeared as a mutant in a naked

mountain barley of Napal or West

China not more than 150 years ago.

Roman Prizes for Large Families

T

H E municipal governm ent of R ome

has established six prizes, to be

awarded annually, over a period of five

years beginning with 1932, to the larg-

est and the healthiest families of the

city. Th e prizes do not consist of

money but of a warranty deed to a

small house or apartment comprising at

least four rooms and a kitchen. F o r

the prizes may compete all parents,

legally married, who, in the preceding

four years, have had at least three chil-

dren born to them in Rome and still

living. Pe rso ns who are affected w ith

transmissible diseases or who are ad-

dicted to alcoholism to such an extent

as to impair the robustness of their

offspring will be excluded from com-

petition for the prizes. T he prize s will

be awarded every year on April 21, the

anniversary of the natal day of Rome.

Jour. Amer. Med. Assoc.

97:1090.

1931.

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the origin of hooded barley

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