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MEMOIRS OF THE
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CONTENTS.
VOL. VI.

Hecror, G. P. Notes on Pollination and Cross-Fertilization in
the Common Rice Plant, Oryza Sativa, Linn.

Suaw, F, J. F. A Sclerotial Disease of Rice (with 3 plates
—1 coloured)...

Howarp, GABRIELLE L,C. Studies in Indian Tobaccos, No. 3,
The Inheritance of Characters in Nicotiana tabacum, L. (with

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Burtxer, E. J.. and Asput Hariz Kwan. Some new Sugarcane
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GraHAmM, R. J.D, Preliminary Note on the Classification of Rice
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Howarp, A., Leake, H. M., and Howarp, GABRIELLE L.C.
The Influence of the Environment on the Milling and Baking
Qualities of Wheat in India. No. 3, The Experiments of
1911—12 (with one coloured plate and a map)

PAGE,

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231

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/NOTES ON POLLINATION AND CROSS-FERTILISA-

TION IN THE COMMON RICE PLANT
ORYZA SATIVA, LINN

BY

G. P. HECTOR, ma., 3.8¢

Economic Botanist, Bengal

AGRICULTURAL RESEARGA INSTITUTE, PUSA

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MEMOIRS OF THE
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IN INDIA

NOTES ON POLLINATION AND CROSS-FERTILISA-
TION IN THE COMMON RICE PLANT,
ORYZA SATIVA, LINN,

BY
GB. HECTOR. AA... B.Sc.

Economie Botanist, Bengal

- AGRICULTURAL RESEARCH INSTITUTE, PUSA

PUBLISHED FOR

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I.
Il,
TE.

Introductory
Pollination
Cross-fertilisation

CONTENTS.

NOTES ON POLLINATION AND CROSS-FERTILISA-
TION IN THE COMMON RICE PLANT, ORYZA
SATIVA, LINN.

BY
G, P. HECTOR, .a., B.sc.,

Economic Botanist, Bengal,

InTRODUCTORY.

r

Dorinea the past two years a large number of varieties of
rice, cultivated in the districts of Lower Bengal, have been grown
on the Dacca farm, with the object of studying their characters
in single plant cultures and of ascertaining to what extent, if
any, natural cross-fertilisation takes place, as a correct estima-
tion of this latter point and of the precautions likely to be
necessary to keep varieties pure, is an essential preliminary to
the work of successfully distributing seed of improved types.
This preliminary work is necessary in the case of all crops, but
particularly so in the case of a crop like rice, in which the
number of cultivated varieties, differing frequently by most
minute points of difference, is very large, and the risk of accidental
mixture of seed very great, owing to the peculiar conditions under
which rice is grown.

Though the results obtained up to date are far from complete,
they have shown that while some types are much superior to
others and self-fertilisation is the normal process, cross-fertilisa-
tion does undoubtedly take place and under certain circum-
stances may considerably affect the successful introduction of an
improved type into any particular locality where other inferior
types are commonly grown. This question of pollination and

—

ya POLLINATION AND CROSS-FERTILISATION IN RICE.

cross-fertilisation in rice has not been studied in detail in India,
and in the following notes it is proposed to give a preliminary
account of our observations on the subject, as far as the districts
of Lower Bengal are concerned.

IT.— Po.iination.

A superficial examination of the flower of rice would lead
one to believe that cross-pollination by the agency of the wind
was the normal method. Shortly after the opening of the glumes
the anthers may be seen hanging downwards from their filaments
in such a way as almost to preclude the pollen from reaching the
stigma of the same flower if dehiscence has not already taken
place before this pendent position is reached, and frequently also
the two feathery stigmas may be seen protruding one on either
side from between the open glumes. Though no work has hitherto
been done in India on the subject, much has been done in other
countries, and a brief summary of previous literature is given
below. Knuth* states that the species is allogamous and that
the anthers are still closed when they emerge from the glumes,
and do not dehisce till after they have bent over and hang down-
wards. More recently a Japanese worker, M. Akemine,} has
given a detailed account of the morphology of the flower and of
the flowering of a race of swamp-rice, known as ‘‘ Akaghé,” oceur-
ring in Northern Japan. Only a summary? of this paper has been
seen by us. According to the summary, the author discusses in
detail the climatic factors affecting the flowering and setting of
the grain, describes the method of pollination, and gives cytolo-
gical details regarding the fertilisation process. He gives no
cases of the occurrence of cross-fertilisation, and concludes, from
observations and experiments on the flower, that self-fertilisation
is what usually occurs, pollination according to the writer taking
place just before the flowers open. Lastly, Fruwirth and Van der

* Hand-book of Flower Pollination, Vol. III, 1909, p. 521.
t ‘* On the Flowers and Flowering of O, sativa,” Agric. Gasette, Nogy6-Sekai, 1910-11.
t Cf, Bot, Centralblatt, No, 41 of 1911, pp, 370-71,

G. P. HECTOR. 3

Stok* summarize fully all the work done up to date, chiefly by
the latter in Java, on the subject of rice-breeding in general,
including the question of pollination and cross-fertilisation.
According to these authorities, self-fertilisation is the usual
process, but cross-fertilisation may not infrequently take place
between neighbouring plants, and must be taken account of in
all breeding work.

According to our observations on the Dacca farm, the actual
process of pollination is as follows :—

The spikelets, or flowers, mature from above downwards in a
fairly regular sequence, and in the case of any individual flower
ihe opening of the glumes and _ pollination takes place, as a rule,
on the same day as that on which it emerges above the level
of the leaf-sheath if this happens before mid-day, or at latest
in the course of the forenoon of the following day. In almost
every normal case, dehiscence of the anthers and _ pollination
take place practically simultaneously with the opening of the
glumes, sometimes even before they open at all, and at the
time of dehiscence the stigmas are still enclosed and the
anthers still within the shelter of the glumes. Hence self-ferti-
lisation appears to be the usual method. In the case of Aus
varieties, which in Lower Bengal flower during the months of
May and June, the opening of the glumes and dehiscence of the
authers commences usually between the hours of 7 and 8 a.M.
and continues till about 10 a.m. when it stops for the day,
but in the case of the Aman varieties which generally begin
to flower in late October or early November, not till later in
the forenoon, beginning usually between 9 and 10 AM. and
continuing till mid-day. The later time of day at which the
flowers of the Aman varieties open is probably due to the much
lower temperatures in October and November.

Immediately before the flowers open and dehiscence takes
place it may be seen that the tup of the anthers is just touching
the concave roof formed by the glumes which are still closed,

* Fruwirth, Die Ziichtung der Landwirtschaftlichen Kulturpflanzen, B. Y., 1912, pp. 36—64,

4 POLLINATION AND CROSS-FERTILISATION IN RICE,

and if at this stage the glumes are gently pulled apart, the
anthers may be seen to dehisce and the pollen to fall in
showers on to the feathery stigmas within. It is probable
that the pressure of the anthers—which at this stage are being
pushed up by the rapidly elongating filaments from below—on
the glumes, together with the slight shock caused by the opening
of the ees and the consequent access of air and sunlight,
are the immediate causes which bring about dehiscence.

Immediately after dehiscence, the glumes diverge a little
further apart, the filaments rapidly elongate till the anthers
protrude about their own length above the tip of the glumes,
and then bend slowly outwards and downwards so_ that the
anthers ultimately assume a pendent position. But before
this stage is reached, in every normal case the anthers have
already lost almost all their pollen and pollination has taken
place before they emerged at all.

Both the extent to which the flowers open and the length of
time they remain open vary considerably, but we have not been
able to discover any differences in varieties with regard to this.
It appears to be largely due to atmospheric conditions at the time.
In the warmer and moister months of May and June the flowers
of the Avs varieties open earlier in the morning, but appear to
open to a less extent, and to remain open a shorter time than in
the case of Aman varieties which flower in the colder and drier
months of October-November. According to our observations,
the whole process from the time of the opening of the glumes
and dehiscence of the anthers till the time when the latter
assume the pendent position, occupies on an average only about
fifteen minutes, in both Aus and Aman varieties, but in the case
of Aus varieties the flowers seldom remain open for a longer
period than half an hour in all, while in the case of Aman varieties
they may remain open for periods varying from about an hour to
an hour anda half. If the weather is wet and rainy at the time
when the flowers should normally open, as is frequently the case
when the Aus varieties flower, they may not open at all, or if
they do, they often do not close again, and a large percentage of

G. P. HECTOR. 5

such flowers set no grain. It is a matter of common observation
that a far larger number of empty husks are to be found in the
heads of Aus varieties than in Aman varieties, due probably to
the wet weather which often occurs in Lower Bengal when the
Aus rices commence to flower.

In about four days a whole inflorescence will ordinarily have
been pollinated in the way described above.

ITJ.—Cross-Ferrinisation.

Though the above is the normal process, the method of
pollination by no means precludes the possibility of cross-polli-
nation taking place. It has been pointed out above that the
stigmas are not infrequently to be seen protruding from between
the open glumes, while the stamens have not lost all their pollen
when they assume the hanging position, so that there is every
possibility, should self-pollination have failed, of cross-pollination
resulting, and we have collected evidence to prove that it is
by no means unknown, and under favourable weather conditions,
bright sunshine with a gentle breeze, possibly takes place to
a greater extent than we have as yet been able to prove. It is
practically certain, however, that such cross-pollination as does
take place is confined to neighbouring plants in the same plot,
aud that the pollen of one plant seldom if ever succeeds in
reaching the stigma of a plant more than a foot or two away.

In the following paragraphs an account is given of the
evidence we have so far collected as to the extent to which cross-
pollination has been found to occur.

In 1911 single plant cultures of 150 types of transplanted
Aman varieties were grown at Dacca farm from single plants
selected in the season 1910. No cases of splitting crosses were
found in these, and all were passed as pure at harvest time, but
in four plots a few variant plants were found. Subsequent
examination, however, in the laboratory, of the seed of certain
white-grained varieties, proved that in some, red-grained seeds
were to be found which had escaped detection at harvest time,
and possibly there were others which were not discovered. Six

6 POLLINATION AND CROSS-FERTILISATION IN RICE.

such cases were found, and in each case as many red-grained seeds
as could be picked out were sown and the seedlings transplanted
separately in 1912, every precaution being taken to prevent
accidental mixture. In three of the six plots the red seeds have
given rise to a few white-grained plants, proving that these were
heterozygous as regards seed colour.

To return to the variant plants mentioned above, seed of
four of these, which it was suspected might be crosses, was care-
fully preserved, sown and transplanted separately in 1912, and all
have given rise to a variety of types as described below :—

The first of these (labelled Ba 1), was found in a variety
named Gobrabali, a coarse rice from Bakarganj, and differed from
the type chiefly in having reddish-coloured awns. This in 1912
split into four types—

(1) Red awns, two inner glumes at flowering time pale green

with red tip, turning yellow when ripe, stigmas black ... 60
(2) Red awns, inner glumes at flowering time dark green with

red tip, turning mottled brown, stigmas black Be tS
(3) White awns, inner glumes at flowering time pale green with

no tip, turning yellow, stigmas white * wit alg
(4) White awns, inner glumes at flowering time dee green with

no tip, turning brown, stigmas white eee i 4
Ratio red awns to white awns, 3:17: 1

The second (labelled R 20) was found ina variety named
Ukulmadhu from Rajshahi. The parent plant had a small, fine
grain, the two small outer glumes yellow, and the inner glumes
yellow with brownish coloured apex. This in 1912 split into
three main types-

(1) Small outer glumes yellow, inner glumes at first green with

reddish tip, turning yellow with reddish tip, stigmas

purple or oy # .. 5 noted,
(2) Ditto, but with stigmas wife ssi - soy, bs
(3) Small outer glumes coloured reddish-brown, inner glumes at

first reddish with no tip, turning black, stigmas white

bo

The only obvious difference between types (1) and (2) in this
case lay in the colour of the stigma which can be accurately de-
termined only before pollination has taken place. Unfortunately

|

G. P. HECTOR.

the plot was not examined in detail till after flowering was
almost over, and the colour of the stigma could be determined in
only a small number of plants.

The third and fourth examples will be treated together.
These were red-grained plants picked from white-grained plots,
the red-grained plants in both cases being otherwise almost
indistinguishable from the remaining white-grained plants in the
plots. The first of these (labelled M 1) was found in a variety
named Aman paddy from Mymensing, and the second (labelled
B 15) ina variety named Bankalam from Bogra. These both
split in 1912 into red and white-grained plants, otherwise almost
indistinguishable except for grain colour, in the proportions given

below—
1, Ml, red-grained, 1911 ... 1912, red-grained ... 145 ) ratio.
white-grained ... 55) 2°6r.: lw.
2. Bld, red-grained, 1911 ... 1912, red-grained ah ae
white-grained ... 68)1°9r.: lw.

Besides these four cases, much further evidence has been
afforded by a series of Dinajpur varieties started from single
plants selected in 1911. The seed of these varieties was collected
from Dinajpur, partly from cultivators and partly from the
Bazaars, and all were very mixed. In 1911, eighty-six single
plants were selected from these Dinajpur plots and these were
sown and transplanted separately in 1912 and no fewer than seven,
or a percentage of 8°15, have given rise to a mixture of types.
These are described below in detail.

In the first (labelled D 28), the parent plant had the small
vuter glumes coloured reddish-brown, and the inner glumes
yellow with a dark apex. This in 1912 split into the following
four main types :—

(1) Outer glumes coloured reddish-brown, inner glumes at flowering time

green with red tip, turning yellow with dark tip, stigmas black... 2 noted.
(2) Ditto with stigmas white “ “AP as tO diy
(3) Taller, more robust plant, outer glumes ebkoned edstion: brown, inner

glumes at flowering time reddish, turning black, stigmas black ... 2
(4) Ditto with stigmas white Ee

8 POLLINATION AND CROSS-FERTILISATION IN RICE.

As in the case of R 20 described above, here again it; was
too late to determine accurately the stigma colour in types (1)
and (2), the only obvious point of difference between them, when
the plot was examined, except in the cases noted.

In the second (labelled D 6), the parent plant hada red
grain and in 1912 split into four main types, two with red and

two with white grains, as noted below.

D 6—Red-grained, 1911.

ee —— pi he So Bed

(1) Reddish leaf-sheaths and (2) Ditto, grain (3) Green leaf- (4) Ditto, grain
stem above nodes, grain amber ... 1. sheaths and white ... 26.
white page: tS stems, grain

red of vari-
ous shades

In the third (labelled D 7), the parent was red-grained, and
split into three main types, two with red grains and one with

white, in the proportions givea—

1. Glumes yellow, grain red se wee . 5O
2. Glumes mottled brown, grain red ... aa sa vee 34
3. Glumes mottled brown, grain white ass aor sestlia

The remaining four Dinajpur examples are similar to those
of M 1 and B 15 described above, viz., red-grained parent plants
splitting into red and white-grained offspring, almost indistin-
guishable except for the colour of the grain. The details of

these four cases are as follows :—

PARENT 1911, OFFSPRING 1912. | Ratio R. : W
Red-grained. Red-grained. White- grained, |

D 17 i 72 21 34 31

D 18 n 72 24 8 31

D 35 " 51 21 2°42: 1

D 38 68 32 21231

From a perusal of the above figures, together with those of
M land B 15 quoted above, it would seem that as regards
colour of grain segregation is taking place in the simple Men-
delian ratio of 3: 1.* epee snion experiments are in

* Cf. Van der Stok, |. c., p. 49.

G. P. -HECTOR, Y

progress with the object of verifying this and also of determining
the mode of inheritance of certain other characters. |

In addition to the examples described above of undoubted
splitting crosses, stray variant plants have also been found in
twelve of the 1912 pure line plots started from single plants
selected in 1910, and in fifteen of the 1912 Dinajpur series started
from single plants selected in 1911, amounting to about fifty in
all. Some of these may prove to be accidental mixtures but
some cannot be matched with any of our other types, and have
all the appearance of being the F, generation of crosses which
must have taken place within our area in 1911. Seed of all these
is being preserved and will be sown in 1913 and the results noted,

From the above cases there is reason to believe that natural
crossing in rice is more common than was at. first supposed,
Moreover, under the conditions in which rice is grown by the
cultivator, whose varieties are never free from mixtures, it would
probably take place to a considerably greater extent than in our
area at Dacca. Even if it did not occur to any greater extent
than the cases cited would indicate, it would be quite sufficient
to account for the extraordinarily large number of types which are
to be found, when one takes into consideration the extent and
great antiquity of the cultivation. Certain facts also which have
been brought to our notice lend further evidence to this belief.
For example, a cultivator from the neighbourhood of Chandpur
recently informed us that on his own land within the past twelve
years the number of distinct types to be found in his fields has
increased from eight or ten to almost a hundred, although he has
imported no new varieties,

The main conclusions to be drawn from the above results
are :—

1. That m Lower Bengal under favourable conditions cross-
fertilisation may take place in rice to an extent which may be
provisionally estimated at about 4 per cent.

2. That this cross-fertilisation takes place wholly through the
agency of the wind and would seem to be effective only between
flowers of adjacent plants to a radius of a few feet,

10 POLLINATION AND CROSS-FERTILISATION IN RICE.

3. That as regards certain characters at least, e.g., grain
colour, segregation along Mendelian lines appears to take place.

4. That so long as seed of a variety is kept free from accei-
dental mixture there is no risk of contamination from cross-
fertilisation, but that if seed gets mixed, cross-fertilisation will
undoubtedly take place between adjacent plants in a plot and to
an extent sufficient in a few years’ time to reduce a variety to
a number of splitting types. Hence the imperative necessity of
taking every precaution to keep seed of varieties free from

accidental mixtures.

Dacca, |
10th January 19138.

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The Wilt Disease of Pigeon-Pea and the Parasitism of MNeocosmospora
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Studies in Indian Tobaccos, No. 1. The Types of Nicotiana rustica, L.,
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Studies in Indian Tobaccos, No, 2. The Types of Nicotiana tabacum, L.,
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Studies in Indian Fibre Plants, No. 1. On two varieties of Sann,
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The Influence of the Environment on the Milling and Baking Qualities of
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The Bud-Rot of Palms in India by E. J. Butter, M.B., F.L.S. Price,
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The Economic Significance of Natural Cross-fertilisation in India by
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Millets of the Genus Sefaria in the Bombay Presidency and Sind
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WolSyRV;, INo; - aI.

Vol. 1V, No. TII.

Wools =Ve2No: FV.

Wol: IV, No. “VY.

Vol. IV,No. VI.

Vol; Va No. ie

Wolssa\n Nos LVe
Vol: V, INo: . Vv.
Vol= VI; No. 4;

Vol. VI,. No. II.

Vol. 1, No. Jip
Vol. I, No. PT:
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ViOlemslNOsn VE
Vol. I, No. V.
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Vol. I, No. VII.

Vol. I, No. VIII.
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Vol. II, No. I,

Vol, Ib No: il:

Vol. If, No. III.

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Studies in Indian Fibre Plants. No. 2.—On Some New Varieties of Hibiscus
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Notes on the Incidence and Effect of Sterility and Cross-fertilisation in
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Note on the Inheritance of Red Colour and the Regularity of Self-fertil-
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Observations on Certain Extra-Indian Asiatic Cottons by H. M. Leake,
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The Morphology and Parasitism of Rhizoctonia by F. J. F. SHAW, B.Sc.,
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The Inheritance of some Characters in Wheat—I, by A. Howarn, M.A.,
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The Influence of the Environment on the Milling and Baking Qualities
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The Varieties of Soy Beans found in Bengal, Bihar and Orissa and their
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On Phytophthora parasitica nov. spec. by J. F. DASTUR, B.Sc. (In the press.)

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Notes on Pollination and Cross-Fertilisation in the Common Rice Plant,
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The Composition of Indian Rain and Dew by J. WALTER LEATHER,
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The Composition of the Oil Seeds of India by J. W. LeaTHER, Ph.D.,,.F.1.C.
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The Pot-Culture House at the Agricultural Research Institute, Puisa, by
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Experiments onthe Availability of Phosphates and Potash in Soils by
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The Construction of Drain Gauges at Pusa by M. H. ARNOTT, M.INST.C.E.,
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The Loss of Water from Soil during Dry Weather by J. WALTER LEATHER,
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The System Water, Calcium Carbonate, Carbonic Acid by J. WALTER
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Water Requirements of Crops in India by J. WALTER LEATHER, Ph.D.,
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The Nature of the Colour of Black Cotton Soil by H. FE. ANNETT, B.sc.
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Water Requirements of Crops in India—II, by J. WALTER LEATHER,
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The Composition of the Milk of some Breeds of Indian Cows and
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Records “of Drainage in India by J. W. LEATHER, Ph.D., F.1.C.
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The fab System of Rice Cultivation in Western India by H. H. MANN,
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Vol.

Vol.

Vol

Vol.

Vol.

Vol.

Vol.

. II, No.
. II, No.

Sh aes
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. II, No.
. II, No.
. II, No.

. II, No.

II, No.

Itt.

lV, No.
IV, No.
IV, No.
IV, No.

IV, No.

II.
III.
Wf

v
VI.
VIl.

. II, No. VIII.

I,

LY,

fF

CHEMICAL SERIES—contd.

The Composition of the Milk of some Breeds of Indian Cows and Buffaloes
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p.sc. Price, Re. 1-8.

A contribution to the knowledge of the Black Cotton Soils of India by
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The Date Sugar Industry in Bengal. An Investigation into its Chemistry
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Evaporation from a plain Water Surface by J. W. LEATHER, Ph.D.,
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Studies in the Chemistry and Physiology of the Leaves of the Betel Vine
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The more Important Insects injurious to Indian Agriculture by H. M.
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The Indian Surface Caterpillars of the Genus Agrotis by H. M. LErrRoy,
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Individual and Seasonal Variations in Helopeltis theivora, Waterhouse,
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The Coccide attacking the Tea Plant in India and Ceylon by E. E. GREEN,
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The Mustard Sawfly by H. M. LEFROY, M.A., F.E.S.,F.Z.S.; andC, C.
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The Rice Bug by H. M. LEFROY, M.A., F.E.S., F.Z.S. Price, Re. 1.

Remarks on Indian Scale Insects (Coccid@) by E. E. GREEN, F.E.S., F.Z.S.
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The Red Cotton Bug by H. M. LEFROY, M.A., F.E.S., F.Z.S. Price, Re. 1.

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The Cotton Leaf-Roller by H. M. LEFROY, M.A., F.E.S., F.Z.8. Price, Re. 1-8.

Notes on Indian Scale Insects (Coccid@) by H. MAXWELL-LEFROY, M.A.,
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Life Histories of Indian Insects (Coleoptera—I), by H. MAXWELL-LEFROY,
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Life Histories of Indian Insects—II. Some Aquatic Rhynchota and

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Life Histories of Indian Insects—III. The Rhtnoceros Beetle (Oryctes
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The Food of Birds in India by C. W. MASON, M.S.E.A.C., edited by H.
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Evi Silk, by H. M. Lerroy, M.A., F.E.S., F.Z.8., Imperial Entomologist ; and
©, C, GuosH, B.A., Assistant to the Imperial Entomologist. Price, Rs. 3.

Tetriginew (Acridinw) in the Agricultural Research Institute, Pusa, with
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The Big Brown Cricket (Brachytrypes achatinus, Stoll), by C. C. GHOSH, B.A.,
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Life Histories of Indian Insects (Hymenoptera), by G. R. Dvurt, B.A.,
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Inquiry into the Insecticidal Action of some Mineral and other compounds
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BACTERIOLOGICAL SERIES,

Vol. I, No. I. Studies in the Bacteriological Analysis of Indian Soils, No. I. By C. M.
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VETERINARY SERIES.

Vol. T.-No. I. Anaphylaxis in the larger animals, by Major J. D. E. HoLmes, D.sc.,
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Vol. I, No. II. Salvarsan in the treatment of Surra in horses, dogs and rabbits, by Major
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Vol. I, No. III. Some more successful Experiments on the treatment of Surra in the
Camel with recommendations for Systemetic treatment, by A. S. LEESE,
M.R.C.V.S. Price, Re. 1.

BULLETINS ISSUED BY THE AGRICULTURAL RESEARCH
INSTITUTE, PUSA.

No. 1. Notes on Cotton in Behar in 1904, by H. M. Lerroy, M.aA., ¥.E.S., F.Z.S., Imperial
Entomologist. Price, As. 4 or 6d.

No. 2. An Outbreak of Cotton Pests in the Punjab, 1905, by H. M. LeEFRoy, M.A., F-.E.S.,
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No. 3. The Extension of Jute Cultivation in India, by R.S. FINLow, B.sc., F.C.S., Jute
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or ls. 2d. (Out of print.)

No. 4. First Report on the Fruit Experiments at Pusa, by A. Howarp, M.A. (Cantab.),
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No. 5. Report on trials of the South African Locust Fungus in India, by FE. J. Burier,.
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Entomologist. Price, As. 2 or 3d.

No. 6. The Ticks Infesting Domesticated Animals in India, by C. WARBURTON, M.A..
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No. 7. A Preliminary Account of the Biting Flies of India, by H. M. Lerroy, M,a., F.E.s..
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No. 8. Official and Recommended Methods for use in Chemical Laboratories of the Depart-
ments of Agriculture in India, by J. WALteER LEATHER, Ph.D., F.LC.,
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No. 9. Report on Cocoanut Palm Disease in Travancore, by E. J. BUTLER, M.B.
Imperial Mycologist. Price, As. 6 or 6d.

No. 10. Treatment and Observation of Crop Pests on the Pusa Farm, by H. M. Lerroy, m.a.,
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No. 11. On Flax Dodder, by A. HOWARD, M.A., A.R.C.S., F.L.S., Imperial Economic Botanist.
Price, As. 4 or 6d.

No. 12. The Making and Care of Lawnsin India, by A. HOWARD, M.A
Imperial Economic Botanist. Price, As. 4 or 6d.

No. 13. Sugarcane at the Partabgarh Experiment Station, by G. CLARKE, F.1.C., Agricultural
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Assistant Director of Agriculture, United Provinces. Price, As. 6 or 6a.

No. 14. The Milling and Baking Qualities of Indian Wheats, by A. HOWARD, M.A., A.R.C.S.,
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No. 15. Note on the Extension of Cultivation of Fibre Plants in India. Price, As. 6 or 8d.
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No. 17. The Milling and Baking Qualities of Indian Wheats, No. 2, by A. HOWARD, M.A.
(Cantab.), A.R.c.S. (Lond.), F.L.S., Imperial Economic Botanist; and GABRIELLE

L. C. Howarb, M.A., Associate and late Fellow of Newnham College, Cambridge.
Price, As. 6 or 8d.

No. 18. Report on the Outbreak of Blister-Blight on Tea in the Darjeeling District in 1908-09,
by W. McRaAgz, M.A., B.Sc. Price, Re. 1 or 1s. 6d.

No. 19. List of Names used in India for Common Insects, compiled in the Laboratory of the
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No. 20. Memorandum on Indian Wheat for the British Market, by Sir JAMES WILSON, K.c.s.1
Price, As. 4 or 6d.

» F.L.S.,

By PASC Se RD S75

No

No. °

No.

No. §

No. %

No.
No. :

No.

No

No.

BULLETINS ISSUED BY THE AGRICULTURAL RESEARCH

33

INSTITUTE, PUSA—contd.

Memorandum regarding Leading Eucalypts suitable for India, by F. Boorn-
TUCKER, Commissioner, Salvation Army, Simla. Price, As. 4 or dd.

The Milling and Baking Qualities of Indian Wheats, No. 3. ‘*Some New Pusa
Hybrids tested in 1910”, by A. HOWARD, M.A., A.R.C.S., F.LS., Imperial
Economic Botanist ; and GABRIELLE L. C. HOWARD, M.A., Personal Asst. to the
Imperial Economic Botanist. Price, As. 7 or 8d.

Insecticides-Mixtures and Recipes for use against Insects in the Field, the Orchard,
the Garden, and the House, by H. MAXWELL-LEFROY, M.A., F.E.S., F.Z.S., Imperial
Entomologist. Second Edition, Revised and Enlarged by T. BAINBRIGGE FLETCHER,
R.N., F-E.8-, F.Z.S- Price, As. 12 or ls, 2d.

The Indian Saltpetre Industry, by J. WALTER LEATHER, Pb.D., F.I.C., Imperial
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ant to the Imperial Agricultural Chemist. Price, As. 8 or 9d. :

Report on the Flax Experiments conducted at Dooriah in 1910-11, by E. M. VANpbsr-
KERKHOVE, Flax Expert to the Bihar Planters’ Association. Price, As. 6 or 7d.

Note on the present position of Cotton Investigation in India, by BERNARD COVENTRY,
Offg. Inspector-General of Agriculture in India. Price, As. 2 or 3”.

Experiments on the Cultivation of Sugarcane at the Partabgarh Experimental
Station, 1909-11, by G. CLARKE, F.I.c.; H. E, ANNETT, B.Sc. ; and SYED ZAMIN
HUSAIN, B.A. Price, As. 5 or 6d.

The Cultivation of Lac in the Plains of India, by C. S. Misra, B.A., First Assistant
to the Imperial Entomologist. Price, As. 8 or 9d.

Directions for the Cultivation of Eri Silk. Price, As. 3 or 4d.

Report on the Flax Experiments conducted at Dooriah in 1911-12, by E. M. VANDE-
KERKHOVE, Fibre Expert to the Bihar Planters’ Association. Price, As. 16 or 2d.

Wheat Experiments on the Botanical Area, Cawnpur, and their bearing on Wheat
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Botanist to the Government of United Provinces, and RAMPRASAD, Assistant
to the Economic Botanist. Price, As. 3 or 4d.

A Note on some Interesting Results following the Internal Administration of Arsenic
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Some Aspects of the Agricultural Development of Bihar, by A. HOWARD, M.A.,
A.R C.S., F-L-S., Imperial Economic BKotanist. Price, As. 4 or 5d.

BOOKS.

“Indian Insect Pests”, by H. M, LEFROY, M.A., F.K.S., F.Z.S. Price, Re. 1-8. (Out of
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“Indian Insect Life”, by H. M. Lerroy, M.A., F.E.S., F.Z.8. 3; and F, M. HOWLETT,
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“Wheat in India”, by A. Howarpb, M.A., A.R.U.S., F.L.8. 3 and GABRIELLE L. C.
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2 suey, 1913. ie Botanical Series. Vol. VI, No, 2.
Memoirs OF the
Department of Agriculture
in India
A SCLEROTIAL DISEASE OF RICE
2 FE J. F, SHAW, ne Cae) A.R.C.S.,. F.LS ,

PUBLISHED | FOR

THE IMPERIAL DEPARTMENT OF AGRICULTURE IN INDIA

BY

€ : Lp . THACKER, SPINK & CO., CALCOTTA
res WwW, THACKER & CO., 2, Cregp Lanz, LONDON
te Price, Re. 1. -
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JULY, 1913. BoTANICAL SERIES. Vol: VE was 2:

MEMOIRS OF THE
DEPARTMENT OF AGRICULTURE
IN INDIA

A SCLEROTIAL DISEASE OF RICE

BY

F. J. F. SHAW, B.sc. (LOND.), A.R.C.S., F.LS

AGRICULTURAL RESEARCH INSTITUTE, PUSA

PUBLISHED FOR

THE IMPERIAL DEPARTMENT OF AGRICULTURE IN INDIA

BY
THACKER, SPINK & CO., CALCUTTA
W. THACKER & CO.,, 2, Creep Lane, LONDON

CALCUTTA :

PRINTED BY THACKER, SPINK AND CO.

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te

A SCLEROTIAL DISKASE OF RICE.

BY

F. J. F. SHAW, Bsc. (Lonp.), a.R.0.8, F.LS.

Recent? investigations in plant pathology have shown that
a not inconsiderable number of the diseases of plants are to be
attributed to the ravages of sclerotial fungi. Considering the
habitat of these parasites it is not surprising to find that tuberous
crops (¢.g., potato, carrot, beetroot) seem peculiarly liable to
attack, the disease in such cases being usually known as “ root

3)
.

rot.” In other cases, however, the host plant is attacked in the
seedling stage, the symptoms closely simulating the ‘“ damping
off” due to Pythium and its allies. Of sclerotial fungi, which
cause such diseases, one of the best known is Rhizoctonia, to
which the “ root rot” of many tubers, as well as the “damping
off” of certain seedlings, is to be attributed. There has, however,
recently been detected in India a disease of rice, due to the
attack of a sclerotial fungus, which seems to present some distine-
tive feature in its effects upon the host plant.

The fungus is known as Sclerotium Oryza, Catt., and was
first described by Cattaneo (2), in 1879, as the cause of extensive
damage to the rice crop in Novara and Lombardy. His descrip-
tion was restricted to the morphology of the fungus on the rice
plant and the symptoms of disease in the crop; as, however, cul-
ture work and inoculations were not attempted, adequate proof of
the parasitic nature of the fungus was not obtained. Since the
work of Cattaneo the fungus has also been discovered in J apan by
Miyake-(7), and has, within the past year, been collected in
India,

a"
to

A SCLEROTIAL DISEASE OF RICE.

Sclerotium Oryza is not, however, the only member of. this
genus which has been recorded as a disease of paddy. Sel. glu-
male, Ces., is known to attack rice in Borneo, while a new species,
Sel, irregulare, is stated by Miyake to be the cause of some loss
in Japan. Both these can easily be distinguished from Sel.
Oryza by the size of the sclerotia. The former has been found
on paddy at Noakhali, Eastern Bengal, but does not appear to
be of any economic significance in India.

The genus Sclerotiuvm was founded by Tode (11) in 1790, and,
at present, includes a number of species of which the fertile
stages are known in only a few cases. In 1816 Esenbeck (8) des-
cribed six of the more important species with figures; he also
includes Thanatophytum Crocorum (Rhizoctonia Crocorum) and
Evrysibe suffulta as near relatives of this genus. In a work pub-
lished a few years later Fries (4) mentions fifty species and classi-
fies them into four tribes, a sub-division which has persisted to the
present day. It is interesting to note that he discards the name
Thanatophytum for the more modern one of Rhizoctonia. In
1869 an enumeration of species peculiar to the Rhine district was
given by Fuckel (5); an earlier publication by Kihn contains,
however, more interesting matter. Kiihn (6) gives some account
of the work of Leveille and Tulasne (12), published a few years
previously, by which the connection between Selerotinm Clavus,
D. C., and Claviceps purpurea, Tul., was established. He also
mentions that other species of the genus Sc/levotiwm have their
perfect stages among the Clavariacea. It is, of course, now
well known that the original genus Sclerotium of Tode is an
artificial one, the different members of which are really the sterile
forms of widely separate fungi. Agaricacea, Polyporacee,
Clavaniacea, LHypocreacee and Pezizacee are the groups
among which the fertile stages of different species of Sclerolium
are to be found,

Tue Drskase 1x tur Figzp.

In India the first collection of Sel. Oryze was made at
Noakhali, About a year later, in December 1912, fresh material

Fr. do We SHAW. 13

was obtained from localities as widely separate as Mandalay,
Samalkota and Pusa; the external symptoms of disease were
in all cases essentially the same. Infected plants can usually be
distinguished from their healthy neighbours by the phenomenon
of “tillering,” that is to say, the development of fresh green
shoots from adventitious buds at the base of the infected culm
(Pl. I, Fig. 1). The infected culm gradually turns yellow and
dies ; any grain which it bears is light and poorly developed, in
fact, there is usually nothing within the glumes. The most dis-
tinetive feature, indeed, the easiest means of detecting infected
plants in the field, is the ‘ tillering” from the base. In Pusa
the disease does not make its appearance until the paddy crop
is fairly well advanced; hence, even supposing that the fresh
shoots remain free from the fungus, an unlikely event, there is no
possibility of their producing any grain. It is the loss of grain
which constitutes the most serious damage due to this fungus.
In Burma the disease is one of the causes producing the condition

?

known as “‘ gwa-bo” ; as, however, a number of insect diseases of
paddy are included under this name, it is ditticult to ascertain
the precise amount of damage due to the fungus. The collec-
tive damage done by the combined insect and fungus attack is
stated to run into hundreds of lakhs of rupees annually. Miyake
in his description of the disease in Japan says: “ Durch diese
Parasitierung wird die Bildung der Reiskérner unvollstinding,
daher ist der Schaden sehr-gross.” He does not appear to have
observed the phenomenon of “tillering,” a fact which Cattaneo
also overlooked.

If a diseased culm is split longitudinally the basal portion is
found to be infested with the fungus. The hyphe form a dark
greyish weft within the hollow stem, and small black sclerotia
can be seen dotted all over the inner surface (Pl. II, Fig. 1).
Sometimes the base of the culm is quite free from the fungus and
the attack begins at a node some distance up the stem. A trans-
verse section through an infected culm shows that hyphe and
sclerotia are not only present on the inner surface of the stem,
but that the hyphz penetrate the cells, and sclerotia are even

14 A SCLEROTIAL DISEASE OF RICE.

formed in the intercellular spaces between the main vascular
bundles. This is clearly shown in PI. III, Fig. 1. The section
shows only the inner portion of the stem; a young sclerotium
can be seen in one of the larger air cavities and the collapsed
nature of the innermost layer of cells, in the vicinity of the two
larger sclerotia, is apparent. In some of the cells hyphe are
distinctly visible; they are, however, more clearly shown in PI.
III, Fig. 2, from the same section. At first sight the sclerotia
strongly resemble that of Rhizoctonia Solan, Kiihn; they are,
however, considerably larger and have a distinctly smooth shiny
surface. In section the younger sclerotia appear to consist of
fairly small parenchymatous cells, the outer cells being more
or less definitely arranged in concentric layers. (Pl. III, Fig. 1).
In fact, at this stage, there is a distinct differentiation into
cortical and medullary zones. In the more mature sclerotia
the differentiation into cortex and medulla is not so apparent,
while the cell walls are thicker and of a sharp black colour

(Pl. ILI, Fig. 3).

CuLrurgs.

From diseased plants, such as those just described, cultures
were obtained on agar. It may be stated here that in all cases,
whether the cultures were made from hyphz or sclerotia, and
whether the material came from Burma, Samalkota or Pusa, the
same fungus was obtained in culture. Cultures were made
on media of widely different composition, and, in some cases, the
nature of the nutrient substratum was not without influence on
the character of the fungal growth.

Growth appeared to be most vigorous on glucose agar. An
infection upon this medium resulted in a copious development of
white hyph, followed, after five or six days, by the appearance
of sclerotia. The sclerotia are at first visible as minute circular
spots of a greyish colour ; subsequently they become black and
shiny, exactly resembling those found in the rice plant. The
hyphe are of the usual type, the cells being about 4—6m« broad
and 150—350 long, they contain numerous oil globules and

F. J. F. SHAW. 15

frequently branch. A transverse septum occurs at the point of
origin of a branch and not some distance from it (PI Il, Fig. 11,
ep. Rhizoctonia). The sclerotia are roughly circular and vary in
diameter from 150—500u. They arise from a plexus of interlacing
hyphe, which continue to branch and intertwine until a small
spherical compact mass is formed. For a time the young
sclerotium increases in size by the adhesion of fresh branches to
the periphery ; ultimately, the cell walls turn black and all
further growth ceases. At this stage the interior of the
sclerotium has a very definite parenchymatous structure, and it is
almost impossible to discern that it has been formed by inter-
weaving hyphe; a thin layer of loosely intertwined hyaline
hyphz can, however, be seen investing the exterior. Not infre-
quently several sclerotia become united, forming an_incrustation
on the medium resembling a stroma: this was particularly
common in cultures on maize neal.

In culture upon glucose agar, at an early stage, before the
appearance of sclerotia, the mycelium along the edge of the aga
appears black. An examination of the hyphe at this region
shows that they are more or less smoky coloured and of very
irregular shape (Pl. lI, Fig. 5); they frequently terminate in
curious appendages. These appendages may consist of one or
more cells ; they are very much darker in colour than the cells of
the hyphe, but resemble them in their irregular form (PI. IT,
Fig. 6). In some cases the parent hypha was attached to the centre
of the appendage in such a way that the latter was borne ina
peltate manner (PI. II, Figs. 7,8). Infection upon glucose agar
from these hyphz and appendages produced cultures of the
normal type resembling the parent culture. The fact that these
structures are produced along the edges of the agar, in contact
with the glass, suggested that they might be appressoria. If this
is the case, it is not easy to understand why their formation
should be restricted to one sort of agar medium: in no case were
they produced upon anything but glucose agar.

Infection upon nutrient agar containing extract of paddy
grains gave rise to a rather different habit of growth. The

16 A SCLEROTIAL DISEASE OF RICE.

hyphe at first spread over the surface of the agar, and slowly turn
« brownish colour where they are in contact with the nutrient
medium ; the aerial hyphw, however, remain white. Sclerotia
arise first upon the surface of the agar, but, ultimately, may be
found imbedded in it at various depths. At a level of about
1 inch below the surface of the agar the hyphe form a dense
brownish black layer, which, upon examination, is seen to consist
of an enormous number of chlamydospores. These chlamydos-
pores are formed by the segmentation of the hyphe intoa
number of short thick barrel-shaped cells. They possess thick
black cell walls and contain food reserve in the form of oil drops
(Pl. II, Fig. 9). Germination gives rise toa culture of the
normal type.

Growth upon the special medium of filter papers was slow
and produced nothing but hyphe and sclerotia. The same may
be said of French bean agar ; in this latter case the fungus seemed
to remain entirely superficial. Upon oat juice agar growth
was very similar to that upon rice agar; in fact, it is difficult to
distinguish between cultures of the same age upon these
two media. Upon lima bean agar the fungus gave rise
to a curious red pigment. This developed about three days
after infection and slowly spread down the tube following the
growth of the hyphe (PI. I, Fig. 3) Growth upon maize
meal was particularly vigorous, a dense white mycelium being
formed in about 12 hours, followed by an abundant pro-
duction of sclerotia. Here, again, the growth of the fungus
is characterised by the production of red pigment in the
meal,

The red pigment is strictly confined to the medium on which
it is produced. If an infection is made from a culture upen Lima
bean agar to glucose agar there is a faint reddening of the
glucose agar just at the seat of infection, but this speedily
dies away and does not spread down the tube, The produc-
tion of the pigment is obviously the result of the changed
metabolism conditioned by the alteration in the nutrient
substratum.

F. J. F. SHAW. 17

INocuLations.

In order to grow the rice plants under sterile conditions the
method used by Ward (13) in his investigations on the rust of
wheat was employed. The seeds were first sterilised in 1% com-
mercial formaline and then sown in sterile potato tubes containing
Knop’s solution. It was found that sterilisation was more
efficient if performed under the air pump, the liquid, in this way,
penetrating the space between the glumes more readily. Owing
to the laboratory temperature in February being rather low for
the growth of rice, the tubes were kept in an incubator at a tem-
perature of 30°C. The incubator was left with its glass door
facing a large north window, and, for several hours in the middle
of the day, the tubes were removed and placed in the sun

The young plants were infected when they were about 7—10
days old and about 3—4 inches high. The first series of infections
was made with small black sclerotia from an agar culture about
one month old. None of these inoculations gave any result, the
sclerotia failing to germinate. Subsequent trials with sclerotia
from old cultures showed that they had, not infrequently, lost the
power of germination. Fresh inoculations were then made from
a culture three days old, in which the hyphe were still growing
vigorously, and sclerotia were not yet formed. A small speck of
agar was removed from such a culture and placed upon a rice
culm, about 1 inch above the remnant of the seed ; hyphz quickly
spread from this centre over the exterior of the culm, which
gradually lost its green colour and turned brown near the seat
of infection. As the outer leaf sheath turns brown, the lamina
attached to it also loses its green colour and wilts (Pl. I, Figs.
2, 3); finally, the process extends to the central leaves, and the
whole plant dies. During the progress of the infection a light
weft of hyphe can be seen investing the culm ; in the later stages
of the disease small, dense, white aggregations of hyphz appear
in this mycelium, and, ultimately, become hard black sclerotia of
the usualtype. This superficial production of sclerotiais a charac-
teristic of the section Libera of this genus. We have, however,
seen that sclerotia may arise in the more deep-seated portions

18 A SCLEROTIAL DISEASE OF RICE.

of the host (PI. ILI, Fig 1), so too much stress must not be laid
on this character. The first sclerotia usually occur about the top
of the first leaf sheath, either on its inner or outer surface ; in the
former case they appear as small dark swellings beneath the dry
and withered leaf base. The time taken from the first infection
until the death of the plant and the preduction of the sclerotia is
about two weeks. |
Portions of dead and dying plants, some of which had not
yet produced sclerotia, were incubated in agar tubes. In all
cases they gave cultures of Sclerotiwm Oryze, which exactly
resembled those from which the inoculations had been made.
Since the infections showed that the fungus was strongly
parasitic, and could penetrate the uninjured external surface of
the plant, it was not considered necessary to make wound inocula-
tions. On the whole, 70—80% of the inoculations proved fatal.
Microscopic examination showed that the behaviour of the
mycelium in the infected culm was not without interest. The

bulk of the hyphze appear to run longitudinally in the large air
cavities of the leaf and in the cells bordering on them (PI. II,
Fig. 11), while a certain number grow outside the leaves in the
spaces between the folds of the lamina; these latter are particu-
larly obvious investing the delicate edges of the inner leaf (see
text figure). It is at spots such as this that the rice plant is

Ff. J. F. SHAW: 19

peculiarly liable to infection by the fungus. On the outer,
dorsal, surface of the leaf sheath the epidermal cell wails are much
thinner in the areas between the vascular bundles than they are
immediately behind them, and, at the edges of the leaf, they are
extremely delicate. Moreover, immediately behind the vascular
bundles, not only the epidermal, but two or more hypodermal|
layers have thickened walls, which give a cellulose reaction with
chlorzine iodide (see text figure). On the ventral surface of the
leaf sheath the epidermis is uniformly thin, and does not show any
such differentiation. Thus, on the outer surface of the leaf
sheath, there are areas which offer a poor resistance to any
parasitic attack. Once the fungus has gained an entry at one of
these spots, progress to the more delicate and deeper seated
tissues 1s easy.

The hyphz are both inter-and intra-cellular. It is by no
means unusual to find that a hypha which has penetrated a cell
wall possesses a pronounced thickening in the portion on that
side of the cell wall from which penetration has taken place (PI.
II, Fig. 11). This swelling may be taken as evidence of the
increase in chemical activity, probably in the direction of
the secretion of enzymes, which precedes the solution and
penetration of a cell wall.

A distinction must be noted here, between the behaviour of
the infected plants used in the inoculation experiments, and the
course of the disease in the field. A successful inoculation killed
the infected plant completely (see Pl. I, Fig. 3); but, in the
field, the result of an attack seemed rather to be a gradual
weakening of the host, culminating in the failure to produce good
seed. Further experiments on a_ field scale are necessary to
elucidate this point. It may perhaps be the case that rice plants
grown under the sterile conditions of our experiments were less
capable of resistance to the disease than under normal cir-
cumstances

ConcLusion,

The above account differs in several important respects from

that of Cattaneo. In particular the Italian author describes

20) A SCLEROTIAL DISEASE OF RICE.

some unusual structures in the sclerotium. He states, that
when the sclerotia are about a fortnight old several vacuoles
inake their appearance in the interior, and, ultimately, coalesce
to form one large central space; the sclerotium thus becomes
hollow. From the solid exterior hyphe now grow into the
central vacuole and bear spores. The spores are circular
structures, about 12 in diameter, and are either borne termin-
ally, or else laterally, on the walls of the hyphe. Such a
method of spore formation was never observed in our cultures,
nor did a series of microtomed sections reveal a sclerotium as
anything but a solid pseudoparenchymatous structure. The only
bodies bearing any resemblance to the spores of Cattaneo were
the oil drops in the hyphe. By crushing a sclerotium the
hyphze become torn and the oil drops set free. Under these
circumstances the oil drops may adhere to the sides and ends
of hyphie, in positions not unlike those which Cattaneo figures
for the spores.

No trace of a perfect stage was ever observed. Brefeld (1)
has pointed out that the sclerotia of certain Basidiomycetes
(Agaricus, Coprinus, Typhula), and of Penicillium and Erysiphe,
arise by the interlacing of branches of a single hypha, while those
of the genus Peziza are formed rather from a plexus of inter-
woven filaments. This comparison has been somewhat erroneous-
ly generalised by some writers (Stout 10) into the statement that
the sclerotia of Basidiomycetes arise from a single hypha and
those of Ascomycetes trom a plexus of hyphe. In the genus
Rhizoctonia it has been found (Shaw 8) that Rhizoctonia Solan,
Kiihn, forms its sclerotia from a single hypha, whereas, in the
macro-sclerotial species, of which Corticixwm vagum is the perfect
stage, the sclerotia arise from a mycelial plexus. It is evident,
therefore, that this character does not afford any basis for
taxonomic consideration, since both methods of sclerotial forma-
tion oceur in both Basidiomycetes and Ascomycetes.

It is interesting to note the changes in the appearance of the
fungus, according to the nature of the nutrient substratum.
These changes are most marked in the colour and form of the

F. J. F. SHAW. el.

hyphe, and, as Stevens and Hall (9) have already pointed out,
afford ground for reflection when we consider the characteristics
on which some groups of the Mung: imperfect: are classified.

Against a parasite such as Sc/. Oryze it is difficult to see
what remedial measures can be employed with success. The
sclerotia of the fungus undoubtedly perennate in the soil, where,
under favourable conditions, they germinate and produce a
mycelium which attacks the paddy crop. Cattaneo suggests
the application of salammoniac, with a view to killing the
sclerotia in the soil. Even if this is successful on a small scaie, it
is manifestly impossible to the extent which would be necessary
in India. Probably the breeding of resistant varieties is the
only method by which any permanent resistance could be made
in the case of a field crop such as rice. Fortunately the damage
done in India at present does not appear to be sufficient to bring
this question within practical consideration.

BIBLIOGRAPHY.

1. Brefeld, O. ... Untersuchungen uber Schimmelpilze, 1881,
p- Ube,

2. Cattaneo, A. ... Sullo Sclerotium Oryze, nuovo parassita vege-
tale. Archiv. d. lab Crittogamia. Pavia,
Wools. 1h Tt, 1879;

3. Esenbeck, Nees Von ... System der Pilze and Schwamme, Wurzberg,
1816.

4, Fries, Eliae ... Summa Vegetabilium Scandinavie, London,
1846, p. 476.

Systema Mycologicum, London, 1823, p, 246.

3. Fuckel, L. ... Symbolee Mycologice, Wiesbaden, 1869,
p. 404,

6, Kuhn, J. ... Krankheiten der Kulturgewachise, Berlin,
1858, p. 113.

7. Miyake, I, ... Studien tiber die Pilze der Reispflanze in

Japan, Journal College of Agriculture,
Tokio, 1910,

22 A SCLEROTIAL DISEASE OF RICE.

BIBLIOGRA PH Y—(con/d.)

8. Shaw, F. J, F, ... Morphology and Parasitism of Rhizoctonia,
Mem. Dept. Agr, India, Vol. IV, No. 6,
1912.
9, Stevens, FF. L, and Variation in fungi due to environment,
Hall, J, C. Botanical Gazette, 48, 1909.
10. Stout, A. B. ... Sclerotium disease of Blue Joint and other

grasses, Bull. 18. Univ. Wisconsin. Agr.
Exp. Sta., June 1911.

ll. Tode, H. J. ... Fungi Mecklenburgenses selecti, 1790,

12. Tulasne, L. R. .. Sur PErgot du seigle, Sclerotium Clavus
D. C., Comp. Rend., 1851.

13. Ward, H. Marshall... Proc. Roy. Soc., Vol. LXIX, 1902.

APPENDIX.
GiucosE AGAR.—
Extract of Lemco ae eos a8 4 grms.
Sodium chloride re an shi Dies,
Peptone aa ee Rae ee LO*
Glucose 4 re aa on ZO > 5;
Agar ase Ati oe a i ..,
Water i ee oie sep 2UOD exc,

Rick AGAR,—
Take 50 gris. of crushed rice seed and boil with 300 c.c. of water for
| hour, strain through a wire gauze. Dissolve 10 grms, agar in
200 c.c. water, add the decoction and heat to mix thoroughly.
Oat Agar, Lima bean agar and French bean agar are prepared in the
same way with the substitution of oats, Lima bean, or French bean for

the rice grains.

Finver PAPpER,—

Ammonium nitrate deo eas ys 10 gems
Magnesium sulphate Lies
Potassium phosphate ase ae 2 os
Lactic acid... ea ats ae le
Water _ ‘a wie use AUU0-0;0,

Take 50 c.c. of above solution, add 10 grms, filter paper and sterilise,

BR. Ji28) SHAW: ae

DESCRIPTION OF PLATES.

Prater’ I,

Fic, 1,—Rice plant infected with Sclerotinm Oryzw—note the young shoots
growing out from the base of the infected culm and the outer sheathing leaf
covered with sclerotia. x 1.

Fies. 2, 3,—Young plants inoculated with Sclerotium Oryze. In 3. scle-
rotia are forming. x 3.

Fie, 4.—Culture on Lima bean agar, x 1,

Puate II.

Fic. 1.—Rice plant with Sclerotiwm Oryzw—note sclerotia formed within
hollow stem. x4.

Figs. 2, 3, 4.— Young and old sclerotia from glucose agar culture. x 50.

Fie. 5,—Hypha from the edge of glucose agar culture. x 700,

Fig. 6.—Appendage on hypha. x 700

Fies.

Fie. 9.

Fic. 10,—Hypha from rice agar culture breaking up into chlamydospores

, 8.—Appendages borne in peltate fashion. x 700,
Chlamydospores from rice agar culture. x 700,

and single cells. x 700.
Fie. 11.—Longitudinal section of leaf showing hypbe in intercellular space

and cells bordering on it. x 700.

Prate ITI.

Fic. 1.—Microphotograph, Transverse section of stem of diseased rice plant
from Pusa crop. Note two sclerotia present on the inner surface of stem and
one small sclerotia in large air cavity. x 50.

Fic. 2.—Microphotugraph. The same showing hyphe in cells, The rounded

bodies with dark centres are starch grains, x 170.
Fic. 3.—Microphotograph. Section through mature sclerotium. x 130,

a
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PLATE I.

PLATE IL.

PLATE Ill,

Fiovot:

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a ~ Vol,

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‘Vol.

e SIN 5
4 rai Sa

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-

1's.” Shae Tul au eye OE Cee Ri Pa, Le Pe i A } |
ba | “ n VARA WAN ie eer hart a He Hash «th ons a4 WY)
a S57 ee ee M Y ] :

/ JULY, 1013 «—=——<“<~S*«~@ottanicl Seerriecss Vol. VI, No. 3.

I

Memoirs of the
Department of Agriculture
in India

; STUDIES IN INDIAN TOBACCOS
q No. 3. THE INHERITANCE OF CHARACTERS IN
i NICOTIANA TABACUM, L.
BY ne?
GABRIELLE L. C. HOWARD, m.a.
Associate of Newnham College, Cambridge and Personal Assistant
to the Imperial Economic Botanist

Bea _ AGRICULTURAL RESEARCH INSTITUTE, PUSA
i PUBLISHED FOR
es) THE IMPERIAL DEPARTMENT OF AGRICULTURE IN INDIA
‘ . : BY
+s) . THACKER, SPINK & CO., CALCUTTA

W. THACKER & CO., 2, CREED Lane, LONDON
~ Price, Rs. 3.

BAT vi
ae

July 1913 BoraNIcAL SERIES Vou. VI, Nox 3.

MEMOIRS OF THE
DEPARTMENT OF AGRICULTURE
IN INDIA

STUDIES IN INDIAN TOBACCOS

No. 3. THE INHERITANCE OF CHARACTERS IN NICOTIANA
TABACUM, L.

BY
GABRIELLE L. C. HOWARD, m.a.
Associate of Newnham College, Cambridge and Personal Assistant
to the Imperial Economic Botanist

AGRICULTURAL RESEARCH INSTITUTE, PUSA

PUBLISHED FOR
THE IMPERIAL DEPARTMENT OF AGRICULTURE IN INDIA
BY
THACKER, SPINK & CO., CALCUTTA
W. THACKER & CO., 2, CreED Lane, LONDON

HEADLEY BROTHERS.
PRINTERS,
BISHOPSGATE, E.C.; AND ASHFORD, KENT,

I

i ks bs ‘1 Ae: Mi " we Ane?

“ah

oe Ba oe

.

pels 4 ¢ @

og eee,

CONTENTS

I. Iyrropuction ae Se ae sh
II. Metsops oF RAISING THE EXPERIMENTAL PLANTS
Ill. THe OccurRRENCE oF PARTHENOGENESIS

LV. Tae EXPERIMENTAL RESULTS a
1. Time or FLOWERING ee fds
2. HeEIaut de ae are uss
3. NuMBER OF LEAVES PER PLANT ae
4, ARRANGEMENT OF LEAVES ON THE STEM
5. INSERTION OF THE LEAVES ON THE STEM
6. VENATION OF THE LEAVES ...
' 7. LEA¥-SHAPE... bape Bee
8. SURFACE AND MARGIN OF THE LEAVES .... ee
9. COROLLA ane om ae sts ues

V. CoNcLUSIONS on Aa aes

APPENDIX. DESCRIPTION OF THE TYPES USED IN HYBRIDIZATION

PAGE.
25

40

OrT 23 1919

STUDIES IN INDIAN TOBACCOS.

No. 3. THE INHERITANCE OF CHARACTERS IN NICOTIANA
TABACUM, L.

BY

GABRIELLE L. C. HOWARD, m.a.,
Associate of Newnham College, Cambridge and Personal Assistant to the
Imperial Economic Botanist.

I. INTRODUCTION.

THE chief direction in which the tobaccos of North-East
India can be improved is in the introduction of superior quality.
Many of the varieties at present grown give large yields and are,
in consequence, very profitable, but the cured leaf produced
from them is, as a rule, of very poor quality and is coarse,
deficient in texture, flavour and aroma. For this reason it can be
used for Indian consumption only, and, consequently, fetches
a very low price. Improvements in the quality of tobacco
may be obtained in three ways: (1) by the discovery of new
methods of cultivation by which healthy growth is promoted
and a larger yield and a better quality of leaf produced ; (2) by
the introduction of improved methods of curing; and (3) by
the growth of superior kinds. Some of the work done at Pusa
on the cultivation of tobacco has already been published." *
The investigations on curing are still in progress and it is hoped
to publish shortly the results obtained. The present paper
deals entirely with the third aspect of the question and is a

1 Howard & Howard, Memoirs of the Department of Agriculture in India, Bot. Ser.
Vol. III, No. 1, 1910.

2 Howard, Agricultural Journal of India, Vol. III, 1912.

26 INTRODUCTION.

continuation of the work already published as “Studies in
Indian Tobaccos, No. 2, The Types of Nicotiana tabacum, L.*”
In that paper an account was given of the work done with regard
to the study of varietal characters and the isolation of pure
forms. The stability of the type was discussed and it was shown
that there is no foundation for the belief, often expressed,
that the uniformity of type in any particular kind is easily
disturbed by its introduction into a new locality. If cross-
pollination be prevented, varieties or types of N. tabacum
remain as constant as those of other species of plants. This
result has recently been confirmed in America by Hasselbring.'
The methods of pollination were also studied and fifty-one pure
types were isolated. These types have since been maintained in
pure culture in the Botanical Area at Pusa and have bred true
to type from year to year. They form the material with which
the investigations, now to be described, have been carried out.

With regard to the improvement of the variety, the
immediate problem at Pusa is the production of a good cigarette
tobacco. The chief requirements in a cigarette tobacco for
growth in Bihar are :—

1. General robustness and rapidity of growth, both in the
seedling and later stages of the plant.

2. A plant of medium height with many leaves and short
internodes,

3. Fairly broad leaves with small veins, so that the cigarette
paper may not be damaged in the process of manufacture.

4. Yellow colour in the cured leat.

5. Good texture in the cured product.

6. Good flavour.

The characters in which the local tobaccos are most
deficient are those of texture and flavour. Several kinds have
been found possessing a fair colour.

3 Howard & Howard, Memoirs of the Department of Agriculture in India (Bot.
Ser., Vol. III, No. 2, 1910.

4 Hasselbring, Botanical Gazette, Vol. LIII, No. 2, 1912.

INTRODUCTION. 27

Many attempts have been made in the past to introduce
into India the best varieties of cigarette tobacco from America,
but the results have been disappointing. This is due to several
causes, some of which are avoidable. In the first case, in-
sufficient care was taken to prevent cross-pollination, and the
introduced varieties, by crossing with the indigenous crop,
deteriorated. In the second place, many of the varieties were
unsuited to Indian conditions by their habit of growth. The
practice in America of topping at a high level has favoured the
growth of tall kinds, which carry their best leaves well above the
-ground, making low or medium topping impossible. This is
a serious disadvantage in the plains of India, where high winds
frequently occur and damage tall varieties or those with thin
leaves. Another defect was noticed in all the American varieties
tried at Pusa, namely, the slow growth of the seedlings.
Although sown at the same time as the indigenous varieties,
transplanting could only be carried out a fortnight or ten days
later in the case of the American kinds, and there was a corres-
ponding lag all through the growth period. This is a very great
disadvantage in Bihar, where one of the secrets of success in
tobacco-growing lies in the maximum utilization of the growth
period from October to mid December. During this period, the
temperature is still high enough for rapid growth to take place
and the soil still contains plenty of moisture. After the
middle of December, when the temperature falls, growth is much
less rapid, and the plant should then be nearing maturity and
be ready to be cured about the end of January, in order to avoid
the dry hot winds which do so much damage during the latter
process. A crop, which through lack of food materials or
through the lateness of the variety makes little growth during
October and November, remains more or less stationery during
December and January and begins to grow again as the temper-
ature rises in February. Such plants seldom attain any great
size, and very frequently do not ripen evenly. The difficulty
in curing the product during the period of the hot west winds is

28 INTRODUCTION.

an additional disadvantage. Owing to lateness and want of
robustness, the yield of the American varieties is far below that
of the coarse local kinds and this would be fatal to their
successtul introduction.

The chances of improving the quality of Indian tobacco
by the introduction of a new variety from America are therefore
not great. It will be necessary to build up, by hybridization,
new kinds of tobacco, suited to Indian conditions of growth,
which possess in addition the qualities necessary to obtain a
better price. Fortunately, the introduced American kinds,
although they lose their colour by the native method of curing,.
nevertheless maintain their good texture and flavour, the chief
points in which the Indian tobaccos are deficient. Thus by
combining these desirable qualities with those of an indigenous
tobacco, which is robust and possesses a suitable habit of
growth, a very great improvement might be effected. Un-
fortunately, however, although tobacco is grown over so large
a portion of the world, very little work has been done on the
hybridization of this crop and little is known as to the
inheritance of the various characters which are of economic
importance. As in hybridization lies the greatest chance of
producing a permanent improvement in the tobacco grown in
the plains, and as it is possible to obtain at Pusa all the facilities
necessary for such an enquiry, it was decided to take up the
question and to make a thorough investigation of inheritance in
this crop, beginning with those morphological characters which
are of economic importance, namely, those concerning the
habit of the plant and the leaf. The subject has proved to be
far more complicated than was at first supposed, and the present
paper must be considered to be a preliminary one only. It will
probably take some years to obtain a complete knowledge of the
subject.

Besides its economic importance, there is another point
of great interest involved in the genetics of N. tabacum. Most
of the characters are concerned with the size of organs, and the

INTRODUCTION. 29

inheritance of these can only be determined by quantitative
means. The importance of a thorough study of the inheritance
of characters, which can be accurately measured, instead of
depending on observation alone, has been pointed out by many
writers." * Until recently, most of the investigations on inheri-
tance, undertaken from the Mendelian standpoint, have dealt
with characters of a qualitative nature, that is, they have
dealt with characters which depend on the presence or absence
of a particular attribute, such as colour, hairs or awns. Forms in
which these attributes are absent occur and can be used as
analysers, and the observations are restricted to a detection of
the presence of the character involved. By a judicious use of the
analysers and a careful analysis of the progeny, it is generally
possible by observation alone to determine the principles under-
lying the inheritance. Characters connected with the size of
organs present more difficulty. In the first place, no absence of
the character is possible and no analysers exist. Take, for
instance, the height of a plant or the length of a leaf, a plant
without height or a leaf without length is inconceivable. Thus
in such characters we are dealing with difference in degree only ;
observation is insufficient and measurement must be employed.
In the second place, such characters are generally very sensitive
to changes in environment and show marked fluctuating
variability. Such fluctuating variability may be inherent in
the plant, or may be directly due to the influence of the environ-
ment on the character under consideration, or indirectly to the
effect of the environment on the general vigour of the plant.
Should the changes due to fluctuating variability be greater or
almost as great as the differences in the characters under investi-
gation, they may obscure or entirely mask the effects of inheri-
tance. Hast® was one of the first to point out that so-called

1 Tammes, Recueil des Travaux Botaniques N éerlandais, Vol. VIII, 1912

2 Nilsson-Ehle, Kreuzungsuntersuchungen an Hafer und Weizen, Lund, 1909,
Kreuzungsuntersuchungen 11, Lund, 1911.

3 East, The American Naturalist, Vol. XLIV, 1910

30. INTRODUCTION.

continuous variation was capable of a Mendelian interpretation.
The whole subject of the inheritance of characters with fluctu-
ating variability has been very ably dealt with recently by both
Tammes'! and Nilsson-Ehle.2 The latter was the first to show
that characters, which to the eye appear similar, may in reality
be due to different genes which are inherited independently.
The red colour of the pericarp of some wheats is composed of
three factors, each of which will independently produce a red
colour, although less intense in tone than that due to the com-
bination. Recent work everywhere endorses the complicated
nature of most characters and has resulted in a large increase
in the number of factors recognised, while at the same time the
visible effect due to each factor appears smaller. In the paper
quoted above, Nilsson-Ehle discusses fully the question of
fluctuating variability and variation in general, and points out
that fluctuating variability may only exist as an effect of
environmental influence. If the number of factors n is large, the
number of homozygotic combinations possible will be much
larger, 7.e., 2? and the differences between these combina-
tions will be smaller than the differences between the factors
themselves. If the heterozygotic forms are intermediate in
value between the homozygotic combinations, we may obtain a
continuous series in the F, generation and as the combinations of
middle values occur most frequently, the form of the curve,
obtained in a graphic representation of the F, generation, will
resemble that of an ordinary frequency curve. The hetero-
zygotic combinations, which occur at different points on this
curve, will in the F, and succeeding generations give a progeny
which varies within much smaller limits than those of the F,.
Assuming a large number of small factors, this is sufficient to
explain all variation which is not induced by environment.
A plant which exhibits small fluctuations in any one character
may be heterozygotic as regards that character and it should be

1 Tammes, /.c.
2 Nilsson-Fhile, l.c.

INTRODUCTION. 31

theoretically possible to extract different types which breed
true from it. In practice, however, errors of measurement and
observation or environmental influence may be too great for such
types and their heterozygotes to be distinguished. From these
considerations as well as the experimental evidence of his own
researches, Nilsson-Ehle concludes that there is no inherent
difference in the mode of inheritance between quantitative and
qualitative characters and that all variations may be placed
in two groups :—

1. Variations which are inherited.

2. Variations which are not inherited and which are prob-
ably entirely due to changes in the environment.

It is obvious that the larger the number of factors in
which the parents differ and the greater the effect of environ-
ment, the more difficult it becomes to separate the factors or to
determine the exact mode of inheritance. If we consider the
case of two parents, which differ from one another in three
factors, there will be in the F, generation eight homozygotic
combinations and nineteen heterozygotic, which, in general,
will lie between the homozygotic forms. Thus we obtain a series
containing twenty-seven stages between the two parent. forms.
It the original difference between the parents is not very large,
these forms will lie very close to one another. If, in addition,
environmental differences supervene, the limits of variation of
one form will very soon overlap those of the next or even of
several others, in fact, in many cases, the limits of the two
parents themselves overlap. :

Taking these facts into consideration, it is not surprising
that up to the present in no case has the inheritance of the size
of an organ been entirely elucidated and the various factors
determined. All that has been possible has been to show that
segregation undoubtedly occurs and that the facts are in accor-
dance with the Mendelian interpretation and with the existence
of many factors, all capable of being inherited independently.
In the investigations described in this paper it has been possible

32 INTRODUCTION.

not only to show segregation in many of the characters, but also
to isolate forms resembling the parents as well as some new
constant forms differing from either parent.

The most thorough examination of the inheritance of
characters connected with the size of organs are the investiga-
tions of Tammes' on the length and breadth of the seed and
the length and breadth of the petal in Linum. The size of the
seed was found to be practically unaffected by the environment
and therefore formed very suitable material for such work. The
limits of variation and the co-efficient of variability were care-
fully determined for each parent. It was found that while the F,
generation was intermediate between the parents, the second
generation could not be separated into groups, but formed a
continuous series, and that, in many cases, no individuals could
be found which resembled the parents. In the F, generation
no individuals breeding true or resembling either parent could
be found but the limits of variation were smaller than in the
F, and differed in each case, the combined variation covering the
limit of variation of the F, generation. These results point to
the existence of several factors with segregation. The various
intermediate heterozygotic forms would naturally contain fewer
heterozygotes than the F, generation, and would therefore vary
within smaller limits.

East and Hayes,* in 1911, published similar results on size
characters in maize, such as height of the plant, length of cobs,
weight and size of seeds, but in most cases the investigations
were not taken beyond the second generation.

Three investigations on the characters connected with the
size of the organs in tobacco have been published, but in no case
have the investigations been carried beyond the second gener-
ation. Lock,’ in 1909, published a preliminary note of some
species crosses in the genus Nicotiana, the characters considered

1 Tammes, /.c.
* East and Hayes, Bulletin 167, Connecticut Agricultural Experiment Station, 1911.
5 Lock, Annals Royal Botanic Gardens, Peradeniya, Vol. IV, 1909.

INTRODUCTION. 33

being the colour, shape and size of the corolla. As the investiga-
tion is complicated by the fact that species crosses were
employed with consequent sterility and as it is admittedly only
a preliminary account with very few data, it need not be further
considered here.

A much more important paper dealing with N. tabacum only
was published by Hayes’ in 1912. The correlation and inheri-
tance of various characters such as the height of the plant,
number of leaves, average area of the leaves, average width and
average length of the leaves, were investigated in hybrids
between various pure types of American tobacco. Full details
are given of the measurements, but in no case have the cultures
been carried beyond the second generation. It was found
that the variability in the F, generation and the parents was
similar but much greater in the F, generation. These results
are most easily explained by the presence of a large number of
small factors with segregation. As regards the correlation
between these characters, the co-efficient in all cases was found
to be less than +. 5, except in the case of the length and width of
the leaf, where a distinct plus correlation was found. The
conclusions, as regards the individual characters, will be con-
sidered in more detail in the separate sections dealing with each
character in Chapter IV, but it may be remarked here that
although the results are undoubtedly valuable, many of the
measurements are taken in what appears to be a somewhat
arbitrary manner. For example, the number of leaves counted
is not the total number of leaves borne on the main stem of the
plant, but the number of leaves which occur between the fifth
leaf from the ground and the last leaf on a topped plant, this
representing the total number of leaves generally harvested.
Such numbers have an economic but no physiological meaning.
Similarly the height is measured to the last leaf counted, not
to the end of the main axis of the plant. As the habit of growth
of the American types used is very similar, discrepancies due

1 Hayes, Bulletin 171, Connecticut Agricultural Experiment Station, 1912.

34 INTRODUCTION.

to this method of measurement are not so great as they would
be in the case of many Indian tobaccos (see Plates I and II).
Nevertheless these arbitrarily chosen points cannot give the
expression of the true physiological activity of the plant as
regards height and number of leaves.

A third paper on the inheritance of quantitative characters
in Nicotiana is that published by Goodspeed.’ In the first part
of these investigations a comparison is made between the weight
of the seeds obtained by hybridization and the plants produced
from these seeds. N. tabacum var. macrophylla, a variety with
heavy seed, and N. tabacum var. virginica, a variety with light
seed, were employed as parents. The seeds of the F, generation
were divided into three groups, heavy, light and medium, and it
is stated that in the plants raised from the heavy seeds the
greater number of individuals resembled var. macrophylla,
whilein the culture raised from the light seeds the greater number
resembled var. virginica. The three classes of seeds showed
a difference in germinating power which was largely influenced
by time. The conclusion is therefore drawn that very variable
results may be obtained in the F, generation which are due
merely to the differential germinating power of the various
heterozygotic and homozygotic combinations. The experi-
mental data on which these conclusions are based can, however,
only be regarded as most unsatisfactory. In the first place,
considering the number of characters and the probably infinitely
larger number of factors involved in the difference between two
varieties of tobacco, it would be practically impossible to divide
the F, generation into three well-defined groups and any such
division which might be attempted would have no significance.
In the second place the original division of the hybrid seed
into three groups is open to question for the same reason. The
difference between these seeds probably depends on several
factors, and to these must be added environmental effects due to
nutrition and pollination. Very little meaning can be attached

1 Goodspeed, University of California, Publications in Botany, Vol. v, No. 2, 1912.

INTRODUCTION. 35

to results in which so many approximations have to be made.
In the second part of the paper, an investigation into the in-
heritance of the length of the corolla in three varieties of N.
acuminata is described and the author states that, although this
character shows very small fluctuations in the parents, the
variation in the F, is very great and covers the whole difference
between the two parents. The details of the F, generation
have not yet been published.

The only other investigations on hybridization in the genus
Nicotiana, which are known to me, are those by Jensen’ and
Lodewijks.*” Hybridization experiments were started by Jensen
in 1906 and the crosses investigated in most detail were those
between Peru, a variety with a petiolate leaf, and White
Burley, Peru and Maryland smoking. The intermediate nature
of the F, and the great complexity of the F, generation, with a
total absence of the parent forms was emphasized. The most
interesting point is the appearance of new characters in the
offspring which were not present in either parent.

1 Jensen, Jaarboek van het Department van Landbouw in Neederlandsh-Indie, 1907
1908, 1909, 1910, 1911.

2 Lodewijks, Zeitschrift fiir induktive Abstammungs und Vererbungslehre, Bd. V, 1911.

Il. THE METHODS OF RAISING THE EXPERIMENTAL
PLANTS.

THE methods employed at Pusa in raising the experimental
plants have already been fully described' in a previous paper,
and it will only be necessary briefly to recapitulate them. In
experimental work on tobacco, the two most important points
are: (1) to raise the seedlings without contamination, and (2)
to eliminate, as far as possible, all differences due to environ-
mental influences.

The seeds of the tobacco are so small that they are very
easily carried from one culture to another by wind, rain, earth-
worms, or by the hands of the workmen. The seed is brown and
indistinguishable from the soil, and retains its vitality, even
under adverse conditions, for several years. The practice
adopted at Pusa is to raise the seedlings in large shallow boxes,
and every precaution is taken to collect the earth and leaf-
mould from places where contamination by stray tobacco seed
is impossible. The boxes are made up about six weeks before
sowing and kept moist, so as to cause any stray seeds to germinate.
So far (1908 to 1912) no tobacco seedlings have been found in the
boxes prior to sowing. The boxes are sown, one at a_ time,
and the sower has to wash his hands before sowing another box.
After sowing each box it is immediately removed into the shade
till the seedlings appear. They are then enclosed in a wire
netting fence to keep off animals, and are placed so far apart
that the earth from one box cannot be splashed on to an adjacent
one by the sudden tropical storms which sometimes occur
at this season. Precautions are taken during the process of
thinning to prevent admixture and this operation is only

1 Howard and Howard, /.c.

GABRIELLE L. C. HOWARD. i'l

carried out under personalsupervision. The boys who dothe work
have to wash their hands after finishing each box, as otherwise
a few ungerminated seeds might be carried to other cultures.
After transplanting, the soil in the boxes is thrown away and
the boxes washed. These precautions have proved successful
and in no case has any mixture of. the cultures been discovered.

The elimination or rather the reduction of the differences
due to environmental influence is much more difficult. There is
perhaps no plant which is so sensitive to changes in soil, climate
and external conditions generally as the tobacco. The shortness
of the growth period, the large amount of material in the form of
leaves and stem formed in a short time are probably the reasons
why any check or stimulus has so great an effect. Moreover,
the tobacco plant appears to have an infinite capacity to adapt
itself to conditions, the same pure type giving rise to plants
14 feet or 8 to 10 feet high according to the cultivation. Even
in very adverse circumstances the plant goes through its com-
plete cycle and forms seed. Apart from such extremes, a very
small difference in cultivation is sufficient to induce a very
marked change and to raise a field of plants uniform enough
for accurate measurement is not easy. Nevertheless, in all
plant-breeding experiments, the absolute necessity of normal]
and well-grown plants cannot be emphasized too strongly.
The differences induced in tobacco plants apart from
size are almost incredible. The general effect of un-
favourable environment on WN. tabacum is to wipe out all
differences and to make the plants appear uniform. The
differences between the various types in leaf shape and leaf
surface, which are small, almost disappear in under-developed
plants. Unless the plants are well grown, it would be very
easy to be misled in observations on such characters as the
undulations of the leaf surface. The numbers would probably
show far too great a proportion of flat leaves.

Owing to the large amount of experimental work carried on
in the Botanical Area at Pusa, special care has been taken to

38 STUDIES IN INDIAN TOBACCOS.

render the land as uniform as possible. The plots are all small,
carefully levelled and are well drained and cultivated, and for
most crops they present an ideal experimental basis. In the
present investigations, a certain amount of trouble has been
experienced even on these experimental plots. Slight local un-
evenness of the ground due to ploughing, a difference in the sub-
soil drainage, and the proximity of a hedge have all had an
effect. It is needless to say that all cultures which could
possibly have been affected have been rejected. The cultures
which have to be directly compared are grown on the same
plot in lines and the two parent types are grown at both ends
of the plot, and also in the centre. In this way, should there be
any slight change in the conditions from one side of the plot to
the other, it would be indicated by the range in variation of the
parent forms. The impossibility of obtaining a large piece of
land with uniform drainage and soil must always limit the
number of cultures grown, even if the amount of work entailed
did not do so. One other important point must be mentioned,
namely, the time of transplanting. If many plants die after the
first transplanting and have to be reset, the replaced plants,
even though only a week later in planting, always remain behind
the earlier ones. By adopting a system of furrow irrigation,
and using great care in removing the seedlings from the nursery
boxes, the loss in transplanting in the experimental cultures
has been reduced to a minimum. Only one replacement is
carried out two or three days after the first transplanting.
Should others die, their places remain blank, but the system of
transplanting adopted has proved so successful that the number
of such blanks is very smallindeed ; the number of deaths before
the first replacement is generally not more than one per cent.

' The methods of crossing and raising the self-pollinated seed
are those in ordinary use, and need not be specially mentioned,
Full details have been given in a former paper.’ The only point

1 Howard and Howard, l.c.

GABRIELLE L. C. HOWARD. 39

which calls for comment is the limitation imposed on the
number of possible observations by the labour connected with
the raising of all the seed under bag. Many of the types will
not set good seed unless they are selfed, and the time taken in
carrying out all the details connected with this and the bagging
of a large number of plants is considerable. The raising of the
seed and the observations on the leaves have to be carried out
during the same period, between the formation of the first
flowers and the partial destruction of the leaves by death or
the ordinary chances of breakage. Thisin Bihar is a very short
period, and even by devoting the whole day to the work the
number of observations which can be carried out in the time
falls far short of those desired.

iI. THE OCCURRENCE OF PARTHENOGENESIS.

The question of parthenogenesis in N. tabacum was taken
up in consequence of a paper published in the Mendel Journal.'
In this communication the author stated that she had been able
to obtain parthenogenetic seed with the greatest ease in the case
of N. sylvestris, N. tabacum, N. suavolens, N. sandera, and
hybrids from these. In some cases only the anthers were
removed, but in others both anthers and stigma, while the
ordinary precautions of sterilizing the instruments and enclosing
the flowers in wax-paper bags seem to have been scrupulously
observed. Success did not attend all the experiments, but
“ parthenogenesis was discovered in ten species, varieties and
hybrids of Nicotiana on chosing the right period for trial, 7.e.,
when the plant is beginning to go off its fullest bloom. In the
tabacums success was unfailing.” East® also mentions the
possible production of apogamous seed. “In crossing species
of the genus Nicotiana I have had plants develop from seed
that have apparently been formed apogamously, that is, formed
from an immature egg-cell without fertilization. It is evident
that this is induced by the extraordinary irritation of foreign
pollen.”

Experiments were undertaken both in 1910 and 1911 to
determine whether, under the conditions obtaining in Pusa,
N. tabacum will set seed without pollination. It had already
been observed that castrated flowers prepared for hybridization,
which owing to pressure of work or other reasons had not been
pollinated, invariably dropped without setting any seed. In
order to obtain more definite information on this point, a large

1 Haig Thomas, The Mendel Journal, No. 1, 1909.
2 East, The Popular Science Monthly, 1910.

GABRIELLE L. C. HOWARD. +]

number of flowers on two individuals in nearly all the fifty-one
types of Indian tobaccos, and also in some F, hybrids, were
castrated. About fifty to one hundred flowers were prepared
on each plant under every possible condition. In some the
anthers were removed, in others, both anthers and stigma.
The plants used included types which self-fertilize with great ease
and those which will set hardly any seed unless selfed, as it was
thought that the latter would be the most likely to produce
parthenogenetic seed. Plants were chosen at all periods of their
growth—when in full seed formation, when full of capsules and
going off their bloom, and when very nearly over. In most cases
the plants were heavily pruned, all capsules, flowers and buds
other than the castrated ones being removed (such heavy
pruning ordinarily induces rapid seed formation), others were
lightly pruned. The same methods were adopted in 1911, but
here the number of kinds employed was smaller, only those used
as parents in the hybridization experiments were tested again,
namely, Types 9, 51, 16, 35, 23 and 38. The castrated flowers
were enclosed in parchment bags and these were taken off
at frequent intervals in order that any newly-formed buds
might be removed. In the earlier experiments the bags were
not applied after the corollas had withered, but in the later
experiments bags with perforations were placed over some of the
branches. A great difference was found between the capsules
formed from the castrated flowers and those formed by ordinary
pollination. In the latter case the capsule swells quickly and
remains firmly attached to the plant. No difficulty is
experienced in removing or replacing bags, and the peduncle
would have to be broken before the capsule could be removed.
This is always the case, whether the flower be self- or cross-
pollinated. The capsules of the castrated flowers, on the other
hand, although they also became swollen at first and simulated
the fertilized ones, were very easily detached from the plant. It
was exceedingly difficult to remove the bags, which finally had to
be cut away carefully. The capsules thus exposed to the air were

42 STUDIES IN INDIAN TOBACCOS.

easily blown or knocked off. For this reason half of them were
enclosed in large well-perforated bags as a protection. Some of
the capsules obtained a fair size, but were not so large as the
normal ones. On examination they were, however, found to be
quite empty, the ovules not having developed. On only three
plants did seed set in all the thousands of flowers which had
been castrated, and the total number of capsules was five.
In 1910, on a plant of Type 9, about one hundred flowers were
castrated and one fully formed capsule was found, the seed of
which germinated and produced plants similar to Type 9. In
1911, again, on a plant of Type 9, one capsule containing seed
was found in about one hundred castrated flowers. In this type
the stamens are so much shorter than the style, that if enclosed
in a bag the flowers normally set no seed and the majority of the
capsules drop. It has always been found necessary to self this
type in order to obtain sufficient pure seed to maintain the
culture. Ifthe seedsin these twocapsules were due to apogamy,
this method of seed production must be theexception and not the
rule. It isof course possible that the two capsules were due to errors
in.castration. The other three capsules were found on one plant of
another type, but by an accident these were not examined.
They were large, well-formed capsules, apparently containing
seed, but unfortunately were destroyed before this fact had been
definitely ascertained.

Considering the great number of flowers examined and
the fact that every stimulus to apogamous seed formation
had been given to the plants by pruning and capsule removal,
the results obtained are exceedingly small, even if we assume
that all five capsules contained parthenogenetically formed
seed.

In addition, in all the first generations (nine in number)
which have been raised at Pusa during the last five years, each
culture containing about one hundred plants, no individuals
resembling the mother plant have been detected. The cultures
have been absolutely uniform and the reciprocals identical. I

GABRIELLE L. C. HOWARD. 43

have no hesitation in saying that under the conditions obtaining
at Pusa in ordinary hybridization work, parthenogenesis in
N. tabacum is negligible.

It is interesting to note that in the details given by Jensen!
of an experiment undertaken to test the possibility of wind
pollination in N. tabacum, no mention is made of the formation
of apogamous seed. Four plants bearing 119 castrated flowers
were covered by a net and so placed that the prevailing wind
must bring pollen to them. In none of the flowers did any seed
set. The uniformity of the F, generation and the identity of
the reciprocals has also been mentioned by Jensen and by
Hayes.”

1 Jensen, l.c.
2 Hayes, l.c.

IV. THE EXPERIMENTAL RESULTS.

THE material used in these investigations formed part
of the types, isolated in 1909, which have since been maintained
in pure culture. Since at the time these experiments were begun
nothing was known concerning the correlation or interdependence
ot the various characters in tobacco, it was decided to make a
preliminary survey of the inheritance of practically all the
characters which deal with the stem and leaves. This was done
partly with the object of ascertaining how far the inheritance of
the individual characters could be studied independently, and
partly to determine the most suitable methods of investigation.
It is hoped to follow this preliminary account by a more
comprehensive study of the more important characters. The
characters which have been considered in the present investi-
gation are (1) time of flowering, (2) height of stem, (3) arrange-
ment of the leaves on the stem, (4) length of the decurrent
portion of the lamina, (5) venation of the leaf, (6) leaf-shape,
(7) undulation of the surface and margin of the leaf.

In all, five crosses have been made, Type 9 x Type 51,
Type 16 x Type 35, Type 23 x Type 38, Type 2 x Type 3, and
Type 2 x Type 51. The first, that between Type 9
and Type 5l, has been carried to the F, generation, those
between Type 16 and Type 35 and Type 23 and Type 38 to the
F, generation. Only the F, generation has so far been raised
of the other two. It will be seen from the photographs of
Type 9 and Type 51 (Plates I and II), and from the full
description of the types reprinted as an appendix to this paper
(p. 108), that these two forms differ in almost every character,
from height of plant to the mode of pollination and colour of
the corolla.

Aad

4

PLATE

IX.

ipl =)

PEATE Dkr

eal

ShabCle

PLATE Tie

LYE EV

PLATE

TYPE AAV.

PLATE V;

THXX AdAL

PLATE Wik

LYPE AXKXVINI,

PEATE: Vie

SUB Shep Ait

PEATE VIET

1X

PLATE

‘NW OOVEAVE

“N

NI

Sassou)

A©® NOLLVIaNa

ES MAel

Io

we
apr

Bey

ye
ee

\4

F4
Tos x Ts1

Fi SK,
Tes x Tass “Rye

FIRSD GENERATION IN N. TABACUM.

PEATE 2

PLATE. Qi:

Dic XxX V's;

FIRST GENERATION IN NN. TABACUM.

PEATE 2p

sich AWE

G AdAL

NOILVYANAD LSdIa

GABRIELLE L. C. HOWARD. 45

In Type 16 and Type 35 (Plates III and IV) we have two
forms which, while not very different in height or habit, never-
theless differ markedly in the number of leaves. The main
object of the cross between these two forms was an investigation
of the leaf shape. Type 35 has the broadest and Type 16 one
of the narrowest leaves in all the fifty-one types, while the length
is nearly the same in both. The cross between Type 23 and
Type 38 (Plates V and VI) was primarily made on account of the
difference in the number of leaves on plants almost equal in
height. The average number of leaves in Type 23 is nineteen
or twenty, in Type 38 about thirty-five. Types 2 and 3 (Plates
VII and VIII) are both petiolate forms, but the petioles are
alate to a varying degree. In the cross Type 2 x Type 51
(Plates VII and II) a form with a petiolate leaf was crossed with
a leaf with a broad base. The most striking result of these
hybridizations was the formation of new forms quite outside
the range of either parent. For instance, in the F, generation
of Type 16 x Type 35, dwarf plants, much shorter than Type 35,
and tall forms, almost equal in height to Type 16 and Type 35
combined, were found, and some of these have bred true in the
F,; generation. Similarly, in the cross Type 23 =x Type 38,
petiolate forms breeding true were produced, although both
parents have sessile leaves. Reciprocal crosses were always
made and grown side by side both in the F, and F, generations.
In no case could any differences between the crosses and the
reciprocals be detected. The F, was intermediate between the —
parents in almost all the characters (Plates IX, X, XI, XII).

Details regarding the manner in which the actual measure-
ments were taken will be given in the section on the individual
characters, but a few words of explanation may be given here
on the number of plants used in the experiments. As explained
in a previous chapter (p. 38), the area of uniform land available,
the labour involved in the raising of the seedlings and the
transplanting, but more especially the shortness of the period
during which measurements can be made, limits the amount

46 STUDIES IN INDIAN TOBACCOS.

which can be accomplished. The descriptions of the various
types of tobacco already published show that the different forms
of N. tabacum can be arranged as regards each character in a
series, each member differing very slightly from the next (see
also Tables VII, XII, XX). This would lead one to expect a
large number of factors in connection with each character, and
in the F, generation a long series of intermediates, with the very
rare occurrence of the parent forms. The chances of directly
analysing such an F, generation are small. A very large
number of individuals would have to be raised in the F, genera-
tion to obtain any indication of the percentage of plants
resembling the parents, and, since the range of variation in the
parents due to external influences is considerable, it would be
quite impossible to recognise such forms with certainty. This
being the case, there seems very little point in undertaking
the labour of raising many thousands of plants in the F, gener-
ation. The plan adopted has been to grow not less than 1,000
plants in the F, generation in order to obtain a fair idea of the
extent of variation by eye,and then measure as many as possible
of these, generally 300 to 600. Self-pollinated seed from plants,
which together cover more or less completely the range of
variation, was taken and these cultures raised in the F, generation.
In the F, cultures about 200 plants were grown from each parent,
and 100 to 200 measured. No choice was exercised in connection
with the plants measured. Two or three complete lines of the
culture were taken. The limits of variation in the F, generation
are much smaller. In some cases the cultures apparently bred
true in some one character. A similar procedure was adopted
in the F, generation.

The number of plants used is small and the investigation
cannot pretend to be statistic, but the larger the number of plants
the smaller must be the number of cultures. The number
employed is sufficient to determine whether the culture is
uniform or is splitting within narrow limits. The object in
adopting this procedure is to obtain as many as_ possible

GABRIELLE L. C. HOWARD. 47

of the intermediate forms pure, as the extraction of the various
intermediate homozygotic forms appears to be the easiest and
most conclusive way ot determining the principles underlying the
inheritance. The range of variation of the parents was deter-
mined each year for each character. In 1911, very few of the
parent plants were sufficiently well developed to be measured.
The cultures were transplanted into a field which had been
ereen-manured. Owing to the exceptionally heavy and _ pro-
longed monsoon of that year, the soil conditions were not
favourable to the proper utilization of the green manure, and the
plants only developed well in the high-lying portions of the
field. The F, generation of Type 9 x Type 51 and the crosses of
this back on to the parents were normally developed, but most
of the parent cultures and the F, generation of the other four
crosses were very uneven. This explains why the measurements
given for the F, generation are of a later date than those of the
F,. In all these cases the crosses had to be re-made and the
work repeated. Similarly, in 1913, as there were a large number
of cultures to be grown, some of the lines of the parents were
pushed as near the hedge as appeared safe. Unfortunately,
the influence of the hedge had been under-estimated, and
although the last line was twenty feet from the hedge, the plants
developed very slowly, and were so stunted that they had to be
rejected. These accidents make the number of measurements
carried out on the parents smaller than is desirable.

1. TIME OF FLOWERING.

The date of opening of the first flower on each plant was
taken as the criterion of this character. All the plants were
numbered and the cultures were examined daily at approxi-
mately the same time in the morning. The climate of India
is very well adapted for all such investigations, as the days are
almost invariably sunny, and irregularities due to the periods of
cloud and low temperature consequently negligible. The results
for the three crosses, Type 9 x Type 51, Type 16 « Type 35 and

48 STUDIES IN INDIAN TOBACCOS.

Type 23 x Type 38, are given in Tables I, II, III. In order to
shorten the tables the data are given in three-day periods. The
date of flowering of the parents under the same conditions was
determined each year.

The most interesting results are those for the cross, Type 9
x Type 51. The parents vary oreatly in time of flowering.
The flowering period of the first generation is intermediate
between the two, but nearer that of Type 9, the earlier parent.
The F, generation shows a wide range, but the flowering periods
of the parents is unfortunately not available for this year, 1911.
It may, however, be noted that the flowering period of the F;
covered two months, a period greater than that covered by both

TABL

Date of Flowering.

De TT, DLE

| Date of - | January.
| Flowering of |__— — — ————— —_—_— -
| Parent. | 4-6 7-9 10-12 13-15 16-18 19-21 22-24 25-27 28-30 31-2]
=! | —————
F, Type 9 x Type 51 (1911) | 1 1 aia ll 3
Loin ||
Type 9 (1912) eve Sop — 5 11 22 27 23 6 1 — -=
Type 51 (1912) .. || | 14» 80. ¢425 8 9
F, Type 9 x Type 51 (1912) | ilies — 2 11 26 Dh el! 3 2 —
F, Type 9 x Type 51 (1912) | |
737 | Feb. 8th | — _— — 3 16 32 34 15 14 Le
738 | Feb. 8th | 1 12 hl 26 25 31 26 8 7 2
736 | Feb. 10th | 1 — 1 5 10 24 35 19 23 9
740 | Feb. 13th | - — 25 54 20 21 6
132) | Heb. Ith) — — 7 28 32 22 18
694 | Feb. 15th — 5 14 12 26 21
167 | Feb. 16th _— ] — 2 2 11 24 14 9 14
| Seog as 4
Type 9 (1918) 2)” Fe | 7am: 8 2 3
Type 51 (1913) : Beall | — — -— —
F, Type 9 x Type 51 (1913) |
738 plan. oul — — J 6 12 19 32 30 26
738—19 | Jan. 11th 5 19 36 39 38 28 8
738—81 | Jan. 15th — — 1 6 34 57 50 19 6 4
738—58 | Jan. 16th 7 29 23 25 21
738—96 | Jan. 24th |. —= — — _- 1 — 10 10 18 15
738—76 | Feb. 2nd — — — — as 2 5 15 18
694—1 | Jan. 25th — — — — 7 iy) 23

694—103) Jan. 30th — — _
694—10 | Feb. 3rd — — — —
694—23 | Feb. 4th — — — —— — — — —
694—38 — — — = — — — —

GABRIELLE L. C. HOWARD. 49

parents either in 1912 or 1913. Seven cultures were grown in
the F, generation and here a distinct difference can be seen in the
limits of variation of the individual cultures. Three, namely 738,736
and 167, have a period equal in length to both parents combined,
while 740, 132 and 694 more or less resemble Type 51. Cultures
representative of each of these groups, namely 738 and 694, were
chosen for future experiment, and six plants of the former and
five of the latter were grown in the F,. It will be seen that the
cultures grown from 738 have again segregated. Two, 758-15
and 738-96, cover approximately the period of the two parents.
One, 738-81, is distinctly earlier than the early parent, and the
others vary over different ranges. The culture 694 has, on the

I.

ype 9 X Type 5I.

February. March. | Total
peers = eee |} No. of
3-5 6-8 9-1] 12-14 15-17 18-20 21-23 24-26 27-1M. 2-4 5-7 8-10 11-13 14-16 17-19] Plants.

30 BO 76 74 132 86 64 50 33 15 ili7/ 12 1 4 2 690
a = = — == —- 95
a = 1 as — - = — — — — — 87
= oY = = = = = — 79
1 ase = = — 126
2 — 1 — — = 152
5 5 — = — = — — — = = 137
3- 5 — 1 1 —_ 136
vi 1 2 117
7 ‘i 4 3 1 — — — — == = — 100
6 9 3 4 1 100
2 2 == a= — 37
3 8 5 9 3 3 1 1 3 oo 1 — — — — 37
8 16 12 2, 1 3 — — — — 178
——— ll 177
g == =e ae $e = =a = _— —_ 180
5 U7 11 5 2 — 175
6 24 26 23 a vi 6 1 2 176
3 31 24 19 8 7 5 2 — — 179
9 40 17 13 7 3 — 2 1 — — 179
2 6 10 18 12 18 18 20 9 18 5 2 138
3 8 16 15 19 20 30 31 12 1l 4 4 — _—- 173

f _— 1 17 18 24 343 33 21 24 14 4 4 — il 194

7 2 12 10 1l 21 20 28 22 21 af 6 2 174

TABL

Date of Flowering

Date of January.
Flowering of) =e .
Parent. | 4-6 7-9 10-12 13-15 16-18 19-21 22-24 25-27 28-30 3l=s
|
Type 16 (1913) we: ~- — 1
Type io (19ts)) 2. — — — 1 2 1 2
F, Type 16 x Type 35 (1913) 1 1 11 14
Type 16 (1912) .. a fe gies es 32 16) 20.5 meee gi ae
Type 35 (1912) =i 1 10 18 19 20 1 §
F, Type 16 x Type 35 (1912) — — 6 14 59 75 89. 48 34 2
1912
Type 16 (1913) é ae — - -- _- —- — — — — ;
Type 35 (1913) AF — — — — — — 1 2 1 4
F, Type 16 x Type 35 (191 3)

sy | aeney, ibao, | = 2 (

8 | Jan. 15th — — = — — —
163. Jan. 15th — 3
27 | Jan. 19th == 4 20 15 41 3
190 | Jan. 19th | 1 3 23 3
202 | Jan. 21st = —= = = = — 5 10 22 1
251 | Jan. 21st = — — — 8 1
35 | Jan. 2Zard — —— — = _

9 | Jan. 24th = == == — = — — — 3

231 | Jan. 24th == = = — 2

142 | Jan. 27th — — — — — = .s — 2
200 |} Jan. 31st — — — -
TAE

Date of Flowering

Date of | January.
Flowering of | =
Parent. 4-6 7-9 10-12 13-15 16-18 19-21 22-24 25-27 28-30 3m
Type 23 (1913) ar — — — 10 2 9
Type 38 (1913) Bo a OS ae
F, Type 23 x Type 38 (191 3) —_ — a = = = 10 3 12
Type 23 (1912) Ee aca = 1 — 1 24 27 16 6
Type 38 (1912) ee ee Se ee, ee 1 1 ys ae
F, Type 23 x Type 38 (1912) ; — 1 _- 4 6 62 89 77 49
1912
Type 23 (1913) ofa woe, oma = = = ae 10 2 9
Type 38 (1913) —— ae — = — p= = a2 gi3
F, Type 23 x Type 38(191 3)
104 | Jan. 24th == — — — 1 3 24 18 34
155 | Jan. 24th = — — — — — 5 21 42
213 Jan. 2ist =e = — — 9 11 29
159 | Jan. 23rd == — = 9 17 28
Lid) daaee2oeb 4|)— = — — — — 9 10 32
204 | Feb. 2nd —_ — — — — — 1 3 13
6} Jan. 23rd — — — — — — 3 — 10
117 | Jan. 31st — = — — — — 2 3 16
9 | Jan. 26th —_ — — — — — — — _-

al
Type 16 x Type 35.

February. March. Total
3 abe was ae ——_ : No. of
35 6-8 9-11 12-14 15-17 18-20 21-23 24-26 27-1M. 2-4 5-7 810 11-13 14-16 17-19} Plants.
u 10 10 6 1 1 = — 36
+ 8 9 5 2 2 1 — _ ~ 37
14 18 7 2 2 2 1 — — -- - 73
2 1 98
1 3 = = 100
14 1 1 — - 362
7 10 10 6 1 1 -
4 8 9 5 2 2 if — -- 36
37
19 28 39 30 5 6 3 q 1 — 1 144
1 14 21 32 21 27 10 10 5 3 145
29 47 22 14 14 2 3 1 1 — ] 142
32 16 9 8 1 3 — 179
42 29 12 11 9 7 3 179
38 40 27 16 4 180
23 4] 25 17 8 2 2 — 1 145
1 17 26 40 28 16 7 d 2 1 142
19 31 25 13 12 f 3 — 1 114
22 30 19 27 9 8 2 3 130
9 21 32 32 17 13 6 5 4 1 145
3 10 8 19 26 25 23 21 4 3 2 144

Ill.

Type 23 < Type 38.
Cee

February. March. Total
No. of

3-5 =—«6-8 9-11 12-14 15-17 18-20 21-23 24-26 27-IM. 2-4 5-7 810 11-13 14-16 17-19 | Plants.

— l 27
a vi 4 3 1 26

10 13 3 2 — — — — 76

s9

8 — 1 -— 80

4 344

— 1 27

3 7 7 4 3 1 — — 26

21 10 6 1 2 — 1 — 152
29 19 11 4 1 1 2 — — = 159
31 20 18 4 2 2 — — 156
31 20 9 4 2, 2 157
36 20 7 3 5 2 — — - 155
30 26 13 5 4 5 1 _ 1 — — - = 130
23 27 25 14 9 4 3 1 — — — — 131
30 46 25 20 11 3 2 2 — — —- —- 185
9 16 23 20 21 ll 13 7 3 — 1 — — — — 125

5? STUDIES IN INDIAN TOBACCOS.

other hand, given rise only to late plants. The difference
between the cultures derived from 738 and 694 was most marked
in the plot. There is a good deal of difference between the
individual cultures of 694. No. 694-1 has a maximum flowering
period between the two parents, while 23 and 38 are much later
than Type 51, the late parent. It is impossible to say whether
a culture is pure as regards its time of flowering until it has been
proved impossible to produce from it by selection cultures
differing in this character, but 738-19 and 738-81 may possibly
prove constant, as may also some of the cultures of 694.

If we consider the F, cultures asa whole, we find a very
distinct separation as regards the flowering period. In addition,
while one culture has been isolated, which is slightly earlier
than the early parent, there are several which are decidedly
later than Type 51, the late parent. The range of the earliest
flowering culture does not overlap that of the latest.

Tables II and IIT show the same general results for other
crosses, but the separation between the cultures is not so great,
as the flowering period of the parents is not very different in
either case. The extremes in the F, are represented in the cross
Type 16 and Type 35 (Table IL) by cultures 27 and 200, and in
cross Type 23 x Type 38 by cultures 104 and 9. In all three
the two parents in successive years bear a similar relation to
one another.

2, HEIGHT:

The height was measured from the point where the roots
begin to the apex of the stem, which in N. tabacum ends in a
capsule. Three or four flowering branches arise round this
point and help to define it, but there is no difficulty in seeing
either the capsule or its scar at any period in the life of the mature
plant. In order to count the lower leaves, the plants were up-
rooted and the determinations of the height and the number of
leaves were made simultaneously. The height was measured

PLATE XiMi-

9g]

AdAL “NOILVYHNAD

GQNOOHS

GABRIELLE L. C. HOWARD. 53

to the nearest centimetre, but in order to reduce the number of
data and to remove accidental inequalities, the heights are given
in classes differing by five centimetres in Tables IV, V, and
VI.

Height is one of the few characters which in the F, genera-
tion is not strictly intermediate between the two parents, and
the results in the five crosses were not the same. The F, hybrid
between Type 9 (a dwarf form) and Type 51 (a very tall form)
was nearly as tall as the taller parent. The actual mean
measurements were, in 1910, Type 9—68 cm., F,—140 cm.,
Type 51—165 cm., and in 1912, Type 9—79.9 cm., F,—158.0 cm.
and Type 51—195.4 em. In the cross Type 16 x Type 35, in
which the respective heights were, Type 16—106.3 cm.,
F,—93.7 cm. and Type 35—86.6 cm., the height is inter-
mediate. In the other three crosses the F, was taller than
either parent. This increase in height was slight in the case of
Type 23 x Type 38, but very marked in the other two crosses.
The actual measurements were Type 2, average height, 155 em.,
Type 3, average height, 152 cm., F, 200 cm., Type 51,
178cm., F, Type 2 x Type 51, 195 cm. It has been suggested,
in the case of maize,’ where the F, is also not strictly inter-
mediate, that this is due to the increased vigour of the hybrid
plants. This may be the explanation here, but it is difficult
to see why this increase should be so much more marked in
some cases than in others, unless this increase in vigour be
correlated with the number of factors in which the parents
differ. |

The F, generation in the cross Type 9 x Type 51 presented
no striking features. A continuous series was formed within
the limits of the parents.

The F, generation of the cross between Type 16 and
Type 35 was absolutely different. These two parents do
not differ very greatly in height, the average height of
Type 35 in 1912 was 94.1 cm., with a range of 60 to

1 Hast, l. c.

4

54 STUDIES IN INDIAN TOBACCOS.

110 em., that of Type 16, 125.2 cm., with a range of 110 to
140 cm. The heights in the F, generation varied from 48 cm.
to 215 cm., and the plot presented an extraordinary mixture
of dwarf and tall forms. Plate XIII shows some of the F, forms,
photographed at an equal distance from the camera. A large
number of plants were self-pollinated and twelve cultures from
these were raised in the F,, generation. These cultures could be
easily divided by eye into three groups, those in which all the
plants were short, that is, no taller than either parent, Nos. 251,

TABLE
Height.
i ee
Height Centimetres.
ain |
Parent.| 40 45 50 55 60 65 70 75 80 85 90 95 100 105 110 115 120
Type 9 L910 Mean 68 - — ne
Type 51 1910 —- Se
F, Type 9 x Type 51 1910 —- — —- — =. | Sn
F, Type 9 x Type 51 1911 — — yo 13 14 16 25. 41 42.44 56 49 57 (av 41 32 48
Type 51x F, Type 9x Type 51
1911] —_—- — 3) 1 2 3 2 Soe
Type 9 1912 1 3 136 8.13! 2- 2. re
Type 51 1912 Se = : a.

F, Type 9 X Type 51 1912 = . 2
F, Type 9 x Type 51

738—1912 Sotelo OS Se AOS 21 ALT Ree oli) Soke 9 1

737—1912 114 - : - F 8 .6 2.7 1)l1 38 Soe

694—1912 137 - ima 1 .2 3 3

167—1912 145 | — - 1 2g 2 — Te

740—1912 157 © : Se eae iS SS Se

736—1912 160 — a

132—1912 ? Sh pa == _ =|

Type 9 1913 a be: eee ee ee ee
Type 51 1913 eee SS SS SS i
F, Type 9 x Type 51 |
738— 96—1913 81 9 5 13 29.56 39 23 4— 1 = = => = See

738— 76—1913 36. — 1 2 6 16 26 30 -15 24 Il 12. Osea 1

738— 15—1913 86 5G 179302688 24 7.0 4A 3 1 =) See

738— 19—1913 ? — © 9 28 36 30 12 10 14 8 4&4 2 —) ee

738— S81—1913 9] — = 4 1 8 4 ¥8 28 28 23 24 28 14.) 4 525

738— 58—1913 123 : 3 9 10 23 35 28 29. 16 Iie

694— _ 1—1913 ? font eer byes Be. Beg $= 4qzeyes.. 7 gory 6 eee
694—103—1913 ? oe ee eS ee 2 OB 6) ee

694— 10—1913 9 ee 5) . ee

694— 23—1913 ? 2 : . 2 : . a _— —

Mean height Type 9—1912, 79:9 em .
Type 51—1912, 195°4 em,
F, Type 9 * Type 51—1912, 158 em.
F, culture 740—1912, 195:2 em.

EV:

~

GABRIELLE L. C. HOWARD. 55

200, 142 and 190 ; those in which all the plants were tall, cultures
35, 15, 9, 163; and cultures in which both short and tall
plants occurred, Nos. 202, 27, 231 and 8. In the last four
cultures it was quite easy to separate the tall and short plants by
eye, and although neither class was uniform, the short and tall
plants together did not form a perfect series. Among the
cultures with short plants, two have apparently bred true,
No. 251 and No. 190. In 1912, No. 251, with a height of 48 cm. ,

was the shortest plant in the F, generation ; in 1913, this gave

Type 9 X Type 51.

125 130 135 140

Centimetres.

145 150 155 160 165 170 175 180 185 190 195 200 205 210 215 220 25

Total
No. of

235 240 Plants.

Mean 165

— Mean 140 — —- — —
Zielona. Lb 17 (i 5 3 —- 2 — ~ —~— — 648
beOMeSes (Se) LO) 5) Ol 5) 3 5 2 — 1— — — 107
a — 41
=] Se a 1‘ 1 1 3 do (Gee EE Gy il 1 — 39
ae 1 2-9 DO 1 3 1 5 2 — 38
SS SS See ; = == 143
Gee 0 Oe Bw dee De ond Ce 1 ee, ee 128
NO <7 ae 88 Gee oe 2 == 2 93
Geen De Vly LO” “8) 22> 10 7 aa. Ole eo 1 — — — — 94
= =|] =| |S == 1 3 1 peel vere Semel Ce Gm Omen, econ 5 eOlme asO oN 2) ie 1 136
De ee Oe Se or va 4 GT OM Ore Sr 0) 2! Ss bi Te by 3. 1 2 1 — — 126
— 1 Geeta soe Mal SLO Om SO eSre ei e < 42 OG 1 1 — 1 1 — 115
. 2. 29
S| — = 1 eo faye a Oy e(ie For hit 33
—- — — = = 172
SS SSS SSS —- —- Fe rr er 164
— — —- — 172
—- — — — —- — —_— —_— — 90
—_—- — —- — 173
1—- —~ ~—~ — — — —_- — — 168
15 5 4 — 170
LG oe Che fees 1 6 — —- —- —- —- —- — — 109
Bio 35 J1 1 4 1 —_ — - 165
eis lS V7 20) 528) 21 4, 18 8 5 1 1—- —- —- —- —- —- —- —- 177

Mean height

Type 9—1913, 57:5 cm.

Type 51—1913, 188°3 cm.
F, culture 738—96—1913, 60-1 em.
F,, culture 738—58—1913, 98-7 cm.

Height.
Heteut Centimetres.
o
Parent. | 35 40 45 50 55 60 65 70 75 80 85 90 95 100 105 110
Type 35 1911 | — = SS SS a i a SO Mean 8%) ee
Type 16 . 1911 —- —-— F— Fe Fe ere re —- —- —
F, Type 16 x Type 35 1911 | — FS ee ae a a a a aS a ae eS ee ee
Type 35 1912 | 1 = .— (1 8 Og SIA ee
Type 16 1912 | —- — — — 1
F, Type 35 x Type 16 1912 | — =—- Tf — 3 2 TF 6 20 14 19 239 °36) 25sec 2S
Type 35 1913 | —- — — — 2 4 13 10°94 =
Type 16 1913 | I ‘be 3a) Sa
F, Type 16 x Type 35 1913 | 2 9 21 224 bo Omeene
F, Type 16 x Type 35 |
251—1913 | 48 LOND) 749 eeZOm Sh aeo — —= aa
200—1913 | 73 — — J] — 2 3 11 16 28 19 36 18 G6 “32a
142—1913 | 81 4 9 11 28 #33 31 11 165 2
202—1913 | 83 6 6 18 27 26 20 14 4° 8) ee
27—1913 88 — 1 9 12 12 2) 17-15 19-15-13) <6) see
231—1913 90 — J '2 56 ‘6 7 10 12, 16 12) 16 LO) (SCR
8—I1913 | 116 - 1— — 3 8 138 24 20 18 Il Is
190—1913 131 1 1— 6 15 28 58 2b 2%
35—1913 | 180 — 1
9—1913 182 ee
163—1913 195 aS ee eS SE ee
15—1913 204 —_— — 1... 1 ee
Mean height Type 16—1912, 125-2 cm.
Type 35—1912, 94:1 em.
TABLE
Height.
Parent. Centimetres.
Cm. 65 70 75 80 85 90 95 100 105 10g
Type 23 OU a
Type 38 1911 == —
F, Type 23 x Type 38 LOT —— 3+ ee
Type 23 1912 — — 1 2 5 4
Type 38 1912 —- — — — — — — — 1 4 6
F, Type 23 x Type 38 1912 — — 1 8 18 12 30 21 38 30 28
Type 23 1913 — 2 4
Type 38 1913 i FM
F, Type 23 x Type 38 1913 — \—Saae
F, Type 23 x Type 38 213—1913 117 — 1— 6 6 3 9 Is “Toe
117—1913 102-5 1 1 2 1:10 11. 1) 1G
155—1913 108 2 1 6 2-7 12 10 “eee
111—1913 128 — — — — 1 2 1 65 7 11
159—1913 139 —- — — — — — 1 2 2 7 tt
104—1913 151 —- —- -—- -—- e- Fr r e lU hr S
9—1913 155 —_- — —- —- FF
6—1913 166-5 |} — — — — = —- —- —- —- — =
204—1913 — — — — 1 56 4 12 18 20

Mean height Type 23—1912, 119:1 cm.
Type 38—1912, 133-0 cm.

[ype 16 X Type 35.
Centimetres. Total
o . : No. of
115 120 125 130 135 140 145 150 155 160 165 170 175 180 185 190 195 200 205 210 215: 220 (225 230 | Plants.
fean 12| Mean 135 — — — — — _ = ae
[=a a > 45
7 3 a lize ul 1 - 43
19 14° 7 4 7 ty al 9 8 10 8 8 8 6 9 6) 14 1 1 1— — — 356
= = - 33
4 — — = 32
= 67
— - 144
== = 145
== == SS = 144
1 jeoio ld 5 6 6 et: 3 1 1 - 186
6 6 5 6a) 25, 2 2 1 1 - - 186
2 — 6 6 3 7 5 4 4 2 1 146
2 3 2 1 — 6 3 74 9 4 3 1 148
12 4 4 181
= eS 2 4 9 6 Sele i ass 19 10 6 5 2 1 — = 1 — — 140
= 1 2 1 1 6 7 Oe TAG SG) a3 4 5 2 1 102
— Sn 4: i 2 10 10 14 20 23 19 18 6 5 2, 1 138
5 1 1 Ze 5 6 li SF Il lO) IE ARGS Gee ala! 6 8 5 7 — — 1—- — — 142
Mean height Type 16—1913, 106°3 cm.
Type 35—1913, 86:6 cm.
F, Type 16 x Type 35—1913, 93:7 cm.
F, culture 251—1913. 50°8 cm.
F, culture 190—1913, 102-1 cm.
v I.
ype 23 x Type 38.
Centimetres. Total
No. of
120 125 130 135 140 145 150 155 160 165 170 175 180 185 190 195 200 205) Plants.
— Mean 133 — — — =—- —- S—- S—- —- l
— Mean 130 — — — — - =
= Mean 132 — — —
ma 65 il 1 2 25
mo 4 1 3 5 3 2 — 1 - — 35
mae ro) °621)6«(CdI8- ods «61012 5 8 3 2 4 1 1 331
mo. 4 1 1 23
— 1 — 4 6 2 & 1 - 19
—_ — — — 5 bP k2 fl 5 7 3 2 iL ~ 47
mm ii 9 12 9 6) 4 1 3 1 1 134
ma 14 10 9 7 — 2 3. — 1 1—- — — 175
s 14 9 8 4 6 6 2, ] 1 2 150
mela 18: IL 12 18 8 7 6 1 2 — 1 144
mei? 17 18 LT Li 9 9 6 5 2 1 3 150
_— — 1 2 6 5 3 PL Px0) 88} eee ty 1183 6 6 2 1 I atb3
me ft — 2 6 Se eG be els pals Grated 6 3 —- — — 124
— — 2 6 3 5 Opti Onan als. 9 5 LO 6 6 3 5 118
19 11 3 5 2 1 — —- — —- —- —-— ar ll 117
% Mean beight Type 23—1913, 122-9 em.

Type 38—1913, 142-3 cm.
F, Type 23 x Type 38—1913, 154°6 cm.

oO

5 STUDIES IN INDIAN TOBACCOS.

rise to a culture which appeared absolutely uniform in the field.
The average height of the culture was 50.8 cm., much less than
that of Type 35, and the limits of variation (35 to 65 cm.) do
not overlap those of Type 35. The individual heights of the
plants, if graphically represented, give a curve which resembles
greatly the ordinary frequency curve of a uniform culture.
This fact, combined with the appearance of the culture in the
field and the small range of its variation, makes it probable that
251 represents a culture constant as regards height. We have
thus the formation in the F, of a new form much shorter than
either parent. The factor or combination of factors which
is represented by this average height of about 50 cm. may be
present in both parents, or only partly in both, but the fact that
this combination has been isolated shows that the height of
Type 16 cannot be composed of small factors superposed on those
possessed by Type 35. Even if we suppose that the combination
of factors denoted by a height of 50 cm. is common to both
parents, all the factors which go to form the excess of height
over 50 cm. in Type 35 and Type 16 must be different. This
gives the explanation of the very tall plants in the F, and F,
generations. Since so large a proportion of the height in the
parents is due to different gametes, the combined effect of such
gametes would be additive.

There is no definite evidence as to the number of the
factors involved, but from the appearance of some _ of
the F, cultures they would seem to be few in number.
Culture 190 is also probably uniform and a reproduction of
Type 16, the mean height in the one case being 102.1 cm., in the
other 106.3 cm. The range of variation is greater, but this
may be due to the larger number of plants measured. From
their appearance in the field and the curves given by their height
it is probable that the cultures 142 and 200 are not uniform,
but represent heterozygotes between two nearly allied combina-
tions. After 251, the most interesting culture is 202. Here
we have a distinct break between the tall and the short plants,

‘i
we

; 4

er ee

ee ee ae

GABRIELLE L. C. HOWARD. 59

and the tall plants form exactly one quarter of the whole. The
large range of variation in the tall forms indicates that the ratio
is not a simple 3:1. Cultures 27 and 231 show a similar
result, an accumulation below 95 cm., a total break, or very
few plants about 100 and a rise beyond this. These three
cultures were derived from three F, plants of very similar height,
83, 88 and 90 cm. The curve formed by combining these
cultures is shown in Plate XIV. Culture 8 is very similar to
these three cultures, but the curve appears to be shifted to a
higher value. The cultures 9, 35 and 163 are all similar
and seem to point to at least two homozygotic combinations at
about 120 and 170 to 180 cms., with possible intermediates.
The combined curves from these three cultures is given in
Plate XIV; the individua) cultures all agree with this. A
large number of plants have been self-pollinated and the F;:
should give sufficient uniform cultures to identify all the factors.
The values obtained in some of the cultures seem somewhat
high for a rudimentary height of 5|0cm. common to both parents.
The fact that in all the cultures grown in the F, and F,
nothing shorter than 50 cm. has been thrown out would point
to nothing smaller than this combination existing in either; yet
if this factor or combination. be common to both, the mean of the
highest combination possible can only be (85—50) + (106—50) + 50
cm. The limits of the tallest homozygotic combination
should also throw light on another interesting point. Since
every plant must have height, it is reasonable to suppose that all
types possess a common factor which represents the smallest
height which the tobacco plant can possess. This, either by the
superposition of numerous small factors or by combination with
various independent factors, gives the height of the different
plants. The first supposition of small superposed factors is
entirely disproved by the cross (T'ype 16 « Type 35), in which
plants have been produced almost equivalent to the combined
height of both parents, showing that the constituents of the
height of both parents must be almost all different. Should a

60 STUDIES IN INDIAN TOBACGOS.

homozygotic combination be produced equal or greater than
the sum of both parents, this may be explained by the
greater vigour produced by hybridization or by supposing a
different rudimentary height in different plants, or that the
factor giving height exists in all plants but has no definite
external expression. Both these hypotheses, however, would
introduce many difficulties.

The results in the cross Type 9 « Type 51 (Table IV) need
not be considered in such detail. The range of variation in the
F, was not greater than the range of the parents combined. In
the F, generation, the eight cultures showed a great difference
in the range of variation, but only one appeared to be uniform.
Culture 740 (Plate XV) resembled the parent, Type 51, in
every respect, both in the limits of variation and in the average
height. The average height in the case of 740 was 195.2 cm.,
and the limits of variation 155 to 240 cm.; in the case of
Type 51 the average was 195.4 cm. and the range 160 to
230 cm. Both cultures were grown side by side in the
field and there was no difference in their appearance. A
form resembling one of the parents was therefore isolated
in the F; The difference in the range of variation in
the F, and F, generations distinctly indicates a segregation,
as well as the probable occurrence of homozygotic combinations
representing heights intermediate between the two parents.
One such was isolated in the F, generation (738-58), as well as
a culture resembling Type 9 (738-96). From their appearance in
the field, their range of variation and the form of the curve
obtained by graphic representation, both these cultures appear
to be homozygotic. The average value of the height in 738-96
is 60.1 cm., that of 738-58 is 98.7 cm. The height of culture
738-96 is probably the same as that of Type 9 (average height
57.5 em.) ; the other forms a novum. The remaining cultures
of 738 are not uniform and contain these two combinations with
possibly an intermediate. The F, cultures of 694 are also not
homogeneous, but their limits of variation are very different to

PLATE XV

No. 740.

TYPE +O

THIRD GENERATION. Type 9 x

GABRIELLE L. C. HOWARD. 61

those of the cultures obtained from 738 and indicate the occur-
rence of a number of intermediates. This would point to
factors of small value. From the fact that no plants shorter than
Type 9 have been found in all the generations, it would appear
as if Type 51 must contain the factors which go to build up
the height of Type 9.

In the cross between Type 23 and Type 38 the matter is
complicated by the fact that the F, is taller than either parent.
The F, again shows a greater range of variation than the com-
bined range of the parents and the large number of plants
shorter than Type 23, the shortest parent, is especially notice-
able. Inthe F, cultures the limits of variation are very different,
ef. cultures 204 and 6. The occurrence of plants shorter than
Type 23 is again marked. No cultures uniform in height have
yet been obtained from this cross. Distinct segregation is,
however, apparent, and there are indications that Type 23 and
Type 38 contain certain factors in common which give a height
somewhere about 80-85 cm. and that the other factors involved
in the height of both parents are different, giving rise by
combination to plants taller than either parent.

3, fHE NUMBER OF LEAVES PER PLANT.

This character is somewhat unsatisfactory on account of
the amount of variation in each pure type, in proportion to the
total variation possible between the types. The smallest number
of leaves found in any kind is nineteen, the greatest thirty-four.
Table VII shows the approximate number of leaves in each
type—the average number of leaves in ten plants. Want
of time prevented the measurement of a larger number of
plants. It will be seen that every possible number of leaves
present and all occur with approximately equal frequency.

The total number of leaves on the main stem was measured
in the following manner. The plants were uprooted, the large
roots cut off short, and the base of the plant well washed. It was
then possible to count all the leaf scars as well as the living

62 STUDIES IN INDIAN TOBACCOS.

Taste VII.

Average Number of Leaves in the Types of N. tabacum.

|
No. of | No. of No. of

Leaves. Leaves. Leaves.

Type 41 19°5 Type 17 23 Type 37 26
» 40 20 st 44 23 #. i 26
23 20 | oy 45 23 | ve 10 27

42 20 5 24 SY!) 27

43 20°5 | 8. D7 24 eats 27

34 20-5 | lois 24 | LO 28

35 Ai | = 7 24 ae 046 28

36 21 sis 24 <) Bil 29

47 22 a) = 20 25 - 4 29

48 22 a= RAD 25 OG 30

13 22 pat e153 95:5 Se Jol 31

a 6 22 ae 30 26 a 5 3353
eee 22 | 2 26 | 38 34.

leaves. Enumeration at the base, where the scars are close
together,'is facilitated by marking the point of commencement
with a knife, and each subsequent scar by sticking into it the
point of a lead pencil. Care is necessary not to mistake con-
tractions in the main root for leaf scars, but with a little practice
this can be done fairly easily, except in the case of a few types.
The determination of the last leaf on the axis is more diffi-
cult. In many types of tobacco, the leaves at the top of the
stem are carried up the branches they subtend and therefore
do not appear to be on the main stem. In such cases, however,
there is generally a faint line running from the leaf down to the
main stem. The method of enumeration adopted in these
investigations has an advantage over previous methods used in
that it represents the true physiological activity of the plant
and not an arbitrary number.

As regards the effect of environment on the number of leaves,
no special investigations have been undertaken, but the fact
that the average value of this character and the range of varia-
tion in the parent forms is practically constant in different
years, wouldindicate that changes in a normalenvironment have
little effect. This agrees with the conclusion of Hayes that
the number of leaves per plant was little affected, unless the

GABRIELLE L. C. HOWARD. 63

conditions were so unfavourable as to greatly stunt or dwarf the
growth of the plant. It is possible, however, that the length
of the growth period may have some influence. As stated on
page 38, in 1913 a portion of the parent cultures were grown
too close to a hedge and developed very slowly in consequence.
These plants, when they did attain maturity, all had an
abnormally large number of leaves. For instance, Type 51,
instead of possessing the normal number of twenty-seven to
thirty-one leaves, had thirty-one to thirty-five leaves, and a
similar increase was noticeable in Type 35. Moreover, in 1913, a
late season, the average values of all the parents was about one
leaf greater than in 1912. Again, in 1911, another late
season, the average value of Type 51 agrees with that of 1913.
In the absence of further evidence this can only be put forward
tentatively. The ordinary fluctuations of season have no
appreciable effect. The number of leaves per plant apart from
these abnormal individuals appears to be a very definite inherit-
able character.

Much attention has been paid, in the course of these
investigations, to the question of a correlation between the
the number of leaves and the height of the plant ; but it is quite
clear that each can be inherited independently of the other.
Some of the shortest plants have the largest number of leaves.
Take for instance culture 251 (Tables V and [X) in the F, genera-
tion of Type 16 x Type 35. This culture is uniform as regards
height (av. 50.7 cm.) and uniform as regards leaves (av. 30),
that is, it has a greater average number of leaves than Type 51,
of which the mean height is 195.5 cm. Culture 190 of the
same cross has an average height of 102.1 em. and the average
number of leaves is thirty. Such instances might be
multiplied indefinitely. Cases also occur in which the culture is
uniform as regards height, but not as regards number of leaves
per plant (for example, 740 in the F, generation of Type 9 x
Type 51) and vice versa. The fact that, while the height varies
greatly with the season, the number of leaves remains

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GABRIELLE L. C. HOWARD. 65

practically constant is further evidence that, within the same
type, height and number of leaves are not correlated. This
is a somewhat surprising result. It coincides, however, with
the conclusion of Hayes, that the correlation between height
and number of leaves was less than +.6.

The data as regards this character in the cross Type 9 « Type
51 are given in Table VIII. The F, generation was intermediate
between the parents. The F, generation has a range far outside
those of the parents, one of the plants having as many as
forty leaves. That this is not due to accidental circumstances
is shown by the reappearance of this phenomenon in one of the
cultures in the F, generation, where the number of leaves varies
from twenty-eight to forty. The eight cultures of the F,
differed greatly in their range of variation; none were
uniform. Two of these were continued in the F, generation,
i.e., Nos. 738 and 694. From 738 two cultures were obtained,
Nos. 738-81 and 738-19, which from their range of variation
and the form of the curve appear to be uniform. The first
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of seventeen to twenty-two. The latter culture, 738-19, is
a replica of Type 9 as regards number of leaves. In the culture,
derived from 694 only one can possibly be uniform, No. 694-1,
but as the range of variation is somewhat large it may have
originated from a heterozygote between two forms differing
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of intermediate homozygotic combinations. Thus in the F,
generation there have been isolated a culture (probably unitorm)
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GABRIELLE L. C. HOWARD. O7

the larger number of leaves. The limits of the IF, generation
are as great as those of the parents combined, but no greater.
The curve obtained from its graphic representation (Plate XLV)
points to distinct segregation, as do also the results obtained trom
the F, cultures. The foliowing cultures appear to be uniform,
Nos. 251 and 190, with an average number of thirty leaves: per
plant, and 202, with an average of twenty-six. This is another
instance of the production of a plant possessing a number of leaves
intermediate between those of the parents and breeding true.

Table X gives the results in the cross Type 23 » Type 58,
which are very similar to those in Table LX. The IF, generation
again approaches the parent with the greatest number of leaves.
Distinct segregation occurs in the F,, but no cultures breeding
true were found.

As regards the principles underlying the inheritance of the
number of leaves per plant, there seems no doubt that this
is a definite inheritable character, and that on hybridization
segregation occurs with the formation of new homozygotic
combinations. In dealing with the mode of inheritance we are
confronted with the same difficulty as in the case of the height,
namely, the impossibility of a plant without leaves. Hayes
supposes a basal condition of x leaves which occurs in all tobacco
plants with a number of small interchangeable factors, each ot
which may represent two leaves. Thus the same number of
leaves per plant may in reality have been caused by different
combinations of characters, ie., twenty-two leaves may be
x + AAbb or x + BBaa.

There is nothing in the results obtained in these investi-
gations to contradict this hypothesis of a basal number of leaves
combined with interchangeable factors. The results in the
cross Type 9 x Type 51 appear to be a direct confirmation, But
the form of the curves obtained in the F, generations point
distinctly to the occurrence of factors of different values. — If all
the factors were more or less equal in value the form of the curve
would be more uniform, and would not show such distinct modes.

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GABRIELLE L. C. HOWARD. 69

4. THE ARRANGEMENT OF THE LEAVES ON
THE STEMI.

Besides the height and the number of leaves, the habit of
growth of a tobacco plant depends on the arrangement of the
leaves on the stem and on the inclination of the leaves, i.e., on
their tendency to assume an upright or a drooping position. An
economically profitable plant for India should possess a large
number of upright leaves on a stem of medium height, the
majority of the leaves being borne towards the base of the stem.
The inheritance of the height of stem and of the number of leaves
has been dealt with in sections 2 and 3. No definite results can
be given as to the position of the leaves, but in cross Type 16 ~
Type 35, in which a form with drooping leaves was crossed with
one in which the leaves are upright, distinct segregation with
the formation of intermediates was observed. Similarly in
culture 694-23 (Plate XVI) the upright character of the leaves
of Type 51 was reproduced.

The results concerning the mode of inheritance of the
arrangement of the leaves are not very complete, but some
interesting observations were made. With regard to this
character, the plants can be divided into three groups: (1)
those which carry all their significant leaves at the base of
the stem and in which the leaves consequently lie on the
ground ; (2) those in which the majority of the significant leaves
are borne near the base, the rest up the stem, and (3) those in
which the leaves are borne at equal intervals up the stem.
This latter condition, which may be termed the “ ladder type,”
is well seen in Type 51 and its offspring, No. 740 (Plate XV).
This particular arrangement does not depend on the number of
leaves or on the height of the plant. Short and tall plants alike
are found of this type. Type 16 is an example of ashort plant,
with equal short internodes ; culture No. 740 of one with equal,
long internodes. In every case plants selected with this
arrangement of the leaves have bred true as regards this

5

70 STUDIES IN INDIAN TOBACCOS.

character. The F, cultures 190, 35, 15, 163 and 9, of Type
16 x Type 35 exemplified this.

The first arrangement in which all the leaves are carried
at the base of the plant, and of which Type 9 is a good
example, also does not depend on the number of leaves. In
the F, culture 738-96, which bred true to this arrangement,
the number of leaves varied from twenty-one to thirty-two,
and apparently the plant found no difficulty in crowding
these thirty-two leaves at the extreme base. The leaves were
fairly narrow, otherwise the lower ones would have rapidly
died from want of light and air. This rosette arrangement
has never been met with in a very tall plant. It appears in
plants of medium height (Plate X VI, 694-103-120), but the leaves
are not quite so concentrated. The dwarf rosette forms have
been bred true, but as yet no taller ones.

The second group, those plants in which the majority of
the leaves are concentrated at the base, is a very variable one
and contains a large number of different types which breed
true. The difference in these types does not depend on the
height, but appears to be influenced by large differences in the
number of leaves. The same arrangement with twenty leaves
and thirty-three leaves must appear dissimilar. The F,
generation between Type 9 (rosette) and Type 51 (ladder)
was of this type. It is interesting to note that in the F,
generation besides rosette and ladder types, two different forms
belonging to the second of the above groups have bred true.
One with a few leaves at the base but which otherwise
resembles the ladder type is shown on Plate XVI, 694-38-72.
Another form with a great concentration of leaves at the base,
694-23-1, is shown on the same plate. The latter is an excellent
type for economic purposes and it is hoped to obtain a useful
tobacco from it. The fact that this culture, with such a
profitable habit of growth for economic purposes, has been
bred from two such useless forms as Type 9 and Type 51 is
important.

XVI.

Piece

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GABRIELLE L. C. HOWARD. fil

5. THE INSERTION OF THE LEAVES ON THE STEM.

In N. tabacum the leaves are amplexicaul and in many cases
the lamina is decurrent. The length of this decurrent portion
varies from .5 to 7 or 8 centimetres. Forms with non-decurrent
leaves are also found. As this is one of the few characters in N.
tabacum in which total absence of the character is possible, it
has been investigated in some detail.

Particular attention was paid to the possibility of corre-
lation between this and other characters such as the length
of the internodes or the configuration of the base of the leaf,
but as far as can be ascertained there is no relation between this
character and any other. Table XI shows the lengths of the
decurrent portion of the lamina of three leaves and the
lengths of the three corresponding internodes in the F, culture’
15 from the cross Type 16 « Type 35. It will be seen that there
in no connection between the length of the internode and the
length of the decurrent lamina. Two of the plants are figured
on Plate XVII. Shortness of internode appears to have
no effect in diminishing the length of the decurrent portion.
Should the internode be shorter than the decurrent lamina,
the latter simply grows on into the next internode past the
next leaf (see Plate XVII, 35-3). At the base of the
plant where the leaves are very concentrated it is true that
this character cannot obtain its full expression, but there are
always longer internodes above on which the real length can be
observed. A limiting case might arise in which the number of
leaves being very large, the plant very short and the decurrent
portion very long, the latter was always stopped by the lamina
of the next leaf, but such cases must be exceedingly rare, and
none have been observed up to the present.

It also seems possible that the configuration of the leaf base
might influence this character and that the lamina of a leaf
with large auricles or a broad base would be ;more liable to
grow down the stem. This has, however, not been found to
be the case. No. 159-44, Plate XVII, shows a petiolate leaf

Type 16 x Type 35, culture 15.

GF

TABLE

Length of Internode and of the Decurrent Portion of the Lamina.

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6.6

63

PLATE XV

Te ZoxLoo
I5S9 - 58

T. 23 xT.38

9 =a

T.23xT7.38
iIs9 —44

T.35 xT.I6

1S = 3 ilGxoo

T.35 xT. 16
IS

35-3

DECOR RRENT LEAVES:

GABRIELLE L. C. HOWARD. 13

with a long decurrent portion; No. 159-58 a broad leaf with a
short one; No. 159-11, a leaf with large auricles and a short
prolongation of the lamina.

Further, this character does not appear to be correlated
directly with the width of the leat.

Measurements were taken in two ways, firstly, by measuring
a typical leaf on the living plant; secondly, by measuring
three leaves on the plant when this had been uprooted
for the observations on the height and number of leaves.
The first method has the advantage that information on the
inheritance of this character is available before the seed
plants are chosen, the second is less fatiguing and more
accurate. A large number of cultures have been examined
by both methods and on normal plants either method gives
good results. In this character a large amount of variation
occurs, particularly in those leaves in which the decurrent
portion is long. It is, however, a_ definitely inheritable
character with distinct segregation (see Tables XIII, XV
and XV). The approximate mean measurements of all
the Indian types is given in Table XII (average of ten
plants). It will be seen that the possibilities are numerous.

TEABEE ONL,

Length of the Decurrent Portion of the Lamina in the Types of N. tabacum.

Cm Cm. Cm.

Type 7 — Type 3 135 Type 24 3.0
NS am Le AiG 1.5 t)has oh
ace 1 _ ret 1.6 be BO 3.3
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ioe 20 5 oe iB 2.0 ae ee 3.6
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atl 8 AND 2.5 AR) 4.1
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» 36 i” aa 2.6 | »» 30 4.5
a 8 1.2 eee ot 2.6 | sar 49 5.4
ae 2s Il a8? oe 258 ele Dil 6.0
ie 1.5 a6 4) 2.9 3+ UBS 6.0

74 STUDIES IN INDIAN TOBACCOS.

Table XIII gives the results obtained in the cross Type 9
(non-decurrent) x Type 51 (with an average of 6 cm.). The
I’, was intermediate between the parents and the F, covered the
whole range between non-decurrent forms and those like Type 51.
Several of the plants classed as non-decurrent in the F., however,
gave rise to forms with decurrent laminas later (cultures 738 and
745). This maybe due to the tact that in the early investigations
the number of factors involved was not realized and no measure-
ments of less than .5 centimetres were made. Great differences
were observed in the F, generations, c.t.,'the cultures 738 and
152, but none of the cultures proved uniform. In 1912, unfortu-
nately all plants in which the decurrent portion was shorter
than 5 mm. were again classed as non-decurrent. Cultures
738 and 694 were continued to the F, generation. The results
obtained from culture 738 will be considered first. Two plants
which appeared similar to Type 9, were chosen for future
cultivation, and four plants whose leaves were decurrent to
varying amounts, 5,7, 11 and 15 mm. Of the two cultures with
non-decurrent leaves, one bred true and resembled Type 9.
The other was a heterozygote. A detailed examination of
Type 9 shows that occasionally the method of insertion of the
leaves caused the lamina to be attached to the stem for about
3mm. This makes it almost impossible to distinguish pure
non-decurrent forms from the F, between non-decurrent leaves
and those possessing the smallest factor for a decurrent lamina.
In 1913 most careful measurements of all plants have been
made especially in those cases where the value was below
5mm. Three careful determinations were made in the case
of all the cultures derived from 738, and the mean of these form
the data given in Table XIV. The limits, within which the
splitting takes place, are so small that it is very difficult to obtain
really reliable information as to the extent and number of the
factors, but there is no doubt of the difference between the
cultures. It appears probable that two factors are involved,
one giving alength of about .5 or.7cem.; the other about 1.1 em.,

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76 STUDIES IN INDIAN TOBACCOS.

TABLE

Length of the Decurrent

Length of the Decurrent Portion of the Lamina

Parent : ae
in Millimetres.

mm. 0 ] 2 5 4 D6 7 8 9 10

F, Type 9 x Type 51 745—1912 | Less than 5| 49 — — 1 — 16 1 11 10 J1 14

F, Type 9 x Type 51 738—1912 | Lessthan 5| 46 — — 2 30 2 10 19 — 15 6
KF’, Type 9 x Type 51

738-19—1913 | Less than 4| — — a sill a8 WAS ts 8 7 4

738—76—1913 5 — 1 it 9 12 bale

738-15—1913 7.5 |— — — 2 1 @ 22 28°75 ee

738-81—1913 | 13 oh 2 2 1a

738—58—1913 15 }|—_ — — — — 1

694— 1—1913 | 13 | : =

Type 23 1913 | - - _ 1 1 1 4 3

F', Type 23 x Type 38 155—1913 15 | - = ae

F, Type 16 x Type 35 190—1913 14 {|— « — —— - - 6 2, 12

or there may be two factors of .5 and .7 cm. with a combination
of 1.2 cm. It will be seen from the graphic representations
(Plate XIV) that 738-81 with a mode at 1.1 em. is most probably
uniform, and culture 15 with a mode at .7 cm. possibly so. The
number of plants with non-decurrent leaves in culture 738-19 gives
a ratio, total plants: those with non-decurrent leaves, 15: 1, which
would also indicate the existence of two factors. Further
observations will be made on this point, but on account of the
small size, the difficulties of measurement and interpretation are
very great. Culture 694 must be considered next. This culture
has never given any plants with non-decurrent leaves. Two
cultures appear to be uniform, 694-1 with an average length
of the decurrent portion of 1.8 cm., and 694-23 with an average
of 4.0 cm. The detailed measurements of the culture 694-1
are also given in Table XIV, and show by their range that
this cannot possibly be identical with the combination of factors
in the culture 738. As the parent plant of 694-23 was the
plant with the longest and that of 694-1 the plant with the
shortest decurrent portion in the F; culture 694, it is probable
that 4.0 cm. the mean value of 694-23 represents the
combination of the factor or factors of 694-1 with another or
others. The existence of at least three or four factors which

=~]
~I

GABRIELLE L. C. HOWARD.

ELLY .

Portion of the Lamina.

ee

Length of the Decurrent Portion of the Lamina in Millimetres. Total
No. of
Peeeteoolo 14 5 16. 17 18 19-20 21 929 23 24 25 26 27 28 29 30] Plants.

143

15 9 8 1 3 1 1] — — 1 i :
8 I 2. —— 1 = 146
5 3 2 1 — 1 1 = 105
if 9 5 6 3 1 94
4 3 2 1 - - 101
oe Le 7 8 1 — - - - 96
3 imelveecoe 3 E30 U5 6 i 2, - 100
— 1 2 — Il 3 5s 2 16 7 9 2 6 4 2 i 100
1 8 8 6 5 — 3 — 2 == 44
1 Eye 4: 9 6 2 4 5 ——-— i 3 — 58
Aa 20 9 9 aj) all a — 6 — 2 — 1 ] 108
Ss —

combine to make up the length of the decurrent portion of the
lamina in Type 51 is therefore probable.

The results in cross Type 16 Type 35 (Table XV), are
also interesting. Here both parents have the same average
value, and the range of variation in the F , generation is exactly
similar to that of the parents. The length in the F, generation
varies from .5 cm. to 6.5 cm., far beyond the limits of the
parents. No plants with non-decurrent leaves were found, but

: this may have been due to the comparatively small number
measured. In the third generation there were great differences
among the cultures, and plants with non-decurrent leaves
occurred in three cultures. The numbers in all these cases
(cultures 231, 202, 27) point to a 63: lratio and consequently to
the existence of three factors in these particular cultures. Higher
values were found in the F, than in these cultures indicating
the existence of additional factors in the parents. This supposi-
tion is confirmed by the high values found in the F, culture 163.
The large number of factors involved would explain the absence
of plants with non-decurrent leaves in the F,. The factors which
are responsible for the length of the decurrent portion of the
lamina in Types 16 and 35 respectively would seem to be entirely
different, even though their external expression is identical.

78 STUDIES IN INDIAN TOBACCOS.

Culture 163 probably represents the combination of all the factors
in Type 16 and Type 35. The total variation is no greater than
in Type 51. The average value in this culture is greater than
the combined average values of Type 16« Type 35, but this may
be due to the added vigour due to hybridization. Further
cultures will, it is hoped, put the matter beyond question, and
any explanation must be tentative until these have been
examined. Culture 190 is probably uniform with regard to
this character.

The cross Type 23 « Type 38 (Table XVI) is similar to that
between Type 9 and Type 51, except that neither parent has
non-decurrent leaves. The F, is intermediate and the range of
variation in the F, generation is the same as that covered by both
parents. It would seem therefore as if the same combination
of factors was common to both parents with the addition of extra
factors in one parent. Culture 155 is uniform and like Type 23,
culture 104 is possibly uniform and like Type 38, but this cannot
be assumed safely until further cultures are raised, as the
variability is somewhat great. The parent Type 38 and the
F, generation exhibit very great variation in the same plant.

0. VENATION OF THE LEAVES.

The characters of the lamina itself will now be considered
and of these the venation has proved to be the most constant
and easily measured. A few words of explanation as regards
the choice of the material are necessary, and the following
explanation refers to all measurements on the leaves themselves.
The leaves of a tobacco plant are not all of the same kind, the
lower or ground leaves are very much alike in all the types and
are not characteristic of the variety. They are almost invariably
flat, and have a venation of 60°. This is the case even in Type 9,
in which the upper leaves are twisted by excess of undulation.
These ground leaves pass by a fairly rapid gradation into the
significant leaves, which are typical of the variety, and comprise
most of the leaves on the plant. The significant leaves increase

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8&0 STUDIES IN INDIAN TOBACCOS.

slowly in size from the base and then diminish at the top of
the plant, the diminution increasing very rapidly at the end.
The leaves in the centre of the plant are approximately uniform
in most of the types. Above the significant leaves come a few
very small leaves which vary greatly in shape. In some of the
broad-leaved types, they are small replicas of the large leaves; in
types with lanceolate leaves they are linear; while in many cases
they may be linear even if the significant leaves are not lanceolate.
In Tables XVII, XVIII, and XIX detailed measurements are
given of all the leaves on three plants of each of the parents.
Unfortunately, by the time the plant is mature, the lower leaves
have become damaged. The large number of fairly uniform
leaves in the centre of the plant is, however, evident. Type 9
(Table XVII and Plate XVIII) is an example of an extreme case
in which no two leaves are alike. The best material for all
leaf measurements is undoubtedly to be found in the central
leaves, which are practically uniform, and all the measurements
in this paper have been taken on such typical leaves. It is
impossible to choose any definite leaf (for example, the tenth leat)
suitable to all plants owing to the variations in the number of
leaves per plant, but it is quite easy with practice to pick out the
middle portion of the plant by eye and to select a typical leaf
in this position. The tables show that in this region of the
plant there are a large number of leaves, all of which are suitable.
Measurements carried out on the parents by taking a typical leaf
from a large number of plants have shown that the method is
most satisfactory. The usual practice of obtaining the average
between the bottom, the middle and the top leaf is open to many
objections and would be quite unsuitable for the kinds of tobacco
grown in India. It presupposes a uniform change in the size of
the leaves on the plant, and takes no account of the large number
of uniform leaves in the centre of the plant, which vary in their
effect on the average with the number of leaves per plant, and
the type of the plant. Moreover, the determination of the top leaf
is necessarily arbitrary. The real top leaves are far too small

PEAGE Savill

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THE LEAVES OF A SINGLE PLANT (TYPE 9).

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Tante XV LE

Dimensions of Lamina. Types 9 and 51.

SS

(Measurements in Centimetres.)
No. of Ven@ Breadth. Length. Length: Vena- Breadth. Length. \Length. | Vena- " Breadth. Length. lene
Leaf. tion. Breadth. tion. Breadth. tion Breadth.
iste ee. ee eS
Type 9.
Plant 1. Plant 2. | Plant 3.

1 | Damaged Damaged - Damaged

2 2”? ” oe)

3 2? ” | 2”

+ 40° 14.6 38.5 2.6 352 12.8 33.8 2.6 SHH" 11.6 Bul 2.7
5 30° bas 44.0 2.9 40° 16.0 42.0 2.6 40° 12.6 39.0 Stl!
6 40° 17.6 Bill 27 3.0 40° 15.0 45.4 3.0 40° 15.0 48.0 Bee
7 45° Whats: 57.0 3.2 45° 17.9 49.0 2.8 45° iat) 50.6 3.0
8 45° Ube 56.0 Z.B: | 50° 15.0 53.4 3.0 50° 14.0 — a
9 50° i oe 70.4 A.D 45° ie: 64.1 Bn 7 45° 13.2 58.0 4.4
10 50° Ee 7 65.3 5.6 50° 14.7 70.2 4.8 50° 12.3 597 4.8
11 50° 14.3 60.0 4.2 45° 11.2 60.3 5.4 50° 11.5 61.0 5.9
12 50° 14.9 74.3 5.0 50° 12.0 70.0 5.8 45° 12.1 TRARY 6.0
13 50° 9.3 65.0 ri) 50° 10.6 63.4 6.0 50° 10.7 bane 5.0
14 50° 8.3 63.5 (els: ADY 8.8 69.7 8.0 45% The 59.5 8.5
15 bog 8.2 67.0 8.2 50° 6.1 52.5 8.6 45° Thea 65.8 9.3
16 50° 4.5 53.0 11.8 a 2.9 44.6 Weak) i = 5.0 53.8 10.8
17 — See 51.5 16.1 remainder not measured. —- 3.4 46.0 13.5
18 remainder not measured. | remainder not measured.

| Type 51.
Plant 1. | Plant 2. Plant 3.

1 Damaged | Damaged Damaged

9 | ”

3 3° +) | 9°

+ > 9 9?

5 75° 13.0 37.0 2.8 ibe 17.4 34.5 2.0 5

6 1am 17.9 Sonal 1.9 70° 17.6 35.9 2.0 70° iWin) 35.0 2.0
1 iy 19.1 S70 1.9 De 23.2 42.8 1.8 D9 18.4 36.1 2.0
8 80° 21.4 42.0 1.9 u(y 23.2 44.7 1.9 70° 20.7 41.3 2.0
9 Se 21.5 42.3 a (Giz Don 48.0 1.8 75° 24.7 47.5 1.9
10 80° 23.9 44.3 1.8 80° 26.8 48.8 1.8 80° 23.8 46.1 1)
11 15° 24.2 48.7 2.0 80° 24.6 48.1 1.9 80° 25.0 47.1 1.9
12 80° 24.7 46.0 1.9 80° Pathe MW 50.5 1.9 80° 25.4 47.4 Lg
13 80° 23.8 46.9 2.0 80° 21.6 48.8 1eSeaesOx 26.8 48.6 1.8
14 80° 25.6 46.3 1.8 85° 29.0 49.3 Wee 80° 25.9 47.2 1.8
15 80° 27.1 46.8 lees Tbe 28.3 45.9 1.6 80° 25.6 46.7 1.8
16 80° 25.9 47.2 1.9 80° 26.4 47.0 1.8 80° 2b. 44.8 Li
17 85° 26.0 43.1 ewe 80° PATE 45.2 LG 80° Dien 46.7 Lag
18 85° 22.5 40.7 1.8 80° 28.8 43.4 1.5 80° 25.0 42.6 1a
19 85° 24.2 40.4 i Waar 85° 24.0 42.0 Vib 80° 24.2 40.1 1%
20 85° 24.0 39.2 1.6 80° 26.6 41.8 DG: 85° 22.0 39.2 1.8
21 85° 20.8 36.8 1.8 80° 24.9 39.4 1.6 80° 24.2 38.0 1.6
22 80° 23.5 34.2 1.6 80° 26.1 39.0 1.5 80° ON 7 35.3 18
23 80° 19.8 32.7 Wee 15° 21.8 34.3 62 4) SOE 20.3 32.8 1.6
24 80° 18.3 30.8 ibe 80° 15.3 27.0 LS 80° 18.9 28.6 1.5
25 | 75° 15.5 25.2 1.6 80° 14.6 24.6 eye PP zi)? 13.4 24.6 ly
26 16° 9.9 18.8 1.9 10° 8.4 18.0 eat ee se 9.4 17.8 il
27 | — *6.8 14.8 2.2 — 5.5 14.0 2.5 remainder not measured.
28 | remainder not measured. remainder not measured.

Taste XVIII.

Dimensions of Lamina. Types 35 and 16.

(Measurements in Centimetres).

a e Veus- Breadth. Length. svemg he) Vene- Breadth. Length. Lengthy) ess Breadth. Length. Leneues
Leaf.| tion. ; Breadth. _ tion, Breadth, ton. Breadth-
Type 35.
Plant 1. Plant 2. Plant 3.

1 Damaged Damaged Damaged

2 99 99 99

3 be) 39 >

4 9° ’ 99

5 . 70° 18.3 32a IES 65° Gees 28.6 17

6 10° 22.1 shell 1.4 70° 24.0 Sono 135 65° 20.6 33.0 1.6

ee T° DHE: 34.3 ile Oe 24.7 Bist 56 70° 25.4 39.0 1.5

8 80° 26.8 — — | 75° 31.0 45/45, Wad. 102 26.3 40.8 5

9 | 80° 31.8 44.3 Ibe at 80° Bell! 49.0 1D 75° 25.0 39.8 eG
10 80° 25.5 45.5 Lest! 80° 29.9 45.6 1.5 80° 34.0 49.2 Ped
11 | 80° 31.0 48.7 1.6 80° 34.8 52.6 5 80° 30.4 50.0 1.6
ii S02 30.7 48.0 1.6 80° Poh el 46.4 NR 80° 28.4 48.2 ey
13 80° 28.1 45.3 1.6 80° 30.8 49.9 1.6 80° 33.0 52.3 1.6
14 80° 3333683 49.1 1.5 (a 351.8 54.4 eS 80° 26.7 43.6 1.6
15 80° 29.0 41.2 1.4 80° 2952 4:3'.3 1.5 80° 30.5 49.0 16
16 80° D200 46.3 1.4 80° 322 46.1 eS) 80° 28.0 41.9 1.5
17 80° 29.7 39.5 [Sem esos 26.8 39.5 135 80° 19.3 Sod 1.6
18 80° peng] BBY 1A 85° 16.4 25.05 16 80° 19.5 30.9 E26
Ole SOc 20.0 29.0 PS Ope |e Lies 19.6 1.8 THe 9.8 19.0 Ley
20 ia 123 20.4 Weert remainder not measured. 60° 560 12.8 2.4
21 60° 4.8 12.8 Ye 7/ remainder not measured.
22 remainder not measured.

| Type 16. |
Plant 1. | Plant 2: | Plant 3.

1 | Damaged Damaged | Damaged.

2 3° 99 99

3 99 9° 9°

4 =. 60 6.1 Piles 53510) EP

5 ~ | Damaged z

6 50° 7.8 30.8 4.0 60° 7.8 28.0 3.6 55° Seo 29.7 3.6

7 55° e.O 25.0 SG 50° oats 30.8 4.) 60° 8.6 ol 2 3.0

8 50° 9.1 30.6 Sore i) GO 9.0 36.6 4.1 50° ey 39.0 4.0

9 55e 8.7 BY He 2 aay. | Or 8.8 BY ay | 3.9 60° 9.8 Belo | 4.0
10 50° 10.7 43.1 SOF Wi bbe 9.1 GLI 3 7/ 4.6 55° LOR 41.2 3.9
11 50° 10.2 rib Ber Al 45° 9.5 — — 50° 10.9 46.3 4.2
12 aoe TAO 46.3 4.2 SOP 9.9 41.9 4.2 552 113800) 45.7 3.8
13 50° 12-5 51.6 4.1 50° LORS 44.6 Al De 50° 12.4 48.5 3.9
14 45° a2, 47.4 3.9) |) 502 10.1 41.0 4.1] 55° ile) 48.1 4.0
15 45° tS Dee 4.5 55° 10.5 43.4 ahd! 50° Ae 49.0 3.9
16 50° Tit eat 46.9 AS eee OS 10.4 43.6 4.2 50° LES — —
a7 | 50° PAS 49.4 4.1 50° 10.5 44.4 4.2 55° 1 is 45.7 4.0
H8- | 50° 13.0 Pile 4.0 50° 11.5 43.8 3.8 50° 12.8 50.3 3.9
19 50° 12.4 48.3 3.9 502 9.7 AN 7 4.3 50° 11.3 48.8 4.3
20 50° 13.0 Fa 5) 4.0 | 50° 10.0 42.7 Ae 50° PAE: 47.0 a8
21 45° ihe 48.1 4.2 50° 10.0 41.4 AT | OF 10.3 42.0 4.0
22 55° ies 48.3 AL 50° 9.7 40.0 4.] 60° 10.5 40.3 328
23 50° L250 46.4 3.9 50° 9.2 38.9 4.2 50° 10.5 43.8 4.2
24 50° 9.8 40.4 4 50° 8.6 37.5 4.4 55° 9.5 39.0 “eat |
25 | 50° 10.6 39.3 Sha 50° 8.5 38.9 4.6 50° 9.0 36.6 4.0
26 | 45° a0) SEG 4.5 50° Bod 34.9 4.5 55° 9.2 36.6 4.0
DT 45° 7) 27 0 4.7 45° 6.8 BY AR 4.8 50° 6.5 28.0 4.3
28 — 2.9 19.6 6.8 45° 6.2 29.1 4.7 50° 5.4 24.8 4.6
29 remainder not measured. — 4.4 21.8 5.0 — 2.4 16.8 Fj 8
30 remainder not measured. remainder not measured.

Taste XIX.

Dimensions of Lamina. Types 23 and 38.

(Measurements in Centimetres. )

| |
No. of Vena Breadth. Length Length. |Vena- Breadth. Length Length. |Vena- Breadth. Length Lents
Leaf. | tion. ‘ Breadth.| tion. ’ Breadth.| tion. ' Breadth.
ee | tare Eee = |
_
Type 23
Plant 1. Plant 2. Plant 3.
1 | Damaged Damaged Damaged
2 ”? 9 be]
3 ” ’> ”
4 ” ” ”
5 si of 70° 18.2 33.5 1.8
6 65° 13.2 33.7 Gg | “FOo 22:0 44.8 2.0 65° 22.5 44.8 2.0
7 70° 19.2 — — 702° 25eL 51.0 220 70° 23.4 48.9 rel,
8 Ge 2360 — — 70° Zoro 56.6 Depl! Tie 28.2 54.5 1.9
9 65° 24.6 55.0 2.2 [ae » Waits: 60.6 Well nO 28.4 62.0 py?
10 qe 27.0 66.0 2.4 ia Oe Trae if 2.4 7D- 28.4 Glee 2.2
11 705 27.0 60.5 PA Se 7p° 27.4 69.4 2.6 thom 24.6 58.7 2.4
12 70° 26.4 6225 Yar 75° 27.0 63.0 Dies 70° 26.4 62.7 2.4
13 Tae 27.5 62.5 3 Te wAGS I 61.0 P4e 3) Tar PAT Al 57.4 2.2
14 70° 21.9 56.7 2.5 Tee Pane 55.7 eA 5) 70° 23.0 52.0 2.0
15 65° 23.8 55.1 Paina} 65° 20.9 50.0 2.4 75. Visa 40.3 273
16 70° 18.2 43.2 Die "0ce lo 40.5 2.9 70° 10.7 33.8 3.2
17 — 5.6 26.4 iN fe || — Te) NGA Dine 6.2 29.7 4.8
18 remainder not measured. remainder not measured. remainder not measured.
= er (wi eS eine een ee ee
Type 38.
Plant 1. Plant 2. Plant 3.
1 Damaged / Damaged Damaged
2 9 | > ”
3 > 9 99
+ 9° 99 >
5 ”? | 9° ?
6 : iS 6De S186 43.0 1.4 65° 24.7 35.5 1.4
7 a i Gs 32.0 42.8 th 8? 60° 28.4 43.3 1.5
8 65° 25.7 33.2 lee 65° 35.3 52.3 ib 65° 32.6 48.3 1-6
9 60° 94.5 37.5 eh alos 40.7 59.5 bess 60° 36.4 55.1 i ee 5
10 65° —s(:333..3 45.8 1:4 | 60° 383.6 56.1 La 65° = 333.6 56.0 i
. 65° 35.4 52.5 Lio 65° 38.7 62.1 6 65° 38.1 58.8 1.5
12 60° 32.0 52.0 1.6 65° 39.7 63.2 ae) 65° 36.5 63.7 1
13 65° 35.5 55.2 1 65° Silver 56.1 Wate) 65° 33.2 57.9 i Bh |
14 60° 36.0 60.8 ety 65° 38.6 63.0 eR 65° 39.1 66.2 Va
15 65° 34.0 57.6 ieee 65° 29.1 Bile 1.8 60° 31.2 58.5 128
16 65° 35.5 58.6 1 Wag 65° 31.5 55.8 1.8 65° 34.5 60.8 1.8
ys 60° 27.4 52.0 1.9 60° 38.3 65.0 Maat 60° 37.4 59.5 1.6
18 65° 29.8 50.5 1 a7 65° 28.6 49.9 i hewk 60° 29.6 51.0 1
19 60° 34.2 56.0 1,6 65° 34.5 55.6 Uke 60° 35.3 56.4 1.6
20 60° 30.0 52.2 1 bee 65° SPAR? 52.4 L.G6 65° 32.5 49.8 le
21 60° 25.2 47.1 1.9 65° 27.0 44.2 LG 60° 31.2 51.2 1.6
22 65° Ol. tk 46.6 i 60° 30.2 48.4 6 65° 30.7 48.2 1.6
23 60° 5 AY, 45.1 ie 7 es 65° oe ee | 39.8 Led 65° 23.3 38.0 1.6
24 | 60° 24.4 43.8 1.8 65° 18.8 33.6 rs 60° 23.3 39.5 Lf
25 | 60° 19.7 36.6 1.9 60° 25.2 42.3 Wed 60° 20.8 32.3 1.6
26 60° 18:3 3362 18 60° 16.4 28.1 iit) 65° 14.9 23.5 Lo
27 | 60° 20.3 34.5 iDNay 65° 15.2 25.4 1 bes 60° 14.1 22.6 1.6
28 | 65° 159 Wik iba 60° 1023 18.6 1.8 remainder not measured, ‘
29 65° 12.6 20.0 iow | remainder not measured, .
30 60° 9.5 16.7 1.8
remainder not measured

GABRIELLE L. C. HOWARD. 85

for measurement and in many types the change between these
and the significant leaves is very gradual. It is probable that in
America the general uniformity of the kinds grown makes this
method more applicable.

The actual procedure adopted in these investigations was
as follows. Very small cardboard labels were prepared with
the number of the plant and culture, and these were threaded
with fine copper wire. These labels could be readily affixed to
the leaf by passing the ends of the copper wire through the mid-
rib, and bending them flat on the under side. The individual
leaf chosen by the investigator for measurement was immedi-
ately labelled in this manner while on the plant. The leaves
could then be removed and measured in some convenient place.
The leaf after measurement was pressed with the label still
attached, and thus a complete record of all the plants examined
has been preserved. It is hoped to use this material in dis-
entangling the factors which are concerned in the shape of the
leaf. This method of attaching a label, which lies flat on the
mid-rib, and therefore interferes with no measurement of the
leaf while still on the plant, has proved most useful. The
labour of preparing these labels, threading the wire, and the
subsequent drying of the leaves, however, would not be possible
for several thousand plants except for the cheap labour avail-
able in India. A second and a third leaf can be removed
immediately above or below the first one chosen.

The venation of the leaves was determined by measuring the
angle between the mid-rib and the lateral vein with a horn
protractor. Only differences of 5° were noted. Experience
showed that such determinations gave good results, and that
the error generally was not more than +*5°. These measure-
ments were carried out in two ways, either while the leat was
still attached to the plant or on the detached leaf which had been
selected for other measurements. The first method has the
advantage that all the leaves can be examined in a general

manner by eye at the same time as the measurements are made ;
6

Taprr! XX.
length
Venation and ratio of the leaf in the various types of N. tabacum.
breadth
t noe | = |
Venation. pene Venation. ne Beth: | Venation. ie net Venation Loe
Breadth. Breadth. | Breadth. Breadth
35° Type 19 2.4 50° Type 6 4.1 60-65° Type 38 1.7 75° Type 23 2.3
meer * a SG i ATS ees 3
a ee hes 6d, 1 dG: yee 65° BE bal O iit » aes
fe 8 4.4 se LY ee BD — , Behe aoo
pe ery oe ee QT p78 70° peer celhs ro eee
So aes So wad a 80° », S52. 128
os 12 3.3 | 65° 33. ZOD 2.8 mS 39 139 Pree! / 139
ei eD . —— AD Pied po. aoe
. eSB 60° SE. ays (bas }, 7 Saal or SS ee
ee fg nid , 438 2.2 ee
<. “Si Te Teh. ipa . ACAI) 29.8 90° «=. ’ “oye
cee ee) Gee eee ye ee
45° ge ine 2.3 oF 36 2.0 cs 38 1.8
| a ae |

86 STUDIES IN INDIAN TOBACCOS.

the second, that all the measurements such as width, length
and venation are made on the same leaf. Comparative deter-
minations on the same cultures by both methods give similar
results. In most cases in 1913, the data given are the mean of
two determinations, but experience shows that where time is
limited one careful determination is quite sufficient. The
difference between the determinations was not greater than the
error of measurement. |
The angle of venation appears to be extraordinarily
constant for any one type (see Table XVII) and to be
largely independent of the shape and width of the leaf.
This is shown very well by Type 9, in which the angle remains
constant at 50°, while the shape and width of the leaves varies
greatly. An examination of Table XX shows that the angle of
venation in the Indian types varies from 35° to 90° and that all
the intermediate grades occur. The smallest angle, 35°, is found
on a comparatively broad leaf. In Table X XI the relation
between the venation and the shape of the leaf as expressed by
the ratio length/breadth of the leaf is shown for the F,, generation
of Type 16 and Type 35, and also for all the Indian types on

GABRIELLE L. C. HOWARD. 87

TABLE XXI.

length
Venation and ratio in F, Type 16 X Type 35.
breadth
oe Shel Size of Angle. } No. of
breadth) 49° 45° 50° 55° 60° 65° 70° 75° 80° 85° 90° aa
La | = FF ree ae l1—_—_- — i = 2
1.8 —— 1 — 1 1 — ] 4
Cf a er eee ee | 16
2.0 |; — —- — — ft 3 7 6 12 4 1 37
2.1 | — 1, o— 2 9 8 5 5 4 7 1 42
2.2 — 1 5 4 7 6 19 9 8 ] 1 61
2.3 — 1 2 3 13 8 8 3 se = — 42
2.4 —_—_- — 4 8 26 7 13 6 7 68
2.5 — 1 a 11 10 5 9 3 2 —- — 45
2.6 1 1 8 7 9 7 2 1—_—- —_> — 36
2.7 — 2 6 2 4 5 4 — 1 => = 24
2.8 — 1 10 4 5 3 1 1 1 —- — 26
2.9 —_—_ — 1 1 2 3 —- — 1 —- — 8
3.0 — 2 30 — 4 1—- —- —> — — 10
3.1 — 2 3 1 2 1 LS SS 10
o.2 —_ i 30 — LS Ss ae SS SS | 5
3.3 | — — 3 2 —- —- —- —- =—- — — 5
CO fe a aaa 1 hoes pen Te id
3.56 | — — — I! 1—- —- —> ~—> —>— — 2
3.6 —- —- — 1 —- ~—- —- —- —-> ~— — | 1
ot —- F—- Fe O ae ae ae aee aeeaee —
3.8 —- -—- F—- TF a ae ae ae ae ee —
3.9 -—- F—- Fe a a aeae eeaee aeee —
4.0 —- —- Fe ae ae ae ae ae oe —
4.1 _- — l1—- —- —- —|- —- —-—- =—- — 1
Total 1 VSS Ot 4855 977 65 69 38 43 14 5 447

Table XX. It will be seen that there is no true correlation
between these two characters, and this is confirmed by many
of the F, cultures in which the venation was constant, but the
shape of the leaf very variable—culture 202 and many
others. There does, however, seem to be a limit to the occur-
rence of the higher angles. Those above 75° are only met within
leaves with small ratios. A more detailed examination of the
venation affords a possible explanation of this, for that there is no
incompatibility between alarge angle and a small ratio, is shown
by the banana leaf. There is a striking difference in the leaves of
the tobacco between the behaviour of the lateral veins which arise
at an acute angle and those in which the angle is large. The

88 STUDIES IN INDIAN TOBACCOS.

former always go straight to the margin, which they naturally
meet obliquely, while the latter curve at this point and run
almost parallel to the margin. This may be to avoid rupture of
the lamina, or it may enable the food channels to serve a larger
portion of the leaf for in the case of the acute-angled venation
the veins cover a long distance, while when the angle is 90° the
distance is very short. If the leaf is broad, this curving upwards
of the ends of the veins does not affect the angle near the mid-rib
but if the leaf is narrow the latter would be reduced. This may

Taste XXII.
Venation in Type 9 Type 51.

| Total

[Venation) |
oi || | No. of
| Parent. | 40° 45° 50° 55° 60° 65° ‘70° 75° 80° 85° 90° Plants.
| |
Type 9 1912 | 1 27 26 — — a 54
Type 9 1913 Peao) 16 — — 26
Type 51 1912 | —- —- — 5 23 20 3 51
Type 51 1913 — 14 18, — — 32
F, Type 9 x Type 51 1912 | — — — 19 47 1l tH
F, Type 9 x Type 51
1911 19 12 147 30 200 21 89 29 42 12 7 608
Type 9 x F, Type 9 x |
Type 51 LOLY ti 38 — 173 2 55 — the - 269
Type 51 x F, Type9 x
Type 51 1911 | 1 — 22 5 51 — 37 4 49 — 38 207
F, Type 9 x Type 51
738—1912 | 50° 1?’ 9 “4g "96°39" oF tS 44 ee
745—1912 50° Leis PbSh, 20. 22 1 — 139
736—1912 | 60° 1 2 8 38 36 31 11 — — — — J] 127
737—1912 | 60° Owe Bk OT Us 9 1 — — — —! 160
740—1912 | 70° —_ — 7 18 41 41 22 — — — — 129
694—1912 | ‘75° —- — — 1 TT illise ooy reo. Lb 4 — 100
132—1912 | 80° - 1 — 10 31 °39 21 4 116
167—1912 | 80-85° —- —- ie ee Ls 282246" 35 98
F, Type 9 x Type 51
738-19—1913 | 40° ll 78 1464—- —~—~ —~ ~ — — — — 105
738-96—1913 | 50° 22 40 4 —- —- —- —- —- 66
738—58—1913 | 50° —_—_ — 1 15 80 ll ~—~ — — — — 107
738—-15—1913 | 55° D> Pe 38) 28. aT per yey = a Se 101
738—76—1913 60° —_—: 2 13. 2%. 45 8 7 - —- 102
738-—81—1913 70° —_-_ —_— — 1] 85 44 4—- — — — 134
694— 1—1913 70° —_—_- —_—_ —_— — 1 41 40 21 — — — 103
694-38—1913 | 70° — — - — —-— _——-—~«x—'*«=«KX«10 *45'«O-32 «=«20° — | To7
694-23—1913 | 75° —- — — — — — 4 32 146 3 —| 565
694-103—1913 | 80° —_—_- —_— — 1 S Wb 26-27 20 2— 99
— — . 0

694—10—1913

GABRIELLE L. C. HOWARD. 89

explain why there is a limiting effect without true correlation.

The study of the venation has yielded some most interest-
ing results and has demonstrated very conclusively the existence
of homozygotic combinations with values between those of the
parents. Table XXII gives the results of the cross Type 9 x
Type 51. The F, is intermediate as usual, and the F, covers the
combined range of the parents. In the F,; only one culture,
167, appeared to be uniform, and this had a mean venation
slightly greater than Type 51. The limits of variation were,
however, very different in the various cultures, and two
cultures in which these overlap very slightly, 738 and 694,
were continued. From culture 738, with a range of 40° to 70°
in the F,, the plants with the lowest and highest angles were
grown in the following year, as well as four others. In the F,
generation two cultures 19 and 96 were uniform with an average
venation of 50°, and a range of 40° to 55°, two others 58 and
81 were uniform with an average of 60°, while one culture 15
resembled the parent culture 738 in every respect. The
behaviour of the other culture is not very clear; it may
indicate another homozygotic combination at 55°. Thus
two homozygotic combinations, representing 50° and _ 60°
respectively, have been isolated, one of which is a replica
of one of the parents, the other is new. The F, cultures
of 694 do not give such definite information, but they indicate
the occurrence of another homozygotic combination 65°-70°,
and it is possible that 694-1 represents this.

The cross between Type 16x Type 35 (Table X XIII) gives
very similar results to those described before. All the cultures
(four) with a venation of 50° or 45° have bred true ; one culture
of which the parent plant had a venation of 60° has also bred
true, and here again is the formation of a homozygotic
combination intermediate between the two parents. Cultures
200 and 8 are possibly uniform with an average of 75°, again
a novum. The other cultures are apparently heterozygotes
splitting within different limits.

TastE XXIII.

Venation in Type 16 =< Type 35.

Venation Total
of 40° 45° 50° 55° 60° 65° 70° 75° 80° 85° 90°| No. of
Parent. | Plants.
Type 16 1912 Sees ee an Se
Type 16 1913 — — 22 10 1 —-—- —- —- =—- =— = 33
Type 35 1912 | ;—{}$ S#s —= os — So — — 3) 10> 952 9 — 74
Type 35 1913 | ;}— — — 13 19 2 — 34
F, Type 16 x Type 35
1912 ;|_—} S#s — ss —-— — 7 23 — — — — — 35
F, Type 16 x Type 35
1912 — 6) (32, 429) (58 43.9 410 925: 20 7 1 262
F, Type 16 x Type 35 |
27—1913 45° |— 17 78 14 1—- —- —- — — — 110
202—1913 50° — 22 %3 9 3 o- Se ee ee 107
190—1913 50° — 21 67 21 — ~—~ ~—~ ~—~ ~—~ —~— — 109
231—1913 BOCan Boe OkeGb a a eee
251—1913 See yk SOLA Sisee sty) Rly eS = Sa as 107
142—1913 55° — SF 38: 140 ~20 2—- —- —_ —_—_ — 108
163—1913 | 60° — — — — — 18 67 16 — — — 101
15—1913 70° — 2 6 8s24 30) 18 6 1— — 105
35—1913 | 75° —_- — 1 5 Seas pole 6 5 — 94
9—1913 | 80° —_ —- —_— — 4 6 42 41 14 — — 107
BRD = tOTs. Js e802 4 et ee eo Ds! G's 94: > 19
8—1913 85° — ie 256) Sab 7 — 105
TaBLE XXIV.
Venation in Type 23 x Type 38.
Venation| | Total
of the | 40° 50° 55° 60° 65° 70° 75° 80° 85° 90°} No. of
| Parent. | | Plants.
Type 23 1912 | ees aS. oR 0 ne
Type 23 1913 | Le ay Oe ae eee 26
Type 38 1912 | Se a
Type 38 1913 SN Ry er ia aa Be Eee 4()
F, Type 23 x Type38 1913 a ee A Ck tee
ate et Ea ee . ae [—— Se Se EE EE SSE EEE Eee eee eee
F, Type 23 x Type 38 1912 | i a A 0s AG. 38 6 eee
F, Type 23 x Type 38 |
213—1913 —_ —_— — 38 62 13—- — — — 103
6—1913 75° — 24 69 ll — — 104
155—1913 —_—_ —_— — 1 136215—-_— — 91
159—1913 — — — 12 32 38 23— — — 105
104—1913 70° —- — — — 1° 55 39 650 1 — 146
111—1913 Like F, 60°-80°

204—1913 | 60°—80°

°

Qi- adh]

Aa Ey exglbKe

Gg adAL

91 AdAL “4 NI SHAVAEAT AO SNWYUOS

uoigeuauas®) be

GABRIELLE L. C. HOWARD. bt

As the parents in the cross Type 23 x Type 38 (Table X XIV)
are so similar as regards venation, no investigations on this
character were contemplated. A few measurements were
made, however, in the F, as some other characters had given
curious results in this generation. The results obtained in
this generation afforded very striking evidence of the accuracy
of the method adopted, and so a few cultures were measured
in the F;. Type 23 has a range of 70° to 80°, Type 38 of 60° to
75°, both therefore overlap. The plants in the F, generation
had a range of 60° to 85°, the combined range of the parents.
In the F,, cultures were found like both parents, i.e.,
Nos. 213 and 6, some like the F,, Nos. 111 and 204 and others,
which indicate the possibility of another homozygotic combina-
tion with an average of 70°.

jp mone EEAE- SHAPE:

The most difficult characters to investigate are those
connected with the shape of the leaf. The large number of forms,
all slightly different, with apparently endless modifications of the
individual parts, appear at first to defy analysis. A detailed
study of the existing forms and their behaviour on hybridization
shows that the number of characters is not so great as might be
supposed, although the factors composing such characters
are probably numerous. The form of a tobacco leaf can be
expressed by a determination of the following points: (1)
ratio length/breadth, (2) position of the greatest width, (3)
amount of indentation at the apex, (4) amount of indentation at
the base, (5) shape at the point of insertion, that is, whether
auriculate or not. According to the evidence which has accu-
mulated during these investigations all these characters are
inherited quite independently of one another.

The influence of the ratio length/breadth on the shape of
the leaf is obvious. The ratio depends on the two variable
characters, length and breadth. Apart from environmental

92 STUDIES IN INDIAN TOBACCOS.

influence, the width of the leaf is not dependent on the length.
The Indian types of tobacco, with their great diversity of leaf
shape, illustrate this point well. In countries with a high level of
agriculture, economic considerations have eliminated all varieties
except those possessing a certain type of leaf,a broad leaf witha
large surface. In India, on the other hand, where the conditions
of seed selection are primitive, and where thickness of leaf is |
desired for certain purposes, the cultivation of forms with very
narrow leaves has persisted. Type 9 (Plates land XVIII) is a
very good example of a long narrow leaf and in Types 16 and
35 we have two leaves similar in length (average length, 42.9
cm. and 46.6 cm. respectively) with a very great difference in the

TABLE
Length
Ratio —-—— in
Breadth
eee
Bog
SG 2114 15 16 1.7 18 1.9 2.0 2.1 2.2 2.8 2.4 25 ote
Type 51 1912 —- —- — 1 Word all 3 - Fr O e
Type 51 1913 — — — — 6 610 9 l- -—- - cere
F, Type 9x Type 51 1912 A 8 i eee
F, Type 9 x Type 51
167—1912 —- 2 QPOs 26) 15) 21 8 i 1—- —- — —
132—1912 — — — 2 7 21 29 26 14 5 2 2 — — —
694—1912 —- —_- — 1 Ze elS) 9) 20) 3s 0 5 3 1 1 —
737—1912 — =, = 1 3 6 306 (66. 4 710 0Rme
738—1912 — —- S—-— SS Te ee el i 4 6 10 10 16) i
745—1912 =—=-S SS SC Tre Te lc 1 — 3) 349 20 ie
736—1912 — — FF rT Se ee ee 1 4 7 © 1
740—1912 —- - - hl — 2 3 6 18 20
F, Type 9 x Type 51
738— 81—1913 2.7|/— — - 3 5 412
738— 15—1913 2.9)—- —- —- -—- —- -—- -—- -—- -—- 1 8 9 22
738— 58—1913 3.2);— — — — — — — — — — — — — 4 6
738— 76—1913 3.2); -—- —-— —- —- —- -—- -—- 6 6 Il Jaa
738— 19—1913 3.38) — — a a SS ee 2 39
738— 96—1913 3.8)/— — — — ~—~ —~ — + — — — — — 2 8
694— 23—1913 19|/— 2 ] SP LO 8s . 4 1 2- - - - hc eS
694— 1—1913 2.1);/—- —- —- — — — 3 656 22 28 19 19 6 —aae
694—-103—1913 2.2 | — 1 il 8 17 14-27. 4 10 8 6 1) ae
694— 38—1913 2.2};— — — 2 16 21 27 27 12 2 — — 1—- =
694—- 10—1913 2.2 | — Lol 8 14 26 32 145 56 — — — — =

KXYV.

GABRIELLE L. C. HOWARD. 93

width (average width 11.1 cm. and 29.2 cm. respectively).
On Plate XIX are shown some of the leaves of the F, genera-
tion of this cross, drawn to scale.

All the evidence accumulated during these investigations
tends to show that differences in width can be inherited quite
independently of the length. The very great effect of fluctu-
ating and temporary changes in environment on the size of
the leaves makes a thorough study of the inheritance of the
width or of the length difficult and necessitates a large number
of measurements. Time has not permitted me to measure
sufficient leaves per plant to make any very definite statements
as regards the factors composing these characters. As regards

ype 9 X Type 51.

|

to

Reone! | |

PROUD D0

—

a

3-0 3.1 3.2 3.3 3.4 3.5 3.6 3.7 3.8 3.9 40 41 42 43 44 45 4.6 4.7 4.8 4.9 5.0

a _
oemotol | |

—_

for)

—_
SOW

PRIM t are ac ors

BRoaeol | |

Male eal ries esicoiesies
LST ec marae
Pe | Weonbe esteniks

TABLE

Length
Ratio —-—— in
Breadth
Ratio

teenie ad Le 16 17 18 1.9 20 21 22 2.3 ah 2) 2
——— a eee ssS—~Ssts t
Type 35 1912 | les as 6 ——- — > =e
Type 35 1913 | 2 is 8 — — — — =e
Type 16 1912 = 2 2S SS =| = Se
Type 16 1913 — 2. 2 eS] | > See

F, Type16 x Type 35
1913 | | S24 + 1 3a

F, Type 16 x Type 35
1912 | «4 16 87 «42 «Cl

F, Type 16 x Type 35
9—1913 1.9 2 7 14 23 Sl 16 woes
g 1918, ,|, 2220 0) 88 2
Pen Corie St te eee 07 20 12 10 5 — =
231—1913 i ee
202—1913 a
IOI hoe) “|=. => = 17 16 13 14 19> 005
ETD iO. As Rom oat ae ee 5 7 12 14 20 280am
aan Motor |e ee ee ee 3 11 29 17
nisi | 2 = Se 3 8 17 16 23
aoe | 20 Mia ee ae 1 2 15 16 22 18
190—1913 = is == _ = =| ae
27—1913 2 a — 1 1a

TABI
Length
Ratio i
Breadth
| Ratio |
| of 2 oc j
| Parent. Tpaenles ale Bedeyl les 21-8220 2.1 2.2 2.3 2.4 25 2
Type 23 1912 | 1— 2 6 2) 33
Type 23 1913 == = 9 9 § 4.050
Type 38 1912 | — — @ 14 10 10 4 2 =
Type 38 1913 a 4 4 0 3. 2 8) Se
F, Type 23 x Type 38
1913 94 3) (9-22 (8 4 = 9 ee
F, Type 23 x Type 38 |
1912 | a8 2 618 27 87 “be Claes 32 20 16
F, Type 23 x Type 38
104—1913 1.6 4 43031 2 6 — = = eee
6—1913 1.9 2 11 13 26 16 48 11 ages 1:
117—1913 2.0 — 9 | 12 12 9 2 6
20.4—1913 2.0 — og Ban 92 16 1 1
159—1913 20 |\— — — 4 12 28 19 24 1 8 ae
111—1913 Ba terea) are 2 41 6 21 20 18 143 2
9131913 | 2.2 |— 2 2 16 18 23 19 12 7 3 he
5 ERE VT NOON ge eer terse! 10 16 26 18 19 5 Se
angie | rete es a eo

XXVI.

Type 16 x Type 35.

2.7 2.8 2.9 3.0 3.1 3.2 3.3 3.4 3.5 3.6

3.7 3.8 3.9 4.0

4.1

4.2 4.3 4.4 4.5

Total
No. of
Plants.

ee ee) ee ee 51
a or
a a = SS ee ee ee ee ee ee eT
eee SS Se ee ee ee ee 37
ee ee ee a SO se, gg
DPCOG TO 4, ty 8 202. 1 = 2s Se Fe ek |, gg
se ee, eS =| 08
a a ee es ee SS 0
Sa rt es | 02
Snes 9) Gl 2 —6 yy of =, 9 2S SS ts SS =) 404
MTEC TO, ur > | fe SM Po Bey ek ee 94
rn ee ey Pe ee | 108
CIR eo) ie ee a ee | 69
a8 Ne Ee eG t,t ee eR SS | 1
ne me mote ee Re Ae a OS a ee I 08
mT Gg ee ee 2 er ee SS | 07
MMG—tnIT4 20 16 6 3 7 6 — 2 — 1-1) —- — — —] 108
mEreise 6 15 13 5 2 4 6 1 4 1-2 1.1 — — —| 105
mx VII.
Type 23 = Type 38.

Total
27 2.8 2.9 3.0 3.1 32 3.3 3.4 3.5 3.6 3.7 38 3.9 40 41 42 43 44 45 ad
nn ores A fe ee ee se Ee te SS 63
Pens yee ay CE re Pee ey ee ee 25
NR oot eR Pe a ea Goe =
me See ee AE eee eR ee 24
nr eg Er 5 9g fy 28) Sere ery 5 Pye sy Ue Se 2 |) eR fag
ee fet geht Se a Lae a Raa
ok Be ie BU Soke ott oe eee ee eels Ake see eS 97
Es ag 2 ag eg Bs Rp ae 98
ee SERS SES SME Sra De eee Wet) Ete eee aa Po 72
LE aS a a ee ee een eg ee er ee ey"
er teh anki gee Sein eee ena eh eee a ee Pee" a04
ge EE EE FS SBE a ay eg i ei eee PY
eee ere ry Re rene ee Be OS ae ee e105
tS Pos a a gee Se oe eh pe a I ge
a en la SS Yi ae Me a Ee et Le 99

a

96 STUDIES IN INDIAN TOBACCOS.

the environmental influences, however, both the breadth and
the length appear to be similarly affected, and the ratio between
the two is therefore less dependent on these influences. The
ratio length/breadth has been studied in three crosses (see
Tables XXV, XXVI, and XXVII). The results are very much
the same as in other characters. In Table X XV the ratio from
Type 9 is not given, as it will be evident from Table XVI that
no two leaves are alike in this respect. In Table XXIV the
most interesting culture is 694 which, although almost uniform
in the F, generation, has shown further segregation in the
following year.

Fig. 1. Leaf shape and position of greatest width.

In cross Type 16 x Type 35 where the length of the leaves is
approximately the same, but the width very different, the ratios
formed by a combination of the length of one with the width of
the other are the same as the original ratios. The data of the F,
generation clearly point to the action of several factors in
producing the difference between the widths of the two leaves.

The position of the greatest width is a very important
point in dealing with the leaf shape. The above diagram
shows how the appearance of the leaf can be entirely altered by
varying the position of this point.

SSS

Total
| P ; No. of
5 Giglio 9.18.3 8.5 8.7 8.9 9.1 $.3 9.5 9.7 9:9 10.110.310.5 10-7 10-9 11.1 11.3 11.5 11.7] Plants.
Ss oo ae a i 45

— —| 23

Me We eS =| 090

Bie XXVIII.
Inheritance of the Petiolate Character of the Leaf in Type 23 » Type 38,

Centimetres.

Lees EDM eeE UMBINIUNNES Nw Ler eT. oues 3.7.39 4143 45 4.7 5.5 5.7 5.96.1 63 05 0.7 6.
SS ae Ser fi Se = t=
a a = 21 Pa
10 18 17 13 18 12 5 rye dg 206 1 wl
7 = ae eee SS LS =
—— = a=- - ~ === n= 1 2 ‘
SS Soke SS == i aS hai ty ny See a Sf
2 739 = SS = — ee SSS SSS a = ——_ =
3 = = = = == Sa SS SS =S.= SS
12 26 0 hen oe == SS hh Sa ot ee en a
1.3 3 1 mM Ce os SPURS C49 Me A SAL GT, if Lier <
1.5 1 13 5 3 = ee es ee ey ee a Oa
26 = = i UL ak Teste eekt Ieee tate eek heen th 1
= 4 Sy le 6) 6) ea) ai a ge
= 5 ost m cu op} — 1 1 ote i
oo 2 a it Tu os WI auetal oie

Hanif the width of the Lamina ut the narrowest pomt

20 07

10,910.56 10-7 10.0 11.1

ellli titi

PT
Phtbtadayy

PLATE Xe

OG

Taots2i 694-23 694-23
9 69 30

694-23 694- 23 694-25
a 80 87

LEAVES OF T 51 AND OF CULTURE 694 (F, Ts x Ta).

GABRIELLE L. C. HOWARD. 97

In Type 51 the position of the greatest width has been
determined for a large number of leaves, and is always half-way
between the apex and the base. In Type 9 it is lower at a
point two-fifths from the base of the leaf. Determinations of
this point made in the F, generation on cultures derived from
694 showed that all were uniform in this respect and resembled
Type 9 exactly, although in size and ratio length/breadth they
were very like Type 51. On Plate XX some of these leaves
are shown, contrasted with a leaf of Type 51. Apparently this
character can be inherited quite independently of the ratio
length/breadth and it is possible to combine in the same leaf the
ratio length/breadth of one parent with the greatest width in the
same position as in the other parent. Similarly, very distinct
segregation as regards this character was observed in the cross
Type 23 x Type 38. A large number of cultures have been
measured with reference to this point, but as the number of
tables is already great, the results will not be published until
the fuller investigation of the factors concerned with the
shape of the leaf is completed. The measurements were carried
out by moving a steel measure at right angles up the leaf until
it denoted the greatest width, when its position was marked by
inserting a needle into the mid-rib. The distance between this
point and the base of the leaf was then determined.

The ratio length/breadth and the position of the greatest
width are sufficient to determine the general form of the leaf in
all cases. If we further imagine pieces of varying size to
be cut out at the apex and base of the leaf we are able to
reproduce the leaves of all the existing types. If we postulate
the existence of independent factors the effect of each of which
is to cause a different indentation in the outline, all the facts
which have been discovered during this investigation can be
explained.

The most striking example is that given by the cross
Type 23 x Type 38 (Table XXVIII). Both these forms
have sessile leaves, but the amount of indentation near

98 STUDIES IN INDIAN TOBACCOS.

the base is different (see Plate X XI, in which both parents
and typical leaves from the F, generation are shown). The
measurements were obtained by measuring the lamina on
either side of the mid-rib and obtaining the mean value.
This is preferable to any measurement across the leaf as it
eliminates the width of the mid-rib, which is probably an
independently varying character. The outline of Type 23
is almost straight, while that of Type 38 is suddenly contracted.
On hybridization, the F, has a bigger indentation than either
parent. In the F,, petiolate forms occur in which the amount of
lamina on either side of the petiole is less than .38cm. All stages
between these and Type 23 were found. Nine cultures were
carried on to the F,. The petiolate forms, Nos. 117 and 213, bred
true in the third generation. If the indentation in the parents be
due to two different factors or combinations of factors, then the
effect of their combined presence might be to reduce the lamina
to a negligible amount. These forms possessing both factors
would be homozygotic and breed true. Two cultures, Nos. 155
and 204, which in the F, possessed asmall amount of lamina gave
a progeny of petiolate and sessile forms which could be easily
separated both by eye and by measurement into three distinct
groups inthe ratiol:2:1. The actual numbers were as follows :—
Culture 159—petiolate 25, intermediate 52, sessile 27.
Culture 204—petiolate 24, intermediate 48, sessile 33.
The parent plants were probably homozygotic for one factor and
heterozygotic for the other.

Three other cultures gave progeny in which a certain
number of petiolate forms occurred, but these formed a
series with the sessile forms, and the number of such
plants was much less than a quarter of the whole. In two
cultures, Nos. 104 and 111, no petiolate forms at all occurred.
If the presence of both indentations is possible, the absence of
both must also be possible. Some leaves with an outline
showing even less indentation than Type 23 were found, but
the range of variation of the latter was, in 1913, so great that

FILIP S ROK

"86 AMAL X 66 AMAL “A NI SHAVAT JO SNYOS

‘uolqeaguag %,

I

ams? =

oe

J

>)

GABRIELLE L. C. HOWARD. 99

this could not be confirmed by direct measurement. These
cultures were very luxuriant and over-grew their normal limits.
The question is complicated by the fact that the indentation
of Type 38 is sudden and short, while it is gradual in Type 23.
Indications of a segregation in this direction were also observed.
These results indicate that the apparently stalked varieties
of tobacco are not really petiolate, but sessile. This
explains the alate nature of the petiole and the fact that in many
such types the upper leaves are sessile, the lower petiolate.

Further evidence on this point was obtained from the other
two crosses. Many cases in which the form of the lower part
of the leaf varied, but the indentation factors did not differ
so widely as in Type 23 and Type 38, were observed in the F,
generation of the cross Type 9 x Type 51. No petiolate forms
occurred, showing that the combined effect of the indentation
factors was not great enough to remove all the lamina. Different
ranges were shown by different cultures.

Observations in the field together with a comparison of
Tables XXV, XXVI and X XVII, show that these indentation
factors are inherited quite independently of the leaf ratio and also
independently of the width of the leaf. Environmental in-
fluences would, however, by influencing the vigour of the plant,
affect both alike.

Plate X XII shows three leaves selected from culture 738-81,
in which both width andlength are thesame but theamount of in-
dentation at the base varies. On the same Plate are shown two
leaves of 738-19, in which the length is the same, but the width
and the form of the base vary. In some cultures the configuration
of the base was uniform, such as 694-1. Plate XIX gives a very
good indication of the range obtainedasregards this character in
the cross Type 16x Type 35. The F, cultures showed very differ-
ent limits of variability. Im one case, culture 9, there was
apparently only one heterozygotic factor, a separation by eye
gave twenty-five plants with a similarly broad base in 108
plants.

100 STUDIES IN INDIAN TOBACCOS.

A similar explanation to the series of independent indenta-
tion factors appears to hold good in the case of the leaf apex.
A curious feature of the leaves in some of the Indian tobaccos is a
sudden constriction of the lamina near the apex of the leaf, which
gives the appearance of a prolongation of the surface into a
long slender tip; in others the tip is short. Good examples of both
these cases are seen in the F, generation of Type 16 « Type 35.
(Plate XIX). At first sight it would appear as if this sudden con-
striction were absent in both parents but indications of it can be
detected in the apex of Type 16, although the narrowness of the leaf
has diminished its effect. Evidence as to the independence as
regards inheritance of the factors concerned in the configuration
of the apex is given in this Plate. Other examples are shown
on Plate X XIII. In 738-81, Nos. 14 and 60, are shown two leaves
with similar apices but dissimilar widths, in 738-15, Nos. 38
and 96, are shown two leaves with the same width, but dissimilar
apices, while 738-19, Nos. 68, 47 and 23, show the two extreme
forms of apex found in that culture with an intermediate form.
All these drawings are to scale.

In three F, cultures of the cross Type 16 x Type 35, the
leaves could be divided by eye as regards the configuration of the
apex, into three different groups giving a ratio 1:2:1. This
shows that the plant was heterozygotic as regards one
apical factor only. Culture No. 9 gave 29 leaves with
broad apices; 50 intermediate; 30 with pointed apices.
Culture No. 35 gave 30: 50: 24. Culture 163 bred true as
regards the apex. These three cultures are illustrated in
Plate X XIV.

No information isas yet available as regards the inheritance
of the presence and shape of auricles at the base of the leaf.

It is hoped to publish a further paper on the factors con-
cerned in the leaf shape when the study of the pressed leaves
has been completed, but enough evidence has been given in this
section to show that the form of the leaf can be expressed by a
knowledge of the ratio length/breadth, the position of greatest

PEATE x70

VARIOUS -FORMS OF LEAVES IN, CULTURE 738 (he Ts x Tm).

PLATE. ~Ie

DY YY

738-81 738-81 738-15 758-15
14 60 38 96

N 738-19
73849 235

INDEPENDENCE OF WIDTH AND APEX.

l6X55 — G9

DIFFERENT FORMS OF APEX.

GABRIELLE L. C. HOWARD. 101

width, and the amount of indentation at the apex and base.
All these characters vary independently of one another and can
be inherited separately, and their mode of inheritance can be
explained by the existence of independent interchangeable
factors.

8. CHARACTER OF THE SURFACE AND THE MARGIN
OF THE LEAR;

These characters are very difficult to investigate. In the
first place, the nature of the irregularities in the surface may
be different; in the second place, the various stages and
combinations are very difficult to distinguish by eye ; in the third
place, even the so-called flat leaves have a certain amount of
undulation at the base. From a study of the various Indian
types, the following conditions appear to be the most frequent—
large undulations of the whole leaf as in Type 9, large undulations
confined to the base only, a general puckering of the surface
between the veins and asmall undulation or “‘frilling”’ of the edge.
This frilling of the edge occurs in Type 51, but is best shown by
the photograph of a plant which arose from the cross Type 9 =
Type 51, namely, No. 694-23 in Plate XVI.

The investigations on this point have been confined to
the cross Type 9 x Type 51. Type 9 has a leaf which is very
undulate all over, while in Type 51 the edge is frilled but the
surface is flat except at the extreme base, where an occasional
slight undulation may occur. Taking the leaf surface only,
it was found that the F, was intermediate and that the F,
showed a series of forms intermediate between both parents,
with a slight intensification, a few plants being more undulate
than Type 9. Two hundred and fifty plants were examined by
two observers. It was found possible to distinguish seventeen
plants like Type 9, and three slightly more undulate. Fifty-
seven plants were found with a slight undulation at the base or
quite flat. These two classes could not be sub-divided. The

102 STUDIES IN INDIAN TOBACCOS.

remainder could be classified as follows :—base slightly undulate,
slightly undulate all over, undulate at the base only, undulate
all over, very undulate all over, a little less undulate than Type
9,—showing that in all probability the undulation of the base
is determined by a different factor to that of the general surface
of the leaf. The numbers obtained indicate the existence of two
factorsand aratio of 15:1, namely, 17+ 3 plants like Type 9 in 250
plants gives a ratio 11.5: 1. It is, however, possible that: the
occasional undulation noticed at the base of Type 51 owes its
existence to a third factor and that the plants that were even
more undulate than Type 9 represent the combination of the
factors involved in Type 9 plus this factor. The ratio 247:3
makes this appear possible. The seventeen plants like Type 9
would consist of the homozygotic combination Type 9, the
heterozygotic intermediate between Type 9 and Type 9 intensi-
fied by the factor from Type 51. In a three factor combination
the ratio of these to the total number of plants would be 3: 64,
i.e., there should be twelve plants, not seventeen. The number
of plants examined is, however, two few to enable any definite
conclusion to be drawn.

The “‘frilling”’ of the edge appears to be inherited indepen-
dently of the surface undulations. This character cannot
be observed when the general undulations of the leaf are great.
In the present case this character was observed on fifty-seven
plants. Of these, fifteen had no frilling at the edge, the others
were frilled to a varying degree. This gives a ratio 42: 15, or
2.8: 1,so that here we are dealing with a simple factor. Some
of these plants with flat leaves and frilled margins were
carried on in the F, generation, and the following results were
obtained :—

Culture 694—105 plants, 78 with frilled margins, 27 with flat
margins.

Culture 167—99 plants, 73 with frilled magrins, 26 with flat
margins,

PLATE XXV.

auae

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Length of flower

20
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Soelnial . SOs seeenRenn8 CAT Bale
SEH SE earl Conc ~
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ys latal = OCT Festool ay
= 20 10
Co BeSer sl |
rH sada Sposeceedaecari SececaeciaaaS
Gan HES ENPeooeeeeee oH ]
aeanne sania nae ry NEDSo |
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DEBEao HUGG Ooo oUe oe eee SaeeeeeeoSs
sosgee EERE EEE EEE EEE HEE
ACoEee Bgnea oeoe ga05 a
| | mm 10 WW 12 13 4 15 /6

——-Length of petal
—— Maximum breadth of petal

7 mm. 4 STG 7
Breadth of tube Breadth of petal at base

COROLLA MEASUREMENTS. F, TYPE 9 X TYPE 51.

GABRIELLE L. C. HOWARD. 103
97 COROLLA:

As regards the corolla the following measurements were
made on both parents, the F, and F, generations in the cross
Type 9 x Type 51—length of the flower, length of the individual
petal, maximum breadth of the petal, basal breadth of the
petal, breadth of the corolla tube and breadth of the funnel.
The results observed were similar to those obtained in the leaf
characters. The intermediate nature of the F, was perfect
even to the method of pollination. The F, generation gave a
series covering the limits of both parents. The full data are not
given here as they present no new features, but a graphic
representation is given in Plate XXV. Want of time made
further measurements in the F; and F, generations impossible.

The difference in the colour of the corolla in this case
was probably due to two factors. The F, generation could be
divided into more than three groups, and those classified as
white formed one quarter of the whole, forty-five plants out of
117. The large number of gradations obtained in the F,
generations indicate that this is probably not a simple 3: 1
ratio, but that some of the palest pink combinations were
indistinguishable from white.

V. CONCLUSIONS.

The results obtained in these investigations may be briefly
summed up as follows :— 3

1. In any statistical investigation on the mode of inheri-
tance, the uniformity of the environment in which each set of
cultures is grown is exceedingly important. Comparisons
should not be drawn between cultures unless they are grown
close to one another with full precautions as to uniformity in
environment. By careful attention to cultural details it is
possible to reduce greatly the effect of environmental fluctua-
tions. The importance of using in such investigations only
normal, well grown plants cannot be over-estimated.

2. Parthenogenesis in N. tabacum, under the conditions
obtaining in hybridization work at Pusa, is negligible.

3. In all characters except height, the F, generation is
intermediate between the parents. In the case of the height,
different results were obtained in different crosses. This
may be due to added vigour in the hybrid plants. It is suggested
that the differences in the increase produced by this may depend
on differences in the number of dissimilar factors in the
parents.

4. In all cases the limits of variation in the F, generation
have been as great as those of both parents combined or have
exceeded these in both directions. In some cases, where the
parents and the F, generation were all alike, the variation in the
F, was very great. This can readily be explained by the
hypothesis that most of the factors possessed by the parents are
different.

5. Selected variates of the F, generation gave cultures
which differed in their range of variation from one another, and

GABRIELLE L. C. HOWARD. 105

often from the parents—the range of variation diminishing
with further selection. Certain cultures showed so small a
range of variation as to appear uniform; some of these
resembled the parent forms, some were new.

6. Observations on the time of flowering during four
generations resulted in the isolation of a culture flowering slightly
earlier than one parent, and others flowering much later, together
with some in which the range of variation was great.

7. It has been shown that although the heights of tobacco
plants may only differ slightly, nevertheless the factors on
which such heights depend may be almost all different. In one
cross, a new form (probably uniform) much shorter than the
shorter parent, has been isolated; in another cross, forms
resembling both parents were obtained. If there is a_ basal
condition of height common to all tobacco plantsit must be small.

8. The number of leaves per plant does not depend on the
height of the plant and is practically independent of the environ-
ment. The inheritance of this character can be explained by
a basal condition (of not more than nineteen leaves) common to
all types of N. tabacum, combined with independent factors
which can add to this number. These factors are probably
different in magnitude, that is, they represent an addition of
different numbers of leaves.

9. Distinct segregation has been observed as regards the
arrangement of the leaves on the stem. The arrangement with
internodes of equal length invariably breeds true.

10. The length of the decurrent portion of the lamina is
probably due to the existence of several factors. Hybridization
between plants whose leaves are equally decurrent has produced
forms with nondecurrent leaves. The differences in length due
to the various factors may be very small.

11. The most suitable leaves for measurements of the
lamina are those in the centre of the plant.

12. The venation of the leaves is one of the most constant
characters of the plant. On hybridization, the parent forms

106 STUDIES IN INDIAN TOBACCOS.

have been re-isolated in the F, and F, generations and also
constant forms with intermediate venation. Many of the
factors involved have a very small external effect.

13. The shape of the leaf in NV. tabacum may be defined by
the ratio length/breadth, position of the greatest width, amount
of indentation of the apex, amount of indentation of the base,
nature of the insertion, whether auriculate or not. All these
characters can be inherited independently of one another. By
the hybridization of two forms, in which the indentation factors
of the base differ, ‘“‘ petiolate ”’ forms which at once breed true,
are produced by the combined action of the factors. All
“ petiolate’ leaves in this species are probably sessile leaves
with deep indentations.

14. The irregularities of the surface of the leaves depend
probably on several factors. The undulation of the margin
in the particular case investigated proved to be due to a single
factor which is inherited independently of the factors concerned
in the surface of the leaf.

15. Measurements of the size of the corolla show that this
organ resembles the leaves in its mode of inheritance.

From the above results the following general conclusions may
be drawn :—The data obtained by a study of the characters of
N. tabacum show that there is no inherent difference in the mode
of inheritance of ordinary qualitative characters (such as the
colour of the corolla) and of those characters connected with the
size of the organs which are subject to fluctuating variability.
All the results obtained can be explained by the Mendelian
assumption of segregation of characters, combined with the
hypothesis that in connection with each character a large
number of factors exist, each of which can be inherited indepen-
dently. This conclusion is supported by the great range of
variation in the F, generation, the formation of extreme forms
in this generation far outside the limits of the parents, the differ-

GABRIELLE L. C. HOWARD, 107

ences and diminution in the range of variation in the F, cultures
raised from different variates of the F, generation and by the
isolation in the F, and succeeding generations of forms like
the parents and also of intermediate forms which breed true.
This isolation of new forms can easily be explained by a
rearrangement of the factors.

Pusa,
April 22nd, 1913.

APPENDIX.

DESCRIPTION OF THE TYPES USED IN
HYBRIDIZATION.

Type II, Plants late, tall; height 150 cm.; lower inter-
nodes short, upper ones long; most of the large leaves borne
near the ground; no large leaves in the upper two-thirds of
the plant. Leaves petiolate, petiole is slightly alate in the
lower leaves, more so in the upper ones; the wings are
decurrent down the main stem for about 2.5 em.; leaves inserted
at an angle of 90° and bend downwards from the top of the
petiole, asymmetric; shape varies from ovate to lanceolate
according to the position on the stem; venation acute-angled,
secondary veins arising at an angle of about 60°; apex acute ;
margin entire or slightly undulate ; colour blue-green ; texture
thick; average length of petiole 6 cm.; average iength of
lamina 49 cm.; ratio length/breadth 2.5. Inflorescence leaves
petiolate, petiole not alate, inserted at an angle of 60°—90°,
lanceolate; apex acuminate; margin generally entire, some-
time undulate. Inflorescence raised, side branches borne at
reguar distances up the stem, parallel to but not as long as the
main axis. Flowers a deep pink colour which does not fade
much; length 45 mm. Calyx slightly globular and inflated,
about one-third the length of the corolla ; teeth moderately long
and acute. Corolla with an orifice 8 mm. in diameter, a broad
tube, and the transition between the tube and the dilated
portion abrupt ; limb not very deeply divided with folds at the
junctions of the Jobe ; lobes very rounded at the base; apical
points short and somewhat reflexed. Capsule much longer than
the persistent calyx, conical ; apex blunt.

GABRIELLE L. C. HOWARD. 109

The anthers burst as the flower expands, not in the bud, and
at this period are above the stigma. In the fully open flower the
burst anthers are about 5 mm. above the stigma and project well
beyond the orifice of the corolla.

Type III. Plants very late, tall; height 150 cm.; lower
internodes very short, upper internodes long ; some of the lowest
leaves lie on the ground, the others are borne at long intervals
up the stem. Leaves petiolate with alate petioles, the wings
of the petiole expand on reaching the stem and are amplexicaul
and decurrent for 5em.; leaves inserted at an angle of about
60° and bend downwards ; shape ovate to cordate ; secondary
veins arise at an angle of more than 60°; apex acute; margin
undulate; leaf undulate; surface puckered; texture thick ;
colour dark blue green; average length of petiole 5 cm. ;
average length of leaf 41 cm.; ratio length/breadth 1.5.
Inflorescence leaves petiolate with very short alate petioles,
inserted at an angle of 90°, ovate; apex acuminate; leaf
undulate and surface generally puckered. Inflorescence with
few flowers and with very spreading sideshoots which arise at
regular intervals on the upper half of the main stem. The side
branches bear very few flowers. Flowers pink, the colour easily
fades ; length 42 mm. Calyx globular and inflated, less than one
third the length of the corolla; teeth moderately long and
acute. Corolla with a broad tube and short dilatation, diameter
of orifice 8 mm.; the transition between the tube and the ex-
panded portion abrupt ; limb not very deeply divided ; lobes
much rounded, pointed but with no distinct apical points.
Capsule much shorter than the persistent calyx, conical ; apex
blunt.

The anthers burst as the flower expands, not in the bud,
and occupy a position above the stigma. In the fully open
flower, the burst anthers are about 5 mm. above the stigma and
project much beyond the orifice of the corolla.

Type IX. Plants early, dwarf; height 104 cm.; lower
internodes very short, causing nearly all the Jarge leaves to lie on

110 STUDIES IN INDIAN TOBACCOS.

the ground. Leaves sessile, inserted at an angle of 90°, slightly
amplexicaul, lanceolate, ]amina much narrowed towards the base,
venation acute-angled, secondary veins arising at an angle of 50°;
apex acuminate, median leaves prolonged into very Jong thin
points; margin and lamina with deep undulations, Jamina
raised between the secondary veins, giving the appearance
of folds or ridges; colour dark green; texture very thick ;
average length 56 cm. Inflorescence leaves sessile, in-
serted at an angle of 90° and droop downwards from
the base, linear ; apex acuminate ; the whole leaf is very sinuate
and sometimes even twisted. Inflorescence conspicuous and
raised above the leaves, with numerous side shoots which are
almost as long as the main axis and not very spreading.
Flowers a very pale pink colour, short (36 mm.). Calyx tubular
with long and acute teeth, more than half as long as the corolla.
Corolla with a wide orifice (11 mm.), and a broad tube, the
transition between the tube and the dilated portion somewhat
gradual; limb divided to about half its depth; lobes rounded
at the base; apical points short, straight and only slightly
reflexed. Capsule cylindrical with a somewhat blunt apex ;
persistent calyx longer than the capsule.

In the unopened bud all the stamens are below the stigma.
The anthers burst just as the flower opens while they are still
below the stigma or at the most touch the underside with their
apices. In the open flowers the stigma is much above the burst
anthers and all project beyond the orifice of the corolla. In
some of the buds the stigma is visible between the still closed
lobes of the corolla.

Type XVI. Plants somewhat early; height 141 cm. ;
habit very open, internodes long, only two or three leaves lie on
the ground. Leaves sessile, inserted at an angle of 90° and
droop downwards from near the base, amplexicaul, sometimes
slightly auriculate, decurrent, lanceolate, lamina slightly
narrowed at the base; secondary veins arise at an angle of
50°; apex acuminate; margin entire; lamina flat except

GABRIELLE L. C. HOWARD. 1li

for occasional slight undulations at the base of some of theleaves;
colour very light green; texture medium ; average length
46 cm.; ratio length/breadth 4.1. = Inflorescence leaves
similar to the lower leaves but much narrower, in some cases
linear, and the undulations at the base are more marked.
Inflorescence raised with long side branches which are somewhat
parallel to and not as long as the main axis. Flowers few in
number, pale pink; length 50 mm. Calyx tubular, somewhat
inflated, about one-third the length of the corolla; teeth long
and acute. Corolla slender, with an orifice 8 mm. in diameter ;
tube slender, the transition between the tube and the dilated
portion slightly abrupt ; limb divided to about half its depth ;
lobes rounded at the base; apices very pointed but no apical
point. Capsule shorter than the persistent calyx, conical ;
apex pointed.

The anthers burst in the bud when level with the stigma.
In the fully open flowers the empty anthers are just above the
stigma and generally slightly project from the orifice of the
corolla.

Type XXIII. Plants somewhat early; height 136 cm. ;
leaves few, lower internodes short, causing some of the leaves
to be borne very near the ground, upper internodes long ;
inflorescence raised in a few long slender branches. Leaves sessile,
inserted at an angle of 90° and bend downwards from near the
base, amplexicaul, very slightly decurrent, elliptical, lamina
slightly narrowed towards the base ; secondary veins arise at an
angle of 75°; apex acuminate; margin and base of leat
undulate ; surface puckered ; leaf not fully expanded but folded
on the midrib ; colour yellowish green ; texture thick ; average
length 46 em.; ratio length/breadth 2.3. Inflorescence leaves
similar to the lower leaves but narrower. Jn/florescence raised
on a few, long, spreading branches. Flowers few, pale pink :
length 45 mm. Calyx tubular, somewhat inflated, a little less
than half the length of the corolla ; teeth moderately long and
acute. Corolla with an orifice 8 mm. in diameter; tube broad,

112 STUDIES IN INDIAN TOBACCOS.

the transition between the tube and the dilated portion somewhat
abrupt ; limb divided to about half its depth ; lobes not rounded
at the base ; apical points long and sometimes oblique. Capsule
shorter than the persistent calyx, cylindrical ; apex blunt.

The anthers burst in the bud or as the flower is expanding
and are then just above the stigma. In the fully open flower
the empty anthers and the stigma maintain their relative
positions and are level with the orifice of the corolla.

Type XXXV. Plants somewhat early, short with exceed-
ingly broad leaves; height 106 cm. ; internodes very short,
several leaves lie on the ground. Leaves sessile, inserted at an
angle of 60° afterwards becoming horizontal, auriculate, amplexi-
caul, decurrent for about 1 cm., the decurrent portion of the
lamina being very broad, elliptical ; secondary veins arise at an
angle of 80°; apex acute; margin with slight regular undula-
tions ; surface puckered; colour blue-green; texture thin ;
average length of leaf 48 cm.; ratio length/breadth 1.6.
Inflorescence leaves resemble the lower leaves in every particular
but are smaller. Inflorescence inconspicuous, scarcely raised
and much hidden by the large leaves ; side branches few, short
and spreading. Flowers large, a very deep pink colour which
does not fade; length 50 mm. Calyx globular, inflated, about
one-third the jength of the corolla; teeth short and obtuse.
Corolla with an exceedingly broad tube; diameter of the orifice
about 10 mm.; transition between the tube and the dilated
portion abrupt; limb entire with slight indentations between
the lobes which have no apical points. Capsule equal in length to
persistent calyx, broad, conical ; apex blunt.

The anthers and stigma are at the same level both in the
expanding bud and in the fully open flower. The anthers burst
just as the flower opens. Both anthers and stigma remain just
below the level of the corolla orifice.

Type XX XVIII. Plants somewhat late and tall; height
134 cm. ; internodes short, giving the plant a somewhat bushy
appearance, several large leaves near the ground. Leaves

GABRIELLE L. C. HOWARD. 113

sessile, inserted at an angle of 60°, the upper portions of the leaves
tend to become horizontal, amplexicaul, auriculate, decurrent,
elliptical, lamina only narrowed just at the base; secondary
veins arise at an angle of 70° ; apex acute ; margin with regular,
very small undulations; surface slightly puckered; colour
light green; texture thin; average length 47 cm.; ratio
length/breadth 1.8. Inflorescence leaves similar to the lower
leaves. Inflorescence not conspicuous and not much raised.
Flowers very pale pink in colour; length 45 mm. Calyx
globular, inflated, about one-quarter the length of the corolla ;
teeth moderately Jong and acute. Corolla with a wide orifice
(diameter 10 mm.), tube exceedingly broad and the dilated
portion very short, the transition between the latter and the
tube very abrupt (the shape of the corolla in this type is unique
among the Indian tobaccos); limb quite entire. Capsule
longer than the persistent calyx, conical ; apex pointed.

The anthers do not burst in the bud but as the corolla
expands, when the anthers are well above the stigma. In the
fully open flower the burst anthers are about 5 mm. above the
stigma and project from the orifice of the corolla.

Type LI. Plants somewhat late, very tall ; height 178 cm.;
internodes long, leaves borne at regular intervals up the
stem, none on the ground ; inflorescence not very conspicuous.
Leaves sessile, inserted at an angle of 60°, slightly amplexicaul
and auriculate, decurrent for 5 cm. or more, the decurrent
portion broad; shape elliptical, lamina somewhat narrowed in
the basal third of the leaf; secondary veins arise at an ang'e
of about 80°; apex acute; margin very slightly undulate and
recurved ; surface slightly puffy or puckered; colour dark
blue-green ; texture very thin; average length 43 cm.; ratio
length/breadth 1.8. Inflorescence leaves similar to the lower
leaves, but with more acute apices. Inflorescence with few
flowers; the side branches which are borne at the top of the stem
are few in number and almost level with the main axis and
parallel to it. Flowers very pale pink in colour, fading to

114 STUDIES IN INDIAN TOBACCOS.

white ; length about 45 mm. Calyx globular, inflated, less than
one-third the length of the corolla; teeth short and acute.
Corolla with a wide orifice 11 mm. in diameter and a very broad
tube, the transition between the tube and the _ dilated
portion very abrupt; limb entire but indented and somewhat
folded at the junctions of the lobes ; apical points very short.
Capsule longer than the persistent calyx, conical ; apex pointed.

In the bud before the corolla expands the anthers are found
just above the stigma and burst while in this position before the
bud opens. In the fully open flower the burst anthers are
about 5 mm. above the stigma and project from the orifice of the
corolla.

This type has an exceedingly good texture, the leaves are
much thinner than in the ordinary Indian tobaccos. It has the
great disadvantage, however, that the leaves are much exposed
to the wind and do not protect each other as in the more bushy
types. For this reason they are often badly torn before they
ripen.

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by ALBERT HowaArD, M.A., A.R.C.S., F.L.S.; and GABRIELLE
L. C. Howarp, m.a. Price, Rs. 9.

Studies in Indian Fibre Plants, No. 1. On two varieties of Sann,
Crotalaria juncea, L., by ALBERT HOWARD, M.A., A.R.C.S., F.L.S. ;
and GABRIELLE L. C. Howarp, M.A. Price, Re. 1.

The Influence of the Environment on the Milling and Baking Quali-
ties of Wheat in India, No. 1.—The experiments of 1907-08 and
1908-09. By Atspert Howarpb, M.A., A.R.C.S., F.L.S.; H. M.
LEAKE, M.A., F.L.S.; and GABRIELLE L. C. Howarp, M.A. Price,
Re. 1-8.

The Bud-Rot of Palms in India, by E. J. BurLer, M.B., F.L.s. Price,
Rs; 2:

The Economic Significance of Natural Cross-fertilisation in India by
ALBERT HOWARD, M.A., A.R.C.S., F.L.S. ; GABRIELLE L. C. Howarp,

M.A., and ABDUR RAHMAN Kuan. Price, Rs. 4-8.
8

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BOTANICAL SERIES - contd.

Millets of the Genus Setaria in the Bombay Presidency and Sind, by
G. A. Gamnig, F.L.s., Lnperial Cotton Specialist. Price, Re. 1.

Studies in Indian Fibre Plants, No. 2, On some New Varieties of
Hibiscus cannabinus, L., and Hibiscus Sabdariffa, L., by ALBERT
HOWARD, M.A., A.R.C.S., F.L.S.; and GABRIELLE L. C. HOwArRD,
M.A. Price, Rs. 3.

Notes on the Incidence and Effect of Sterility and Cross-fertilisation in
the Indian Cottons, by H. M. Leaxg, m.a. (Cantab.), r.u.s.; and
Ram PersHapD. Price, Re. 1.

Note on the Inheritance of Red Colour and the Regularity of Self-Fer-
tilization in Corchorus capsularis,—the common Jute Plant, by
I. H. BURKILL, M.A.; and R. S. FInLow, B.sc., F.c.s. Price, Re. 1.

Observations on certain Extra-Indian Asiatic Cottons, by H. M.
LEAKE, M.A., F.L.S.; and Ram PeEersHAD, Assist. to the Economic
Botanist, U.P. Price, Re. 1-8.

The Morphology and Parasitism of Rhizoctonia, by F. J. F. SuHaw,
BSc PACR CG LSee Mt S. mlerice: dusiens

The Inheritance of some characters in Wheat—I, by A. Howarp, M.A.,
A.R.C.S., F.L.S. ; and GABRIELLE L. C. HowARD, M.A. Price, Re. 1.

The Influence of the Environment on the Milling and Baking Qualities
of Indian Wheats, No. 2. The Experiments of 1909-10 and 1910-11,
by A. HowArbD, M.A., A.R.C.S., F.L.S.; GABRIELLE L. C. Howarp,
M.A., and H. M. LEAKE, M.A., F.L.S. Price, Re. l.

The Varieties of Soy Beans found in Bengal, Bihar and Orissa and
their Commercial Possibilities, by E. J. WoopHousg, M.A.; and
C.S. Tayitor, B.A. Price, Rs. 2.

On Phytophthora parasitica, nov. spec., by J. F. Dastur, B.se.,
Price, Rs. 2.

Studies in Peronosporacew, by E. J. BuTLER, M.B., F.L.S., Imperial
Mycologist; and G. 8. Kuikarnt, t.Ag., Mycological Assistant,
Bombay. Price, Rs. 2.

Notes on Pollination and Cross-Fertilisation in the Common Rice Plant
Oryza sativa, Linn., by G. P. Hector, M.A., B.sc. Price, Re. 1.

A Sclerotial Disease of Rice, by F. J. F. SHaw, B.sc. (Lond.), A.R.¢.s.,
F.L.S. (In the press.)

Studies in Indian Tobaceos. No. 3. The Inheritance of Characters
in Nicotiana tabacum, by GABRIELLE L. C. HOwarpb, M.A.
Price, Rs. 3.

Studies in Indian Cottons, Part I.—The Vegetative Characters, by
H. M. LEAKE, M.A., F.L.S.; and Ram Prasap. (In the press.)

CHEMICAL SERIES.

The Composition of Indian Rain and Dew, by J. Watrer LEATHER,
Ph.D., F.I.c. Price, Re. 1.

The Composition of the Oil-seeds of India, by J. W. LeatueEr, Ph.p.,
F.1.c. Price, Re. 1.

The Pot-Culture House at the Agricultural Research Institute, Pasa,
by J. W. LEATHER, Ph.D., F.1.c. Price, Rs. 3.

Experiments on the Availability of Phosphates and Potash in Soils, by
J. W. LEATHER, Ph.D., F.I.c. Price, Re. 1-8.

The Construction of Drain Gauges at Pisa, by M. H. Arnorr, M.INST.
C.E., with a Preface by J. W. LEATHER, Ph.D., F.I.c. Price, Rs. 3.
The Loss of Water from Soil during Dry Weather, by J. WauTmr

LEATHER, Ph.D., F.1.c. Price, Rs. 2.

The System Water, Calcium Carbonate, Carbonic Acid, by J. WALTER
LEATHER, Ph.D., F.1.c., and JATINDRA NATH SEN, M.A., F.C.S. Price,
Re: i.

Mol. - I, No

Vol. I, No.
Vol. I, No.
Vol. II, No

Vol. II, No.
Vol. II, No.
Vol. II, No.
Vol. It, No.
Vol. II. No.
Vol, III, No.
Vol. III, No.
Vol: I; No:
Viole ek No:
Vols i, No:
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Vols alle No:
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5 WARE

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CHEMICAL SERIES—contd.

Water Requirements of Crops in India, by J. WALTER LEATHER, Ph.D.,
HAE Cs ErICOwhseoe

The Nature of the Colour of Black Cotton Soil, by H. E, ANNETT, B.sc
(Lond.), F.c.S., M.S.E.A.c. Price, Re. 1.

Water Requirements of Crops in India—II, by J. Warrer LEATHER,
ph.p., F.1.c., Imperial Agricultural Chemist. Price, Rs. 2-8.

The Composition of the Milk of some Breeds of Indian Cows and
Buffaloes and its Variations, Part I, by A. A. Meaerrr, B.se. (Lond.),
and H. H. MANN, D.sc. Price, Re. 1-8.

Records of Drainage in India, by J. W. LEATHER, Ph.D., F.1.c., Price,
Re. 1.

The Rab System of Rice Cultivation in Western India, by H. H. Mann,
p.sc; N. V. JOSHI, B.A., B.Sc. L.ag.; and N. V. KanirKar, B.Ag.
Price, Re. 1.

The Composition of the Milk of some Breeds of Indian Cows and
Buffaloes and its Variations, Part II, by A. A. Meraarrv, B.sc.,
and H. H. Mann, p.se. Price, Re. 1-8.

A Contribution to the Knowledge of the Black Cotton Soils of India,
by W. H. Harrison, M.se., Government Agricultural Chemist,
Madras, and M. R. Ramaswami SIvAn, B.A., Chief Assistant to
the Agricultural Chemist. Price, Re. 1.

The Date-Sugar Industry in Bengal—an investigation into its,
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assisted by G. K. LELE, b.ag. ; and Buartat M. AMIN, B.A. Price,
Rs. 3.

Evaporation from a plain water surface, by J. WALTER LEATHER
Ph.D., F.I.c. Price, Re. I.

Studies in Chemistry and Physiology of the Leaves of the Betel Vine,
by H. H. Mann, p.sc.; D. L. SanAsRABUDHE and V. G.
PATWARDHAN. (Jn the press.)

ENTOMOLOGICAL SERIES.

The Bombay Locust, by H. M. Lrrroy, M.A., F.E.S., F.Z.S. Price,
Rs. 2-8.

The more important Insects injurious to Indian Agriculture, by H. M.
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The Indian Surface Caterpillars of the Genus Agrotis, by H. M. Lerroy,
M.A., F.E.S., F.Z.S.; and C.°C. GHOsSH, B.A. Price, Re. 1-8.

Individual and Seasonal Variations in Helopeltis Theivora, Waterhouse
with description of a new species of Helopeltis, by Harozp H.
MANN, D.se. Price, Re. 1-8.

The Coccide attacking the Tea Plant in India and Ceylon, by E. E.
GREEN, F.E.S.; and Haro~p H. Mann, p.se. Price, Re. 1.

The Mustard Sawfly, by H. M. Lerroy, M.A., F.E.S., F.Z.S.; and C. C.
GHOSH, B.A. Price, Re. 1.

The Rice Bug, by H. M. LEerroy, M.4., F.E.S., F.z.s. Price, Re. 1.

Remarks on Indian Seale Insects (Coccidw), by E. E. GREEN, F.E.S.,
F.z.S. Price, Re. 1-8.

The Red Cotton Bug, by H. M. Lerroy, M.4., F.E.S.,F.Z.S. Price, Re. 1.

The Castor Semi-Looper, by H. M. LErroy, M.A., F.E.S., F.Z.S._ Price,
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The Tobacco Caterpillar, by H. M. Lerroy, M.A., F.E.S., F.Z.S. Price
Re. 1-8,

The Cotton Leaf-Roller, by H. M. LEFRoy, M.A.,F.E.S., F.Z.S. Price,
Re. 1-8

Vol.

Vol.

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II, No.

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IV. No.

EVieNo-
IV, No.

IV. No.

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I, No.

I, No.

I, No.

I, No.

I, No.

II, No.

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Bons

ENTOMOLOGICAL SERIES—contd.

Notes on Indian Seale Insects (Coccidw), by H. Maxweri-LEerroy?
M-A.,F.E.S., 8.2.5. Price, Re. 1-8.

Life Histories of Indian Insects (Coleoptera—l), by H. MaxweE.t-
LEFROY, M.A., F.E.S.,F.Z.S. Price, Rs. 2.

Life Histories of Indian Insects—IT. Some Aquatic Rhynchota and
Coleoptera, by Dr. NowrosEexr, B.A. Assistant to the Imperial
Entomologist. Price, Re. 1.

Life Histories of Indian Insects—IIf. The Rhinoceros Beetle (Oryctes
Rhinoceros) and the Red or Palm Weevil (Rhynchophorus Ferru-
gineus), by C. C. Guosx, B.A., Asst. to the Imperial Entomologist.
Price, Rs. 2.

The Food of Birds in India, by C. W. Mason, M.s.E.A.C

., edited by
H. MaxweEtu-LEFRoy, M.A., F.E.S., F.z.S. Price, Rs. 7-8.

Eri Silk by H. Maxwett-Lerroy, M.a., F.E.S., F.z.S., Imperial
Entomologist ; and C. C. GuHosH, B.A., Assistant to the Imperial
Entomologist. Price, Rs. 3.

Tetrigine (Acridine) in the Agricultural Research Institute, Pasa,
Bihar, with description of new species, by J. L. HANCOCK, F.Z.S.
Price, Re. 1.

The Big Brown Cricket, by C. C. GHosu, B.A. Price, Re. 1.

Life Histories of Indian Insects (Hymenoptera), by G. R. DuTTA, B.A.,
Assistant to the Imperial Entomologist. Price, Rs. 2.

Inquiry into the Insecticidal action of some Mineral and other Com-
pounds on Caterpillars, by H. M. Lerroy, M.A., F.E.S., F.Z.8.; and
R. S. FINtow, B.se., F.c.s. Price, Re. 1-8.

BACTERIOLOGICAL SERIES.

Studies in Bacteriological Analysis of Indian Soils, No. 1, 1910-11, by
C. M. Hurcuinson, B.A., Imperial Agricultural Bacteriologist.
Price, Rs. 2-8.

Rangpur Tobacco Wilt, by C. M. Hurcuinson, B.4., Imperial Agr.
Bacteriologist. (In the press.)

VETERINARY SERIES.

Anaphylaxis in the larger Animals, by Major J. D., E. HOLMEs, M.A.,
p.se., 1.¢.v.D., Imperial Bacteriologist, Muktesar. Price, Rs. 2.

Salvarsan in the Treatment of Surra in Horses, Dogs and Rabbits, by
Major J. D. E. Hotes, M.A., D.sc., 1.c.v.D., Imperial Bacteriologist
Muktesar. Price, Re. 1-4.

Some more successful Experiments on the treatment of Surra in the
Camel with recommendations for Systematic treatment, by A. 8.
LEESE, M.R.C.V.S. Price, Re. 1.

Contribution to our Knowledge of the Immune Bodies oceurring in
anti-rinderpest serum and of the variations occurring in the serum
proteins of animals during Rinderpest and during immunization
and hyper-immunization, by Dr. P. Harriey, D.se. (Jn the press.)

Some cases of Surra treated in the field and in the laboratory during
the autumn of 1911, by Major J. D. E. HoLMEs, M.A., D.S¢., I.C.V.D.
(In the press.)

“BULLETINS ISSUED BY THE AGRICULTURAL RESEARCH

INSTITUTE, PUSA.

Notes on Cotton in Bihar in 1904, by H. M. LEFROY, M.A., F.2.S., ¥.Z.8., Imperial
Entomologist. Price, As. 4 or 6d.

An Outbreak of Cotton Pests in the Punjab, 1905, by H. M. Lerroy, m.A., F..8.,
¥.z.S., Imperial Entomologist. _ Price, As. 4 or 6d.

The Extension of Jute Cultivation in India, by R. S. Frytow, B.sc., F.0.s., Jute
Specialist to the Government of Eastern Bengal and Assam. Price, As. 12 or
1s. 2d. (Out of print.) %

First Report on the Fruit Experiments at Pisa, by A. Howarp, m.a. (Cantab.),
A.B.0.8. (Lond), ¥.t.s., Imperial Economic Botanist. Price, As. 6 or 6d.

Report on Trials of the South African Locust Fungus in India, by E. J. Burizr,
M.B., F.1.S., Imperial Mycologist; andH.M. Lerroy, M.A.,¥.E,S., F.z.S., Imperial
Entomologist. Price, As. 2 or 3d.

The Ticks Infesting Domesticated Animals in India, by C. WARBURTON, M.A.g
Zoologist to the Royal Agricultural Society of England. Price, As. 4 or 6d.

A Preliminary Account of the Biting Flies of India, by H. M. Lerroy, M.A., F.z.s.,
¥.z.8., Imperial Entomologist, Price, Re. 1 or ls. 6d. (Owt of print.

Official and Recommended Methods for use in Chemical Laboratories of the
Departments of Agriculture in India, by J. Watrer LEATHER, Ph.D., F.I.C.,
Imperial Agricultural Chemist. Price, As. 4 or 6d.

Report on Cocoanut Palm Disease in Travancore, by BE, J. BuTLeER, M.B., F.L.S.,
Imperial Mycologist. Price, As. 6 or 6d.

Treatment and Observation of Crop Pests on the Pisa Farm, by H. M. Lzrroy,
M.A., F.E.S., F.Z.8., Imperial Entomologist; and C.S. Misra, B.a. Price, As. 6
or 7d.

On Flax Dodder, By A. Howarp.. M.A., A.RB.C.S., F.L.8., Imperial Economic
Botanist. Price, As. 4 or 6d.

The Making and Care of Lawns in India, by A. HowarpD, M.A., A.R.C.S., F.L.S.,
Imperial Economic Botanist. Price, As. 4 or 6d.

Sugarcane at the Partabgarh Experimental Station, by G. CrarKE, F.1.c., Agri-
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The Milling and Baking Qualities of Indian Wheats, by A. HowArp, M.A., A.R.0.8.,
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Newnham College, Cambridge. Price, As. 4 or 6d.

Note on the Extension of Cultivation of Fibre Plantsin India. Price, As. 6 or 8d,

Second Report on the Fruit Experiments at Pisa, by A. Howarp, M.A. (Cantab.),
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The Milling and Baking Qualities of Indian Wheats, No. 2, by A. Howarp, M.a.
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L. C. Howarp, M.A.,'Associate and late Fellow of Newnham College, Cambridge.
Price, As. 6’or 8d. . -~

Report on the Outbreak of Blister-Blight on Tea in the Darjeeling District in
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List of Names used in India for Common Insects, compiled in the Laboratory of
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Memorandum on Indian Wheat for the British Market, by Sir Jamzs Witson,
K.¢.s.1. Price, As, 4 or 6d.

Memorandum regarding Leading Eucalypts suitable for India, by F. Bootn-
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The Milling and Baking Qualities of Indian Wheats, No. 3, by A. Howarp, m.a.,
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Personal Assistant to the Imperial Economic Botanist. Price, As. 7 or 8d.

Insecticides-Mixtures and Recipes for use against Insects, in the Field, the
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Imperial Entomologist. Second edition revised with many additions by T.
BAINBRIGGE FLETCHER, R.N., F.E.S., F.Z.S., Officiating Imperial Entomologist,
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Nt) a Pee wee ae Ve et ES Bact RI ER als SA ee RT Dk DE We ee RR TSB ei A et ee ee ee STAR
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BULLETINS ISSUED BY THE AGRICULTURAL RESEARCH
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No. 24. The Indian Saltpetre Industry, by J. W. Learner, ph.p., F.1.c., Imperial Ade. ;
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Imperial Agricultural Chemist.. Price, As. 8 or 9d.

No. 25. Report on the Flax Experiments conducted at Dooriah in 1910-1911, by BE. M.
VANDEKERKEHOVE, Flax Expert to the Bihar Planters’ Association, Price, |
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No. 26. Note on the Present Position of Cotton Investigation in India, by BueNagp.
Coventry, Offg. Inspector-General of Agriculturein India. Price, As. 2 or 3d.
, No. 27, Experiments on the Cultivation of Sugarcane at the Partabgarh Experimental
vt Station, 1909-11, by G. CrarKkE, F.t.c.; H. E. ANNETa, B.sc; SyED ZAmmN
HUSSAIN, B.A. Price, As. 5 or 6d.

No. 28. The Cultivation of Lacin the Plains of India, by C, S. Misra, B.a., First Assistant ay
to the Imperial Entomologist. Price, As. 8 or 9d. Oy

No. 29. Directions for the Cultivation of Eri Silk. Price, As. 3 or 4d.
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. No. 31. Wheat Experiments on the Botanical Area, Cawnpore, and their bearing on Wheat
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an Assistant to the Economic Botanist. Price, As. 3 or 4d.. Nea)

No. 32. A note on some interesting results following the internal Administration of Arsenio.
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No. 33. Some Aspects of the Agricultural Development of Biba. by A. Hiwatt, M.A,
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No. 35, Report on the Flax Experiments conducted at Dooriah during the year 1912-13,
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ant Wie No. 36. Note on the MacFadyean Stasinine reaction for Anthrax Bacilli, by Major J. D. E.
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~ Memoirs of the
Department of Agriculture

ra

"STUDIES IN INDIAN COT TONS

| PART 1
THE VEGETATIVE CHARACTERS

4 BY
~H. MARTIN LEAKE, M.A., F.L.S

Economie Botanist to Government, United Provinces
. AND |

RAM PRASAD

Assistant to the Economie Botanist

AGRICULTURAL RESBARCH INSTITUTE, PUSA

PUBLISHED FOR
fh THE IMPERIAL DEPARTMENT OF AGRICULTURE IN INDIA

BY
‘THACKER, SPINK & CO., CALCUTTA

OW. THACKER & CO., 2, Crezp Lanz, LONDON

February, L914. BovrANICAL SERIES Vol. VI, No. 4

MEMOIRS OF THE
DEPARTMENT OF AGRICULTURE
IN INDIA |

STUDIES IN INDIAN COTTONS
PART I
THE VEGETATIVE CHARACTERS
e4
H. MARTIN LEAKH, M.A., F.L.S
Economic Botanist to Government, United Provinces
AND

RAM PRASAD

Assistant to the Economic Botanist

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SS = ~
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R $5 Sy y

AGRICULTURAL RESEARCH INSTITUTE, PUSA

PUBLISHED FOR

THE IMPERIAL DEPARTMENT OF AGRICULTURE IN INDIA

BY

THACKER, SPINK & CO., CALCUTTA
W. THACKER & CO., 2, Creep Lanr, LONDON

. CALCUTTA :
PRINTED BY THACKER, SPINK & CO,

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Text
Tables
Maps

Plates

CONTENTS.

PAGEs.

115—141

APR 29 1914

(1) Jour. Asiatic Soc. of Bengal,
Do. do. do. Vic pe 23:
(3) Proc. Roy. Soc., B. 83, p. 447.
(4) Journal of Genetics I (ii), 3.

(5) Memoirs of the Dept. of Agri. in India, Botanical Series,

STUDIES IN INDIAN COTTONS
PART I
THe VeGEerativE CHARACTERS

BY

H. MARTIN LEAKE, M.A., F.L.S.

EHeonomic Botanist to Government, United Provinces
AND

RAM PRASAD,

Assistant to the Economic Botanist.

THE present note is intended to bring up to date the results of
the experiments in the improvement of the cottons in the United
Provinces of Agra and Oudh.

bY; No.3.

The work which has been in steady
progress since the autumn of 1904, was commenced at Saharanpur
and continued since 1906, on the Botanical area of the Govern-
ment Agricultural Station, Cawnpore.
number of publications based on this work have been made and
are here noted :—

During this period a certain

Woaw pails:

116 STUDIES IN INDIAN COTTONS

(6) The Problem of the Improvement of Cotton m the United
Provinces of Agra and Oudh. Pamphlet published in
conjunction with Dr. Parr, Deputy Director, Western
Circle, in connection with the U. P. Exhibition,
Allahabad, and reprinted in the Agricultural Journal
of india Vol! VI, Pt.

(7) Memoirs of the Dept. of Agri. in India, Botanical Series,
PY, Novo:

In addition, an article of a more general nature on the breeding
of cotton has been written for Prof. Fruwirth’s “ Die Ziichtung der
landwirtschaftliche Kulturpflanzen.”

Of the above. No. 4 contains the fullest detail of the experi-
ments. but is limited to those problems of which the scientific
aspect only had been fully developed ; moreover, in it the results
are only carried down to the I, generation. The economic aspect
received little notice and has only been treated in briefest outline
in the pamphlet (6) and from a very special point of view in the
recent memoir (5).

The work divides itself broadly into two sections, the Scientific
and the Economic, the former dealing with all those points con-
cerning the characters, and behaviour of these, which it is necessary
to ascertain to indicate the most promising lines on which to develop
the economic work and the conditions under which this work must
be carried out, the latter, with the actual experiments undertaken
to effect the desired improvement.

The first step in any work, such as the present, is to ascertain
the extent of the material at hand for the purpose of experiment,
or, in other words, using the plant as a unit, to determine the
number of forms or types it is possible to isolate and cultivate in a
condition of purity. Throughout India there are cultivated a
series of cottons including numerous and diverse forms. The
grouping of these into a systematic series of which the unit is the
type, or that form of plant which can be recognised as distinct and
remains thus distinct when cultivated under conditions which
ensure purity, is the first object and the one which forms the basis
for subsequent experiments,

LEAKE AND RAM PRASAD 117

Having ascertained the types it is next necessary, by a com-
parative study of the points or characters, by which these differ,
to determine the main characteristics. The unit of this series of
observations is now the character. But the investigation is not
limited to the determination of these alone; by crossing types
which differ in certain definite characters and tracing the be-
haviour of these in the offspring, it is possible, not only to ascer-
tain with some clearness what constitutes a unit character but,
also, to obtain information as to the behaviour of these units under
definite conditions—information which permits efforts aimed at
improvement to be undertaken with some confidence.

The plant may be imagined to consist of an aggregate of unit
characters,—the presence or absence of an ultimate common base
is not here a matter for concern c.f. de Vries (11)-~certain combi-
nations of which are to be found in nature. Improvement consists
in a rearrangement of these units with the production of a plant
more suited to the conditions of growth and having a more valu-
able produce. Regarded in this light the possibility of effecting
any Improvement is seen to depend very largely on the clearness
with which the objective receives definition in the mind. In this
way only can the most suitable method of obtaming that combi-
nation, and the most suitable types to use as parents, be selected
with any degree of certainty.

Although the plant forms the central object of study it is not
possible to limit observation to this one aspect. The inter-action
between the plant and its surroundings is intimate. It is further
necessary, therefore, to determine the nature of the surroundings
or environment and also, if possible, the conditions which render
one type more suitable than another for growth under a particular
set of conditions. In the broadest sense the environment includes
not only the physical conditions by which the plant is surrounded
but also the organic. In other words, there exists an inter-action,
not only between the plant and the soil, climate and such physical
conditions of growth but also between the plant and its neighbours
—in the case of a field crop usually plants of the same species—
insects, harmful or otherwise, and organisms causing disease,

118 STUDIES IN INDIAN COTTONS

This line of investigation has not in the present case received
more than general attention. The physical conditions as existing
in the province at the present time have been accepted as standard.
It is unlikely that any large change either in methods of cultivation
or in the extended use of manures will take place in the near future.
The main object seems, therefore, to be rather the production of an
improved plant under these conditions than the adaptation of the
conditions to meet the needs of the plant. The organic conditions
in like manner have not formed the subject of especial investigation
except in as far as they affect the methods of experiment. Of
major importance here 1s the relation of the plant to its neighbours
and to insects. On this relation depends the extent of the occur-
rence of cross-fertilization in the field. This subject has been treated
somewhat fully in a previous memoir (20) and will not receive more
than a passing reference here.

Before entering in detail into an account of the experimental
work which has been undertaken it will not be amiss to review
briefly the conditions under which cotton is grown in the Provinces.
These Provinces occupy the large alluvial tract lymg between the
Himalayas on the north and the uplands of Central India. Through-
out the entire area cotton is grown, but the relative area under the
crop varies considerably. In map A is given the percentage of the
kharif area under cotton for each district, the figures being derived
from the Season and Crop Report for the year 1909-10, in which
the cotton crop was 108 per cent. of the normal. From this map
it is seen that the largest proportionate area under cotton occurs
in the Muttra district at the south-west corner of the Province.
Here over 30 per cent. of the kharif area is sown to cotton. From
this centre the percentage of the kharif area under the crop gradu-
ally decreases in passing eastwards until, in the east, the crop is
practically absent, the total acreage in the Ghazipur district being
3 only. From the shading of the areas showing a certain range of
percentages it further appears that, were the units of determination
smaller than a district, a series of lines of equal percentages could
be drawn which would be found to be V-shaped with their apices
pointing towards the east. In map B are given the isotherms

LEAKE AND RAM PRASAD 119

(average mean temperature) for the months June—August, and in
Map C the lines of equal rainfall for June to October. The similarity
between the direction of these lines and those indicated for equal
percentages of the cotton crop is somewhat striking. It seems to
follow that,as far as the United Provinces are concerned, the climatic
(rainfall and temperature), rather than any other, conditions are
the principal factors in determining the percentage area of cotton
grown in the various districts. A problem of somewhat different
nature refers to the conditions affecting the area sown to cotton
in the different seasons. The statistical aspect of this problem has
been dealt with by Moreland (25), by whom also reference is made
to the aggregation of the crop into certain areas or cotton tracts.

The Genus Gossypium and the Types used in the investigations.

The distribution of the genus Gossypium throughout tropical
and semi-tropical climates 1s nearly universal, the cultivation of
the numerous forms extends back to the earliest times, and the
origin of these forms, as with so many agricultural plants, is lost in
antiquity. Owing, moreover, to the commercial value of the crop
and to the amount of attention which has been devoted to it
from the time that the fibre began to be used as the raw product
of an ever-increasing industry, the original limits of cultivation of
each form have been broken down so that at the present time it is
possible to find im such a country as India examples of all the larger
groups of the genus. In many cases the history of these intro-
ductions is wanting and the presence of the plant alone bears testi-
mony to the fact. In spite of this intermingling of types, however,
it is still possible to recognise two fairly marked groups of culti-
vated forms which may be termed the New and Old World groups
respectively. Perhaps the most constant character by which these
two groups can be distinguished is to be found in the bracts which.
in the former, are free and, in the latter, united. It is impossible
here to enter into a revision of the genus, though there is little doubt
that such a revision is necessary. Such revision can only be made
after personal study of all the forms in the field, opportunity for
which has not been forthcoming. As so commonly occurs in plants

120 STUDIES IN INDIAN COTTONS

cultivated over wide areas there arises a considerable degree of adap-
tation to particular conditions. and no single area will be found in
which the numerous forms can be cultivated under conditions
which adnut of successful study. Even among the Indian indi-
genous forms certain occur which, owing to the lengthened period
of vegetative growth, it is impossible to study in full detail under
the chmatic conditions prevalent at Cawnpore.

While. therefore, opportunity for such revision is lacking, it Is
possible to indicate reasons which would seem to throw doubts on
the present systems. The two most fully developed are those of
Todaro, published in 1877 in his “ Rel. sulla Cultura dei Cotoni in
Italia”, and by Watt in 1907 in his “ Wild and Cultivated Cottons of
the World.” The former classification is based on the observation
of living plants grown in Italy. The chief defect is the failure to
distinguish clearly between the two types of branches —especially
the secondary branches. These may be monopodial or sympodial.
The value of this particular character in a systematic scheme may
be open to some doubt, but the economic importance of clearly
recognising the two types is indisputable. This point will be re-
ferred to in some detail later. In the latter, this question of habit
is again relegated to a position of insignificance. The author has
an extended knowledge of the Indian cottons from personal expe-
rience and he has made an exhaustive study of the material col-
lected in the vast majority of the herbaria of the world. We are
tempted to think that, in a plant like cotton, the material thus pre-
served may be very misleading. It is not too much to say that.
owing to this difference in habit, two totally distinct types might be
represented in a herbarium by material showing no recognisable
differences. Thus the secondary branch of a sympodial type may
appear identical with the tertiary branch of a monopodial type and
unless the exact position from which it has been taken has been
noted it will be impossible to determine the type to which it belonged.
The author attaches considerable importance to the presence or
absence of a fuzz in addition to the floss of the cultivated forms.
Thus his section II is characterised as Fuzzy-seeded cottons with
united bracteoles: his section Ill as Fuzzy-seeded cottons with

LEAKE AND RAM PRASAD LAR

free bracteoles, and his section IV as naked-seeded cottons with the
bracteoles free or nearly so. We have recently been in a position
to grow and examine a series of cottons from China and among
these occur a series of forms which are undoubtedly related to the
members forming the author’s section II, but the seeds of which
are quite naked (21).

From the limited number of general observations we have
been in a position to make, it would appear that among the culti-
vated forms the condition of the bracts is a character of consider-
able systematic importance. The two groups so separated agree in
the further point of their geographical distribution. The origin of
most of the cultivated forms is, it is true, unknown or speculative ;
moreover the numerous efforts to acclimatise exotic forms in various
regions of the globe have led to extensive intermingling of types.
Nevertheless the group of plants with united bracteoles appear to be
typical of the Old World, while that of plants with free bracteoles is
similarly typical of the New World, though the natural limits of
this group are not so well defined as those of the latter.

The various types used in the course of these earlier stages of
these investigations have been given in a previous publication by
one of us (22). As has been there stated, the difficulty of culti-
vating many, and especially the monopodial, types renders a com-
plete survey of the genus at present impossible.

The list of types there given may be here quoted.

Monopodial Types.

Perennial ; secondary branches ascending sharply at an acute
angle. Leaf factor is less than 2; plant almost glabrous. Brac-
teoles small, triangular ; margin entire or dentate. Corolla yellow.
(Plate 1).

This plant is the G. obtusifolium-Roxburgh Flora Indica of
Gammie (15) and Watt (32). The various forms to which the spe-
cific name obtusifolium has been given at different times have
been dealt with by Burkill (5)... A. .. Type 1.

Perennial; with secondary branches spreading. Leaf with
a factor less than 2. Stem and leaves densely covered with short

122 STUDIES IN INDIAN COTTONS

hairs. Bracteoles deeply auriculate or reniform, deeply serrate,
spreading in fruit. Corolla yellow, petals small. Stigma heavily
glandular. Capsule inflated and nearly spherical with a sharp
mucronate apex. (Plate II).

This plant is the G. herbaceum, Linn, of Todaro (28) and
Gammie and the G. obtusifolium var. Wightiana of Watt
(32). ar ie 7 Type 2.
Perennial ‘‘ tree cotton; ’’ secondary branches ascending
sharply at an acute angle. The entire plant of a deep-red, or
purple colour. Leaf with a factor greater than 3; frequently
with an extra tooth on one or both sides of the central lobe.
Bracteoles small, triangular ; margins entire or with the tip dentate.
Corolla deep-red. Stigma eglandular. Capsule usually 3 celled,
ovate. (Plate III).

This plant is the Gossypium arboreum of Linn. Sp. PL ;
Parlatore (26) ; Todaro (28); and the G. arboreum type of Gammie
(15) and Watt (32)... LG st | Lypevs-

Sympodial Types.

Annuals with a few only, or none, of the lowest secondary
branches monopodia, the remainder sympodia; the monopodial
branches ascending and the sympodial spreading.

A tall plant, in later stages drooping under the weight of fruit.
Leaf large, with factor less than 2; lobes commonly 3 or with 2
small accessory basal lebes. Young stem and leaves sparsely hairy.
Bracteoles small, entire or with few small apical teeth, closely
enveloping bud and fruit. Corolla yellow with deep red “‘ eye.’’
Petals large, semi-transparent. Stigma eglandular or with few
glands only. Capsule commonly 3 celled, ovate. (Plate IV).

This plant is the Gossypium indicum, Lamk. of Gammie (15)
and G. Nanking var. bani of Watt (32). es .. Type 4.

An erect plant, in later stages drooping under the weight of
fruit. Leaf factor less than 2; lobes 5-7. Young stem and leaves
hairy. Bracteoles large, entire or with few small apical teeth
loosely enveloping bud and in fruit sometimes reflexed. Corolla
yellow with deep red ‘* eye; *’ petals opaque. Stigma eglandular

LEAKE AND RAM PRASAD 123

or with few glands only. Capsule ee 3-4 celled, ovate.
(Plates V & VI). i :.abype 5.

An erect plant differing deat Type 4 in the greater rigidity
of the main stem and the greater angle at which the secondary
monopodia arise, in this case about 45, and in the corolla: which is
white. The petals are small, scarcely projecting beyond the brac-
teoles. (Plates VII & VIII). .. sic Ly pe: 6.

Plant erect with secondary naencnidial branching, when
developed, sharply ascending. Leaf factor less than 2 ; flower white.
This type differs from the last in two respects. The secondary
monopodial branches, if developed, are sharply ascending. Fre-
quently, however, they are absent, and even when present reduced
in number in plants where the growth of the main axis has not
received a check, to one, or at most, two with vigorous growth.
The plant is consequently strongly asymmetrical. For the same
reason the length of the vegetative period is very brief and the
first flowers develop while the plant is still quite small. Growth
continues throughout the season, the plant maintaining a mar-
vellous fertility. (Plate IX). zs .. Type 7.

A tall plant, in later stages drooping ences the weight of fruit.
Leaf factor greater than 3; lobes 5-7 with an extra tooth, on
one, or both sides of the central lobe frequently developed.
Young stem and leaves hairy. Bracteoles entire or with few
apical teeth. Corolla yellow with deep red ‘“‘eye.’’ Stigma
eglandular or with few glands only. Capsule 3-4 celled, ovate.
(Pleo Xe)... - .. Type 8.

A plant differing cam (8) in atte colour of corolla only which
is white and scarcely protrudes beyond the bracteoles .. Type 9.

Types (4)-(9) fall into the G. neglectum and G. roseum of
Todaro (28), the G. neglectum, Tod. of Gammie (15) and the G.
arboreum vars. neglecta and rosea of Watt (32).

A tall plant with the main stem weak and early drooping.
Leaf factor greater than 3; lobes 5-7. Bracteoles entire or with
few apical teeth, large and continuing to grow with the developing
boll. Corolla pale yellow with deep ‘‘ eye.’’ Stigma eglandular.
Capsule ovate, very large with numerous seeds. (Plate XI).

194 STUDIES IN INDIAN CO'TTONS,

The plant is the G. cernwum of Todaro and Gammie and the @.
arboreum var. assamica of Watt (32). at .. Type 10.
A tall plant with leaf factor greater than 3 ; leaf lobes 5-7,
stem and leaves of a deep red or purple colour; bracteoles entire
or with few apical teeth. Corolla with deep-red ‘‘ eye,’’ petals
white, tinged with pink along margin and the portions exposed
in the bud. (Plate XII). |
This plant is the G. sanguineum Hassk. var. minor of Gammie
(15) us oy Be ne . Type.
In addition to the above, the following forms have since been

subjected to experiment.

Monopodial Types.

Plant tall with long monopodial branches sharply ascending.
Leaf with a factor greater than 3, wrinkled. Leaf and stem covered
with short hairs. Bracteoles triangular with margin dentate.
Flowers yellow or white.

This group appears to contain a number of type which, however,
owing to their monopodial habit it has been impossible to isolate
with certainty. The commonest form possesses the habit of Type 3
(G. arboreum, Linn.), which it also resembles in the form of the
bracts, while the wrinkled leaf and presence of short hairs give a
marked resemblance to Type 2(G. herbaceum, Linn). The relatively
large value for the leaf factor measurement (/) gives the leaf a
characteristic appearance which is frequently obtained in the
ofisprmg of the cross Type 2 x Type 3. The group is grown
usually as a mixed field crop in the east of the United Provinces
and in Bengal.

The types appear to differ chiefly in the density of the hairs
on leaf and stem and in the length of the internode and of the
petiole. The group comprises the G. intermedium of Gammie

(15).
Sympodial Types.

(1) Plant small with monopodial secondary branches few
or none. Leaves 3-5 lobed with factor less than 2. Flower

LEAKE AND RAM PRASAD 125

vellow. Boll large. This group includes a series of types received
from China, of which the following have been employed :

Corolla, yellow eved ; seed with fuzz wont sbypeyken
naked J ype 143:
without eye ; seed naked 27 Type 4.

(2) Plant differing from Type 2 mainly in the character of
the secondary branches. The group includes a complex series of
forms from Persia which has been dealt with by us elsewhere (21).
The simplest form is the true G. herbaceum of Todaro.

(3) Gossypium hirsutum, Linn.—This plant, which vields the
bulk of the American cotton crop, has formed the basis of numerous
experiments both in the United Provinces and in other cotton tracts
of India and is to be found locally throughout the former. Selected
plants from cultures derived from various sources have been
used.

(4) Gossypium Stocksii, Max. Mast. A wild form of Gossy-
pium found on the limestone hills around Karachi from whence

seed was obtained.

Pollination.

The cotton flower is hermaphrodite (Plate XII1). The stigma
becomes receptive and the stamens liberate their pollen shortly
after the flower opens. These organs are so distributed that self-
pollination follows almost immediately. Associated with this
arrangement of organs we find that in almost every case self-ferti-
lization is effective. A detailed discussion has been given on a
former occasion (20).

When, however, the question of effecting cross-fertilization is
considered, the genus Gossypium, at least as far as the cultivated
forms are concerned (for of wild species G. Stocksii, M. Mast. alone
has been studied), appears to fall into two marked groups, the
members of each of which are fertile /vter se, but exhibit complete
sterility when attempts are made to effect a cross between the two
groups. These two groups have already been referred to above as
characteristic of the Old and the New World and as respectively
characterised by united and free bracts.

126 STUDIES IN INDIAN COTTONS

The Experiments (A).—The Colour of the Corolla.

The corolla of the cotton flower consists of five petals arising
from the base of the staminal column. The petal is frequently,
and in the true Indian cottons invariably, possessed of a deep red,
or purple eye situated onthe claw. In the extra-Indian cottons
this eye may be absent, and in this case the petal is self-coloured.
The presence or absence of the ‘‘ eye ’’ appears to be independent
of the general colour of the petal and requires to be treated
separately.

Omitting for the moment consideration of the eye, the petal
may be yellow, white or red. The latter colour is present in Types
3 & 11 only and is not limited to the petals alone, but is here a local
manifestation of the presence of a red sap colour which occurs
throughout the plants of these types and will be more conveniently
treated in detail in a following section (C).

The two petals, then, may be either white or yellow, and of
the latter type two forms are readily distinguished. In the one
the colour is a full yellow, described in the subsequent text simply
as © yellow,’ while in the other, found only in Type 10, the colour is
very light—a condition which will be denoted below as ‘ pale-yellow.’

The yellow colour is dependent on a colour—producing factor
which, according to its presence or absence, gives rise to a simple
pair of alellomorphic characters, of which the presence of the colour—
producing factor is dominant in both cases. That is, the yellow
is dominant both to pale-yellow and to white.

In Table I are given the results obtained from a cross between
Type 8 (yellow) and Type 10 (pale-yellow), while in Table II is given
the offspring from a cross between Type 4 (yellow) and Type 6
(white). In both cases the numbers are small and insufficient to
give reliable numerical values to indicate the proportion existing
between the pairs of characters. A clearer indication is given in
Table III, which is derived from the crosses between Type 3 and
Type 10 and between Type 3 and Types 7 & 9. These crosses are
dealt with in fuller detail below, since, owing to the presence of
the red colouring matter found in the sap of Type 3, a pair of
factors 1s here involved.

LEAKE AND RAM PRASAD 127

Type 10 has in no case been crossed with a type possessing a
white petal nor, in the constant propagation of this type from
natural seed, has a single natural cross with a white flowered type
been recognised in spite of the fact that it has been each year culti-
vated in close proximity to Type 9. This is the more remarkable
in that. when self-fertilization is not resorted to. there occur a con-
siderable number of plants with the full yellow petal, which are pre-
sumably natural crosses with a yellow flowered parent, though
no such parent is growing in the same proximity as is the case with
the white flowered type. For the present. therefore, it is impos-
sible to say what relation exists between the two conditions which
may be indicated by the presence and absence of the pale-yellow
factor. This point is now under investigation. — It is clear, however,
that this condition does not correspond with that denoted by Balls
(2) as lemon yellow which he found to represent the unpure stage
in the cross between full yellow and white flowered parents in the

Keyptian cottons.

(B).—The Eye of the Petal.

As has been noted under the description of types all true
Indian cottons possess an eye situated at the base of the petal.
In some of the forms received from China, however, the petal is
self-coloured yellow. These cottons exhibit complete fertility. in
the cross with the Indian types and the combined group, therefore,
Is, In respect to this character, comparable to the Hgyptian series
of cotton in which the eve may or may not be present. Balls
(2) has shown that in the latter group the heterozygote is repre-
sented by plant in which the intensity of the eye colour is much
reduced. In the examples we have observed such an intermediate
condition appears to be rare or non-existent. At the present time
crosses in which the eyeless form figures as parent have not been
varied sufficiently far—the eyeless type being only recently
acquired—to supply definite information of the behaviour of this
character. In the F, generation of a cross between this form and
Type 3 the eye is fully developed, but it is possible that the red sap
colour of Type 3 may act as a masking character. Indirect evidence

128 STUDIES IN INDIAN COTTONS

is given in Table IV. From this table it is seen that, of 15 eyed
parents grown under conditions admitting of cross-fertilization
taking place naturally. 14 have given eyed forms only and are
therefore pure. One only has given eyeless plants in numbers
which closely approximate to expectation on the supposition that
the parent is impure and presence of the eye is fully dominant over
absence. Two only of the eyeless parents have bred pure. Of the
200 offspring of the remaining 4 plants 19 have the eye fully devel-
oped, while in the remainder it is absent, a number readily accounted
for by the amount of cross-fertilization known to occur, but incom-
patible with any simple Mendelian ratio. In a single case only
has a form appeared in which the intensity of the eye is definitely
reduced. This form is bemg submitted to further investigation.*
It would appear. therefore, that the intermediate form of eye found
by Balls does not occur in the present group.*

(C)..-The Red colouring-matter in the Sap.

Type 3 and Type 11 are: characterised by the presence of a
red anthocyanic colouring-matter in the sap which imparts to the
entire plant, stem, foliage and flower, a deep red colour. In these
types the intensity of the red in the petal is sufficient to mask the
true petal colour entirely, and it is only ina few cases of diseased
flowers and of such as open out of season that the presence of
the yellow petal colour in Type 3 can be determined by direct
observation. When a plant. which breeds pure to this character,
is crossed by one in which the red colour is absent, the F,
generation bears the red colour which may be said to be domi-
nant.

The intensity of the red colour is, however, definitely dimi-
nished and the petal attains a condition which has been denoted
in the case of a cross with a yellow flowered plant as * red on yellow.’
It is consequently possible to say, from an examination of the petals
of plants derived from such a cross, whether the plant is pure with
regard to this character or not. From an examination of Table V it

condition, It must, therefore, represent a further type.

LEAKE AND RAM PRASAD 129

appears, however, that the determination so made is not invariably
correct and the method is subject to an error of 22 per cent.
A more accurate method has been found for this determination in
the young leaves. The red colouring matter in the leaf, from the
time this first opens until full development is reached, has been
found to have a different distribution which varies with the intensity
of the colour. In those plants in which this is developed in greatest
intensity the entire leaf, ribs, veins and lamina, are all suffused
with a deep red colour. With a diminished intensity the colour
becomes restricted to the ribs and veins and, in limiting cases, to
the ribs only. These three stages have received the notation of
colour to lamina, to veins and to ribs respectively. Plate XIV
illustrates 3 examples—the parents of a cross between a pure red
form (Type 3) and a form in which the colour is absent (Type 9)
with the F, in which the colour extends to the ribs. In Table V
are collected the records of the determinations of the colour in the
young leaf from which it appears that the error is reduced to slightly
under 7 per cent.

A series of crosses have been made between Type 3, bearing
the red colour, and types in which this is absent and the behaviour
of this character may be most readily considered in conjunction
with that of the petal colour.

In the cross between Type 3 and Type 4 both parents bear the
yellow factor, and in this case a simple pair of allelomorphic charac-
ters is under consideration.

In the crosses between Type 3 and the two Types 7 & 9, the
yellow factor is present in Type 3 only, while in the cross between
Type 3 and Type 10 the yellow factor is again present in Type 3
only, but the case is complicated by the presence of the pale-yellow
factor of Type 10. .

Such crosses between a red sap coloured plant (Type 3) and
one in which this colour is absent have been more than once at-
tempted by other investigators. The earliest on record is that
made by Major Trever Clarke in 1867 (Watt 32, p. 336). Refer-
ences to these experiments are scattered throughout the Journal of
the Agri. Horticultural Society of India for that period as also in

130 STUDIE£S IN INDIAN COTYTONS

the Cotton Commissioner’s report for 1869 but no detailed report
has been traceable. A similar cross is referred to by Fletcher (12),
but here again full details are not given. In as much as, however,
both red and yellow flowered plants appeared in the F, generation,
it would appear that the red flowered parent was a heterozygous
form. More recently similar crosses have been effected by Main
(23). Here, again, the presence of yellow flowered forms in the F,
seems to indicate that the red flowered parent was not pure but it
must be admitted that the numerical results do not agree with
any Mendelian expectation. Further the appearance of plants
with white flowers in the F, generation would indicate that one of
the parents was heterozygous with regard to the yellow factor also.
That such heterozygous forms are of common occurrence in the
field is clear from our own experiments, and the latter case appears
to be similar to that observed and detailed by us (20 p. 52).

(1) Type 3 x Type 4.—This cross has been carried as far as
F, generation and the results are tabulated in Tables VII, VIII
& IX. It is evident that in this instance a single pair only of alle-
lomorphic character is concerned. The two characters comprising
this pair are presence and absence of the red colouring-matter—the
former condition showing partial dominance over the latter. In
Type 3, therefore, the red sap colour masks the yellow colour in
the petal, which colour is present in both parents. The F, gene-
ration is composed of two groups, one derived from protected
flowers of the F, parent, the second from unprotected parents.
The latter group, as Table VIII shows, contains numerous impure
plants, the result of natural cross-fertilization. In 1911 when the
F, generation of this cross was raised, the entire plants were pro-
tected under fine mosquito netting. A comparison of Table IX,
in which the results of this year are given, with section B of Table
VIII indicates that this method of protection, while it does not
entirely check cross-fertilization, reduces the amount very consi-
derably. The petal colours of the parents and of the first two
generations are illustrated in Plates XV and XVI.

(2) Type 3 x Type 9.—This cross also has now reached the
Ff’, generation, and the results are tabulated in Tables X to XIII.

LEAKE AND RAM PRASAD 131

It differs from the above in that Type 4, which has a yellow petal,
is replaced by Type 9, in which the petal is white. As far as the
present discussion is concerned, therefore, the two parents differ in
two characters. Type 3 possesses the two dominant factors re-
presenting presence of the red and of the yellow colour, which are
both absent in Type 9. Type 3, therefore, represents the condition
RRYY and Type 9 the condition rryy. Since it is possible to distin-
guish the impure form Rr from the pure form RR, it should be
possible to recognise in the F', generation 6 groups which should

possess the following numerical proportions :—

Flower. Foliage. Form.
(1) Corolla red ... Colour extending to lamina | & REY; ; 3 /
\9
(
(2) Corolla red ... Colour extending to veins & a vy , 6
(3) Corolla red on white ... Colour extending to lamina RRyy 1
3 3
(4) Corolla red on white... Colour extending to veins Rryy 2
(5) Corolla yellow ... Colourless {yy : 3 3
(6) Corolla white .»» Colourless rryy 1 1 1

That this expectation is fulfilled is clear from a consideration
of Table X, while the close agreement between the expectation
and actual results obtained in the subsequent generations, as indi-
cated in Tables XI & XIII, leaves no doubt as to the correctness of
this interpretation. Plates XVII and XIX illustrate the different
forms of petal arising in this cross.

(3) Type 7 x Type 3.—This cross, as far as the points at
present under consideration are concerned, is in all respects similar
to the above. In it Type 7 lacks both factors and therefore pos-
sesses the constitution rryy, while Type 3, as has already been shown,
has the constitution RRYY. ‘The results of the cross, which has
been carried as far as the F, generation, are tabulated in Table XIV
from which the agreement is apparent.

(4) Type 3x Type 10.—The doubt as to the exact constitution
of Type 10 as regards the petal colour, has already been referred
to above (page 126). In the present case the pale yellow condition

132 STUDIES IN INDIAN COTTONS

of the petal appears to correspond to the white condition as illus-
trated in the above two cases (2 & 3). This is illustrated by Table
XV. From this it would appear that the pale yellow factor is also
present in Type 3 which will be represented by the formula RRYYPP,
Type 10 having the constitution rryyPP. That this does not truly
represent the constitution of Type 3, appears to be indicated by a
cross between Type 3 and Type 9. Assuming Type 9 to have a
form rryypp we should expect this cross to produce plants with
pale yellow petals. This, however, does not occur. For the present
the true constitution of the pale yellow form Type 10 must remain
in doubt until the examination of the cross between this type and a
white flowered form has been completed.

D.—The Leaf Factor.

This character has been described in detail on a former occa-
sion (22). Opportunity for further investigation has been lacking
and the results there recorded need not be reconsidered in this
paper.

E.The Type of branching and the length of Vegetative period.

The inter-dependence between the type of branching and the
length of vegetative period is a matter of the utmost practical im-
portance. It is essential that a plant which is to be cultivated on
a field scale in the United Provinces should pass through the entire
stages and produce an abundance of fruit between the time of
sowing in May or, in the case of unirrigated lands, at the beginning
of the monsoon and the end of the year. Under these conditions,
if a remunerative yield per acre is to be obtained, the plant must
commence to ripen its fruit by the middle of October at latest. This
means that flowering should commence in the end of August, giving
a maximum vegetative period, 7.e., the period between the date of
sowing and the appearance of the first flowers—of 80 to 90 days—
a period which must be considerably reduced in the case of a crop
grown on barani lands. Table XVI shows that in the monopodial
types vegetative period is considerably longer than that of the
sympodial types, and too long to render the cultivation of such forms
on a field scale practicable, In crosses, therefore, which are pro-

LEAKE AND RAM PRASAD Loe

duced with the object of transferrmg the long staple of the mono-
podial Type 3 to a plant having the sympodial habit, a knowledge
of the exact conditions which determine the length of the vegetative
period is essential.

In the axil of the leaf of the cotton plant there occur’two buds,
one main bud to which the second is accessory. Vegetative growth
is effected by the development of a monopodium from either of
these two buds. Reproductive growth is effected by the develop-
ment of a sympodium from the former bud only. According as the
secondary branch derived from the main bud in the axil of the leaf
of the main stem develops into a sympodium or a monopodium, so
will the appearance of the first flowers be accelerated or retarded,—
in other words, the length of the vegetative period is controlled
by the form of the secondary branches. If these branches are all
monopodial, the appearance of the first flower will be delayed until
the appearance of tertiary, or even more remote, branches. IH,
on the other hand, the early secondary branches are sympodial
the first flowers will appear at a very early stage and the vegetative
period will be reduced to a minimum. The extreme types are well
illustrated in Plates III & VI. This inter-relation between the type of
branching and the length of the vegetative period was first noted by
Thompson (30) in 1841. A similar inter-rejation has been indicated
by Balls (1) in the case of Egyptian and American Upland forms.
The same point has more recently been recorded by Cook (6, 7, 8).
In the latter publication it is stated that all the branches that bear
flowers and fruit before again branching come from lateral or extra-
axillary (accessory) buds.* This statement is in direct contradic-
tion to our own observations, according to which the main bud may
give rise to monopodial or sympodial branches, while the accessory
bud when it develops, invariablyt gives rise to a monopodium.t

* See also McLachlan Bull, 249, Bureau of Plant Industry, U. 8S. Dept. of Agriculture.

+ In the Indian types, It is generally true of all types, but we have observed forms of G,
hirsutum in which the sympodial secondary branch produces lateral sympodia, As the sympo-
dium is itself built up by the growth of the main axillary bud, the terminal bud forming the
flower, this lateral branch, which arises apparently in the axil of the leaf, is developed from the
accessory bud.

~ See postscript, page 139.

134 STUDIES IN INDIAN COTTONS

In the pure types the difference between the two groups in
which the secondary branches are monopodial and sympodial res-
pectively is readily distinguished. In the monopodial types even
the ultimate secondary branches are monopodial. In the sympo-
cial types, however, although single plants are to be found in which
no monopodial secondary branches occur, in general a few of the
lowest of these branches are monopodial. Thus in Plate VI a
single short monopodial branch is developed, and, similarly, one
such branch only occurs in Plate LX. Plate VIII demonstrates the
maximum development of monopodial branching found in the
sympodial types but, even here, the type is quite distinct from the
monopodial type illustrated m Plate II]. When, however, the
progeny of crosses between types belonging to these two groups 1s
considered, every gradation between the two extreme forms is
found. In as much as, however, the change from monopodial
to sympodial secondary branching is abrupt, the main stem
is divisible into two portions, a lower bearing monopodia and
an upper bearing sympodia, and this character can be con-
veniently expressed as the percentage of the stem bearing
monopodial branches. Thus 100 will represent, in this ter-
minology, the full monopodial type and 0 the full sympodial
type which, however, includes the forms with a few basal monopodia
such as are found in pure types. Plate XVIII illustrates an inter-
mediate type represented by the symbol 70. It is clear that this
determination can only be made at the end of the growing period
and also that, to obtain such a symbol in any particular case, the
main axis of the plant must continue to grow. Unfortunately, in
practice, this occurs in comparatively few cases. Through various
natural conditions, and especially owing to the attacks of larvae of
Earias sp., which penetrate the leaf axil of the young cotton plant
and bore their way down the stem, the main axis is frequently
destroyed and growth carried on by lateral branches. In such
cases, this determination is not possible, and it is necessary to resort
to some other method for determining the character of the plant.

Such is to be found in direct measurement of the length of the vege-
tative period.

LEAKE AND RAM PRASAD 135

The vegetative period has been defined as the period between
the date of sowing and the date of appearance of the first flower and
may be measured as the number of days. This period is in itself
definite and also is determinable early in the season. It is, however,
subject to the influence of numerous subsidiary forces which make
the actual figure obtained for the plant merely the net result of the
action of these forces and not an exact measure of the true character.
Thus considerable seasonal variations have been found to occur.
These are indicated in Table XVI. A comparison between
the figures there given and the climatic conditions for the years
concerned indicates that a delay in the arrival of the monsoon
leads to a considerable shortening of the vegetative period, but
that the period is also influenced by the strength of the monsoon
is Indicated when the amount and frequency of the rainfall is con-
sidered. A strong and early monsoon with liberal rainfall leads
to vigorous vegetative growth and a delay in the flowering period ;
with a delayed monsoon the plants remain small and they flower
early.

The difference which is induced in the length of the vegetative
period by these causes may amount to as much as a month or even
more ; and the introduction of a seasonal factor becomes necessary
in any comparison between cultures of different seasons.

Minor disturbances in the length of the vegetative period may
be produced by such causes as the failure of the earliest flower buds
to develop into mature flowers. The bud withers and falls, and the
true vegetative period is in such cases less than that actually
recorded.

The inter-dependence between the type of branching and the
length of vegetative period may be indicated by a correlation
coefficient. Tabulating the figures obtained from a series of plants
derived from a cross between Type 3 and Type 4, of which the type
of branching has been determined, as above described, and of
which the length of the vegetative period has also been recorded
(Table XVII) the correlation coefficient is found to be *6628, while,
for a similar series, a correlation coefficient as much as °8589 has
been obtained. The inter-relation is, therefore, definite, and the two

136 STUDIFS IN INDIAN COTTONS

characters are merely two outward expressions of the same struc-
tural peculiarity of which the vegetative period, being most readily
determinable, has been adopted in the present work.

Balls (2) has adopted the same method of record. He further
has traced outa certain periodicity in the appearance of the subse-
quent flowers—a periodicity which has not been investigated by us.

When a cross is effected between two plants, one of which
belongs to a sympodial, and the other to a monopodial type, the
plants of the F, generation possess a vegetative period which is
intermediate between those of the two parental types. This inter:
mediate position, however, does not correspond to the mean of the
two parental values, but invariably approximates in a greater or
less degree to that of the sympodial parent.

In the F, generation the plants form a continuous series in
which every degree of length of vegetative period is obtained. It
is noticeable, however, that while those individuals of the I, series
which have the shortest vegetative period are in flower as soon as,
or even before, the plants of the parental type, in no case does the
vegetative period equal in length that of the monopodial parental
type. In other words, while the full sympodial type appears com-
paratively frequently the full monopodial type only rarely or never
does so. The divergence from the mean length of the parental
vegetative periods noticed in the F, generation is here even more
marked.

Table XIX illustrates the point for 5 separate crosses in
which Type 3 is taken as the monopodial parent with a prolonged
vegetative period. Owing to the seasonal variation which has been
referred to above, it 1s not possible to make a direct comparison
between the successive generations of such crosses. For this reason
the comparison must be effected indirectly by reference to the figures
obtaied in successive years for the parent types. In the present
case we appear to possess an example of partial dominance com-
bined with incomplete resolution of the component factors in subse-
quent generations, though it may be questioned whether such in-
complete resolution is a reality. We are dealing here with a case
in which the experimental error of the method of record is inde-

LEAKE AND RAM PRASAD 137

terminate, while, from numerous reasons, it is undoubtedly consi-
derable. It is possible, therefore, that it may be sufficient to act
as a masking factor.

The plants of the F, generation of such a cross may be said in
general terms to possess a vegetative period the mean of which will
approximate to that of the F, parent, from which they were derived.
In other words, an F, plant with a short vegetative period, will
produce offspring which possesses a short vegetative period and in
like manner those possessing a long vegetative period will give rise
to F, offspring with a prolonged period of vegetative growth. This
is shown in Table XVIII. In the majority of families given in that
table the average length of the vegetative period of the offspring
approximates to that of the parents. In certain cases this figure
for the offspring is considerably less than the corresponding figure
for the parent. This is easily understood when the various influ-

ences which affect the date of appearance of the first flower—e.y., fall
of the immature flower buds—are considered. In a few cases only
does the figure for the offspring show any considerable excess over
that of the parent. This difference is not so readily explained.
The difference does not become apparent until the F, generation
is well advanced, and as, in all probability, the influences which
produce this difference are such as affect the F, plant in its early
stages, it 1s impossible to ascertain them with any degree of
certainty.
CONCLUSION.

In the above we have attempted to give an account of that
part of our work which deals with the vegetative characters of the
Indian cottons. Though these do not directly concern that portion
of the crop which is commercially valuable, yet we have said suffi-
cient to show that they are of considerable indirect importance.
The habit of the plant is, as we have shown, dependent in great
measure on the method of branching—and on this habit depend
such vital points as the suitability of the plant for field culture, and
the yield of kapas per acre.

The study of the commercially valuable portion of the crop
is far more intricate and while a considerable amount of information

138

STUDIES IN INDIAN COTTONS

has already been gleaned, further study is necessary before it will

be possible to put forward a clear and succinet account of the

results obtained in this section of the work. We hope, however,

to be ina position to do this at no distant date in a second

Part.

LireRATURE.

Balls, W. L, Journ, of Agricultural Science, Vol, II, No. 2. ;

Balls, W. L. Year Book of Khedivial Agricultural Society, 1909.

Bateson, W. Mendel’s Principles of Heredity.

Burkill, I. H Journ. and Proc, Asiatic Society of Bengal (New
Series), Vol. III, No. 7, p. 517.

Burkill, I. H. Memoirs of the Department of Agriculture in India
(Botanical Series), Vol, I, No. 4.

Cook, O. F, U.S. Department of Agri. Bureau of Plant Industry,
Bulletin 147.

Cook, O. F. U.S. Department of Agri. Bureau of Plant Industry,
Bulletin 155,

Cook, O. F. U.S. Department of Agri. Bureau of Plant Industry,
Bulletin 222,

Darwin, ©. Effects of Cross and Self-fertilisation in the Vegetable
Kingdom.

Davenport, C. B. Statistical Methods,

de Vries, H. Intracellular Pangenesis.

Fletcher, F. Journ. of Agricultural Science, Vol, II, p. 281.

Fyson, P. F. Memoirs of the Department of Agriculture in India
(Botanical Series), Vol. II, No. 6.

Gammie, G. A, The Indian Cottons.

Gammie, G. A. Memoirs of the Department of Agriculture in India
(Botanical Series), Vol. II, No. 2.

Hartley, C. P. U. S, Department of Agriculture, Bureau of Plant
Industry, Bulletin No, 22,

Johannsen, W. Ueber Erblichkeit in Populationenund in reinen
Linien. Jena, 1903.

Leake, H. M, Journ, and Proc. Asiatic Society of Bengal (New
Series), Vol. IV, No. 1, p, 13.

Leake, H. M, Journ. and Proc, Asiatic Society of Bengal (New
Series), Vol. V, No. 1, p. 23.

Leake, H. M., and Ram Prasad. Memoirs, Department of Agri, in
India (Botanical Series), Vol. IV, No, 3.

Leake, H. M., and Ram Prasad, Memoirs, Department of Agri, in
India (Botanical Series), Vol. IV, No. 5.

LEAKE AND RAM PRASAD, 139

22. Leake, H.M. Journ. of Genetics, Vol, I, No. 3.

23. Main, T. F. Dharwar Agricultural Station Reports, 1909-10, 1910-11.

Z4. Middleton, T. H. The Agricultural Ledger, 1895, No. 8.

25. Moreland, W. H. Agricultural Journ. of India I (06), p, 37.

26. Parlatore, F. Le Specie dei Cotoni. Firenze, 1866.

27. Todaro, A. Osservy. sui Specie dei Cotoni Coltivati in Palermo, 1863.

28. Todaro, A. Relazione sulla Cultura dei Coteni, 1877-78.

29, Todaro, A. Prodromus Monographie Generis Gossypii.

30, Thomson, J. V. Proc. Agricultural and Horticultural Society of
India, 1841, Dee., p. 15.

31. Watt, Sir G. Dictionary of the Economic Products of India. Article
on Gossypium,

32. Watt, SirG. The Wild and Cultivated Cotton Plants of the World.

ROSTSCRILT.

Since the pesent paper went to press a further note by the
same author (O. F. Cook, Circular No, 109, U.S. Dept. of Agr.
Bureau of Plant Industry) has appeared and is mainly devoted to
a criticism vf our interpretation of the system of branching as
here propounded. As this circular has reached me in England,
where access to our detailed records is not possible, 1 am unable
to enter in any detail into the points involved. I may, however,
take this opportunity to deal briefly with the general questions
raised,

In this circular (p. 11) it is stated “some stalks are right-
handed and others left-handed with respect to the position of the
extra-axillary buds and the branches produced by these buds’—
in other words, the extra-axillary, or accessory bud, will for a
given branch, lie either to the right or to the left of the main
axillary bud. This is in full agreement with our observations
(Jour. and Proc. Asiatic Soc. of Bengal, New Series, Vol. V, 1,
1909, p. 23). The author proceeds to state that “the axillary

buds may be developed into vegetative branches...... The fruiting
branches arise from extra-axillary buds...... ” The conditional

tense here used implies that, if buds develop, they will give rise
to vegetative branches, but that development need not necessarily

take place.

140 STUDIES IN INDIAN COTTONS

Assuming the correctness of these two statements, let us
see what must follow in—to take a particular case—a ‘sympo-
dial’ plant of the Asiatic type of cotton. In sucha plant, as I
have shewn elsewhere (Jour. of Genetics I, p. 232), the passage
from the lower vegetative, 1o the upper, fruiting, branches is
abrupt and, further, when grown under normal conditions, a
single branch only develops, the second bud at each leaf axil
remains dormant. As the vegetative branch is developed from
the axillary, and the fruiting branch from the extra-axillary bud,
and as the extra-axillary bud lies constantly either to the right,
or to the left of the main bud, it follows that, when the dormant
bud on the lower, vegetative, portion of the stem lies to the
right, the dormant bud on the upper, fruiting, portion of the
stem must lie to the left, and that, conversely, when the dormant
bud on the vegetative portion of the stem les to the left, that
on the upper, fruiting, portion of the stem must lie to the right.
Observation, however, shews that, in the type of cotton we are
considering, the position of the dormant bud is constant through-
out the shoot,—a condition I am inclined to believe obtains in the
American series of cottons also, though our observations of these
are less extensive. It is impossible, therefore, that the two pro-
positions I have quoted can both hold good.

One further point only can be referred to by me here. On
p. 12, a reference is made to the influence of environment on the
character of the branches; under conditions which dwarf the
plants, few, or no, vegetative branches are produced while, under
conditions favouring luxuriant growth, the fruiting branches may
be replaced or transformed into vegetative branches.

In the sympodial types, as defined by us, the extra-axillary,
and even the lower axillary, buds remain dormant if the condi-
tions of cultivation are such as to check growth. Under such
circumstances no vegetative branches are produced. When, how-
ever, the same types are grown under conditions favouring luxu-
riant growth both buds develop, giving, in the lower portion of
the stem, two vegetative shoots—a condition incompatible with
the second statement quoted—and, in the upper portion, one

LEAKE AND RAM PRASAD. 141

reproductive and one vegetative shoot; further, the vigorous
srowth of the vegetative branches leads, owing to the exclusion
of light and air, to the dwarfing, and even to the complete death,
of the weaker reproductive branches. Here the vegetative, re-
places the reproductive, branch, but does not arise by any process
of transformation, for the dormant bud, frequently represented
by a weak or dead shoot, lies on the wrong side of the vigorous
vegetative branch. <A single case only of transformation has
been observed by us. It occurs among the Persian series of
‘“herbaceum” cottons, and is referred to in our paper on these
types (Memoirs, Dept. of Agr. in India, Bot. Series IV, No. 5).
It is in this series of cottons that the widest range of diversity in
form of branching is developed, and, in consequence, the series
affords a valuable field for observation of the methods of
branching.

That environment is able to produce a change of habit is
seen to be true for plants of the ‘sympodial’ type. When, how-
ever, we turn to the true ‘ monopodial’ types, no such change can
be induced. Under adverse conditions, no, or a single, vegeta-
tive branch is developed in each leaf axil of the main stem ;
while, under conditions of vigorous growth, two vegetative branch-
es are produced.

For the reasons already given I am unable to follow the
author in detail into the minute structure of the branches. I[
have shown that the two primary assumptions are not supported
by observation and that the change of habit induced by varying
conditions of environment are due to the relative development
of the two types of branches and not to any transformation of the
one into the other. These facts point to the fundamental nature
of the difference of the two types of branches. The reproduc-
tive branches are admittedly sympodial in structure and it seems
more natural to infer that they are truly such than to hypothecate
a series of torsions in the earlier stages of development.

Hao, Lb.

142 STUDIES IN INDIAN COTTONS

Ta Bw Al

Type 10 (pale yellow) x Type 8 (full yellow).

“FP 11 plants all full yellow.

fe 99 plants full yellow. 41 plants pale yellow.
Ratio 2°4 1
Expectation 3 1

Tapper it:

Flower Colour. Type 4 (yellow coloured) x Type 6 (white flowered).

Fi 68 plants all yellow flowered.
Po { 109 plants yellow flowered. 52 plants white flowered.
P ratio 72 1
Fz plants used as parents 21 13
5 10? 6 13
pr, fyellow 65 35 34 0
- \ white 0 0 11 100

1 No difference has been observed between the direct cross and its reciprocal. The two have,

therefore, been grouped together in this and subsequent tables,
2 Number of offspring too small to be a reliable guide to purity of parent.

PABcEe Lil:

i
Parents, Fi. Fo. Fs.
z-| Lal
g|8 (88
oo ¢ >=)
Type 7 x Type3 15 plants red Pure yellows Gilecculy Gee All yellows ... 271} ...
| on yellow. Impure yellows]... 16} .. . | Yellows LF 680))'2:9)\ aes
_ Whites sve 1290) plea
Unchecked yellows 618 |
| Total yellows’... 630 | 3'2| 3
| White .. 196} 1, 1] Allwhite ... 121
Type 9 x Type 3 67 plants red Pure yellows gee hE | cae | eee, | ALL Yellows! co. 2 OGOUe-sulmare
and reciprocal. } on yellow. Impure yellows ... 169; ... ... | Yellows og 226 14/)/2:3))) ns
Whites ... 808 Led
Unchecked yellows 135
Total yellows... 318/2°9| 3
Whites «» Jdd} 1] 1 | All whites ... 1366
Type 10 x Type 3 | 50 plants red | Pure yellows sory OR | cee | ce ALD VOLlOW. “ade COUN osrey ae
and reciprocal. | on yellow. Impure yellows .. 15) ... . | Yellows Saar ZOD! | eon |) cee
Pale yellows... 78| 1) 1
Unchecked yellows 310
—|
Total yellows .. o2oial1| 8
Pale yellows .. 103) 1] 1 All pale yellows 35

LEAKE AND RAM PRASAD 142a

ABU Mey

PARENTS. OFFSPRING.
Number. Character. Eyed. Eyeless.
|
= | Be
14 Eyed | 454
1 Eyed a) (= 40
| 21 |
|
1 Eyed + 6 Kye -
| lightly | |
shaded |
2 Eyeless | “ee 34
4 Eyeless resis | 181

iABEE.. V:

The intensity of the red colouring matter in the petal as an indication of purity.

Flower of #2 parent Constitution, as determined Total
recorded as by F3 offspring, of the form a

——S SS

RR Rr

(a) Type 3xType 4
Red

a 28 2 :
Red on yellow ca 35 136 171
Total B 63 ~ 138 201
Ratio = 1 22 See
(b) Type 3x Type 9
Red a3 1l 3 14
Red on yellow =u 46 136 182
Total as 57 139 196
Ratio & 1 i2°4 ae

Taste VI.

The intensity of the red colouring matter in the leaf as an indication of purity.

Leaf of #2 parent Constitution, as determined Total
recorded as by /’3 offspring, of the form bate
——_———
RR Rr
(a) Type 3x Type 4
Lamina ret 61 5 66
Veins a 2 20 22
Ribs se 0 116 116
Total ees 63 141 204
Ratio Me 1 2°2 A
(b) Type 3xType 9 g
Lamina si 59 4 63
Veins 13 2 15
Ribs 9 188 197
Total Laan 191 275

Ratio Sot 1 24

143 STUDIES IN JNDIAN COTTONS

Taste VII.
Sap Colour. Type 3 (red flowered) x Type 4 (yellow flowered).

F! 38 plants with flowers red on yellow and the red colouring matter extending to veins.
RR
RR Rr RR+Rr ir
F; Foliage (lamina) (Ribs or Veins) (Total (Colourless)
77 147 Coloured)
Ratio 163] el 224 69
Used as hs Lamina 5 Lamina ae 1
parents J 2 Veins 136 Veins ne +7
204 68
Reference to F ‘ =
Table VIII. . 2 2 é 5
RR RR Rr RR+ Rr rr rv
F; Foliage (Lamina) (Lamina) (Ribs or (Total (Colourless) (Colourless)
Veins) coloured)
1328 &32 1692 2524 773 1245!
RACLO es uss e sss 1°07 2°18 ove 100 sé

Seen eee
1 And4red plants. A consideration of other characters indicates that 2 of these are without
doubt either volunteer plants or have arisen through an accidental mixing of seed,

Tapre VV LL:

F, of Type 3 x Type 4.

PARENTS. | OFFSPRING.
| Column Serial
[aNinmbers|Moftntite No. Lamina Veins | Colourless
| VII A. B. |
: |
(a) Selfed Series F 13 1 271 |
8 2 2 194 a Se0
12 3 167 St a 39
Expectation | Sl oe eel 170 340 | 170
13 4 4 | * See 155
6 5 5 120
(b) Natural Series 7’; 5 1 6 108 (1)
27 2 7 625 (83) Bs
37 3 8 203 | 393 | 176
22 4 9 om (50) 493
| 5 5 10 a (3) | 132

AB ee.

F. of Type 3 x Type 4.

PARENTS. OFFSPRING,
Composition, Rararante
No to Lamina. Veins. Colourless,
Table VITT.
|

RR 11 LA 267 (2 ao
RR 3 3A 182 oie ah
Rr 5 3B 76 158 86
rr 6 5 C aay 595
RR 12 7A 4 af ey
RR 10 8S A 131 (4) ae
Rr 4 8 B 16 27 17
rr 9 8 C a (1) 143
rr 7 9C | 175

143a@

LEAKE AND RAM PRASAD

*dnoas oY} JO UOTZVOYUApr O[qeITA.A AOF [[VUS 003 Suradsyo Jo aoquinyy +

a
a Dales | T T rt T $ | Z pe Ney a T Zz I I g | dnois yore
| | | | | Ut uol1yB Ded xX
See abel toca tcc a Soul atel 1 LSPeG:S Ost |e es oeateeeme cere TH 96 Perce
% 1 1GG | 1ol | | [0G lf cl z ,
e1¢ | Sta| 89 | 6c GSI | chr | F0G| SIZ | 06 | eee | 1s% |96 | 9¢9 2 | bat | ves | LIL | SFI | 86 | 102 IIX ¥ IX
0G 61 | SL LI | 9L cI lal tl Gl Ul OL 6 | 8 L | 9 |G Pb to G i! S9[QE,L 10j VOUDIOJOY
| | | | |
| ea aite es a0 ae oa es eS | ; | eer ;
re — fas pana > ete | eetiesva lees |e ess = > > = ie = =
| 4) 22 SUS es (aya | S eee | ae | Sa | a e2| m | 2P| & | a
(eG EB) 1 oO) & |e |e | go pan)| & > |sa|/s2|)/ oe] @ SNS pas] Deo eects ee Es grec
s| 212) & |®e |Se ee eeett | gh Nig, |S | cen ins || eee ees cl ey en
— = es en matte fi oe I | a S| aS [=] Ss | a a) . oa)
| | ies | sl .
|p 5 — poe = 7 | a arp ie me = a = Aa See es lS = |
On lo- Sq |S ‘ AS eels Bes [See Sel Ge Cs | Bey [Stel oe |
(userp)|f 2/25 2)22 Sx (ae |e & | (eurmwy) Oz PEE a2 | SE SPREE OF | SP) ER ESS aR] Bx | asvrloy
kaa |82/B Sts Se [Fu lec | MMH | gee [ett See | Be Sita ee] S| Set | ai Eee [Be | Be | a
Sea Soe ea ae ae eens ae PSP eS amy 24
6G & y ae Fe 6 el 83 leer sjuosed sv pass)
a = Ll LI LE
T | $ S I 9 e | uotyej00d xm
8-0 0-€ BG 0-1 0-9 8G i ts |
&% 96 SL 0g £61 68 |
| ; | s(oqryas : £ 19MO0
( ue *(MOT[9X) (971YM UO poy) | uo pay) (‘Ao[]eA UO pay) (poy) A la
pee net: Z Neate ee
*(u9a.14)) *(ugaa19) *(SUTOA ) | *(euruery) *(SUIAA ) *(vurmMerq) asellod
AK at | &X at‘ AR Ul AK ay | AL a AX AW ‘AA AY AK UU ‘AA UD oH
*MOT[OA UO pos STeJed YIIM JOAOG OY} PUT SUIOA SB Avy SU 109}vUI SULINO[OD pos YQIM syuR[d 79
61 ¢ eddy, x 6 Od4 I,
SP 6 edky, x ¢ edhL ‘al

OST,

*(pazamoy oy14yM) 6 adAy X (paraMmoy poi) € adAy Inojo) JaMOTY

144

STUDIES IN

INDIAN

(PA BLEs

F, Type 3 x Type 9 (Selfed Series 1910).

F, PaReENtTs,

Character.

Number.

Red Lamina
Do.
Do.
Do.

Do.
Do.

Red Veins
Do.
Do.
Do.
Red on white Lamina
Do. do.

Do. ;
Do do.

Red on white Veins ...

do.

Do.

Yellow
Do.

Serial : re
: | Composition,

I RR YY
II RR YY
Itl RR YY ss
LV RR Yy ee
Expectation
V RR YY .
VI RR Yy a:
Expectation
Vil nai NENG Bs
Expectation
VILL | Rr vy wah
Expectation
IX Rr Yy Se
Expectation
xX Rr YY* or Rr
Yy.
XI RR yy
xi RR yy
X!It | RR yy
XIV | RRyy
XV Lay eayay ie
Expectation
XVI | Rr yy ae
Expectation
XV} rr VY he
XVILI! rr YY
SKS alii
XX rr Yy ve
Expectation
ok i ee :
XXII | rr Yy me
Expectation
XXIII} rv yy
XXIV | rr yy
XXV irryy
XXVI\ rr yy

COTTONS

- F, OrrspRING,

oOo a + oa = ee | = aos

g Red on Red on

co) ? 2 ;

5 yellow Hike Yellow, | White.
Oe ee .
= S| aS : = ps 3
=|) 2/2/28] & 5
S| =i a |'s & 3 £ 2
“i ge 4 > 2 > id) o
3 AS aT PF) 7 .
7, S$ A B | (6 D E I
13 1 86 a i
4 4 38 e Loe
2 2 10 a
2391 NG Sul: b
ele 6 Sui ke
Sls 7 198 | ae ;
LOR 7 (Se PAI ae
ae Ge OGal.. S25 ae
Sih 25 18 | 2% =: & 7
a ae 18 | 26 exc 138
2 8 4/14 ee 4
164|"8S 52 | 98 16 | 34 44 16
Ret liees 48 | 96 16/382! 48 16
9) 8 9] 17 Ne ||, ace 3 bee
30 | 13 11Gs) & 5
Sal axe SOF) te. re
16 | 14 LOSS e (2 aoe
9 9 148 - ‘x as
MW | 15 | P20 ie eee 10
E 3 | 9/18 se 9
16 } 10 26) az ie 36
on ae 383 66 = 33
14 | 17 on on 67 ie
ee 88 (1)
15 |} 11 158 ist
10} 11 40 m1
be ? 45 15
8] 18 | 47 Bae
17 | 18 54+ 19
ms oA 54 18
3} 20 Be 10
8 | 16 19
8 | 19 37
10 } 12 50

* Number of offspring too small for reliable identification of group.

LEAKE AND RAM PRASAD 144a

Taste XII.

F, Type 3 X Type 9 (Natural Series 1910).

(0)

F’, PARENTS. ee F, OFFSPRING,
— - ST eal lit | a - —— = :
$s Red on Red on |, | :
f 3 | 2 yellow Hite || Yellow, | White.
° Se | /© |
2 Se sil ics | E i i
Character. 5 | Composition. © | be = f 3 = 5 w 5 e
a | e@igo| = |/2| <¢ |S} ss oe
= | 2 |= — i < a
2 | 3 52 desahit Aaes - aed VS:
Bz | ae pea ee Cy Ba age lien
Red Lamina 5 I RR YY 6.| 1 | 135 3
Do. = II RR YY 5 | 4 2 8
Do. See YUAN a ee Le ee2 |e 2
Do. Fer IV |RRYy yee 95) Te S18 3
Do. arte Vv RR YY Gol epee 10
Do. HPN In ah Re vy (oa | 40 | 2) 46
Expectation eee |e ae ke .
Red Veins ool! SAVIN iid eae EN" SH OB See ae Lege al eae ve tice 81
Expectation we | 765 [153 y "65 :
Do. sce NOU E lo e'nG e} Tt) S|) 64 2 (143 x =o 56 a)
Expectation... .. | 66 |182; .. be, 66 re
Do. use 1X Rr Yy 2153) 8) 55° (133, 2 | 27 79 D4
| Expectation |... ..| 68 (126) 21 |42 63 21
Red on white Lamina el RR yy aed lstayenul 5. | 149 9 si
Do. do. 5a.) TIE WR AY 3 Ie ate 3
Do. do. ... | XIE | RR yy. | 14 2 | 105236 3
Do. do. sf) XIV | RRayy ie Be 99 2). 80 1
Red on white Veins .... XV_ |Rryy | aSe| Ws WS | 2893) 214 16 125
Expectation | .. lier -. | 122 |244 122
Do. do. Seen NOV) Ties ER VAY : 3 | 10 2 Ga ite Ie 24 1 15
Expectation st realise Tetley ee br: 21
Yellow soe |} eS ATIE | pe yeaah ets OF Na Ue a 2 . 124
Do. $5 | 2 SAYIN TENANT Ee: per dale MOL |Povis =e 1 225 (1)
Do. foo |) OKTENOM TR NOY ssi ores plik eee. UGGS), 272
Do. co eee heyy eect) dle lire. we (ho) 69 18
Expectation as¥ | “+ 66 22
Do. By) |e: D.4 al | Th ans E | 40} 18) 2 90) a
Do. ao |e. TNL |e ay BATA T Srl. wee teeee tet : 415 122
; Expectation 5 dom lant 4, 4 ve 432 144
Whites . | XXIII] rr yy £ ie ie Ua) eee aa Ngee ; . 39
Do. | XXIV] rr yy ee PES TelG le 6. | Sages Me ee 21 R45
Do. ai 7 XXV | rryy ¥. Gre lOh) se. %: bd 6 211
Do. "| XXVIj rr yy Bi al NS Fe a = ' 29

eee Ee

145 STUDIES IN INDIAN COTTONS

Taste XIII.

F. Type 3 x Type 9.

PARENTS J’, | OVFSPRING,
| Red.on),.|~ Rediot | -velicw:|-iiuttel
Reference inaz=he Sl
Character, Composition. (cee gee) = ’ ; a a
fete OX NE sm: |, 5) slower Soft e | eee ee teens | el ge An
B/S |} 2 | ee 18 | 22 eee
5 = (3) e cs) of of
A =) > | > oO o
be be | ,
Red Lamina Pl) eo Way Bee as A DO ires “a
Do, es PRR MONS As Ms AC) 6 ie 5 F
Do. Aas SISSON ae DL) Ver A0| o1 2 ~
Do. 2 | eee Was SA AE ONS 26 | oe
Do. | RR YN Seo IL ATE BEA Se 52 me :
Do seul Eee Way: 1A 2A] 2 2D) It ase Whe esc
Do. See [PR NOG ve {12 FX Asl 2 7 ae ne fe
Do. seal ee Nm | TOES WAN ICS2 | “AlOb cee: BO ier:
Red Veins STP a ees XS Base? ORNTAS) | celles 80
Do. ee VAY IS SES BTSs 45 | 87 13 ) 31 35 15
Do. Fa ERA AS Beil? TEX SBiine 9 | 26 Sau 1 1
Red on white Lamina RR yy eee PE! ee @a) “2 2))(1) | 191 | -.. ;
Do. do. .. | RR yy OL RECN so MA
Do. do. .. | RR yy Jo EE XLV iC) 4 206
Do. do. ..| RR yy Jos he ex Ve C 1 58
Do. do. ..; RR yy . (12 XVEO! 4 On| ee ee oh
Red on white Veins .. | Rr yy Set) ODEN: ITDAT eres 63 |152 of 68
Yellows NT oe ahs we el 1) GN Yt Reel ea Re. _ i|leee- 7 aa
O. Se NOG ro LL LPB) pail 38
Do jie Nas LIZA Le Dp 6
Do rr YY 1LXVITIE! 6 259
Do By aN |11 XXI E/ 1 12
Do nr vy |12 XVII E| 4 30
Do Troy wi S02) SX | 2 | a >:
Do rr YY (12 XXE H | 12 : 52 (3)
Do rv Yy pL x aE) 3 432 126
Do rr Yy Ve; ae Bh 2 12 ¢
Do, rer Yy }1) XXTIITE| 2 25 9
Do rr Yy I2eexex 1D) 2 1 4
Do rr Yy 12XXII | 1 8 2
White ... | tr yy SV TOE he. a ee aa ea nese ee ace: s 14
Do. ely Ay ae |e ey | <4 omogelieeds aol EEE bt | 42
Do. rr yy Fol) VEU. @ Cle (97) Fh) I Reena Neco etn Loe | ay | 39
| PES Re

SE

145a

LEAKE AND RAM PRASAD

I 8 T ae | Hie g Bat g I 8 “uoreqoodxam
I | LE l Ste ale eas l | &G “G GP GZ I 2 dnoaz
| | ove UT 019%]
9G cé | 601 #8 €G ¢9 It lee | OL 91 69 | LP Sli GLI £6P CG
ee ies | ee [oS Pe = | ee = 2 == we
} } | 6 . lite j ‘3 |
"OUI AA “OFT AA |ANOTIOK [*MOTIOR | “OFT AL | io nae anes | OUY MA | MOTLIA as pew MOTI A ‘poy cocpes ‘poy oo: IOMOLT
| | |
—_ 2 ee ee ee — => || ees SSeS
Uda) “UIIAH | ‘Ud9Ix) | ‘UIdIH | 'U9d.1 eee ‘euruer| “U9e19 | ‘U9eIH % eaaiey crys) aaa ‘RULE VUTMeT | -euIMe] “-eseIoq sy
| t . | L
Ea =: a = Is
¢ G to 9 I | ¢ | rs 6 g oo squespd
| | | |s@ pesgQ
eamee ze | ease $a) Aenea : , 2
T $ v4 | I 9 $ uoTzBj00dxH
OT Gg 9.6 LI 0.9 0-8 [++ dnoad
oF SFI 401 | OP } §96 6cL | (Ou Ur OFVY
ees ——
*(99TU AA) | *(MOT[OX ) *(9JLYM WO poy) Peal *(MOT[9A UO poy) *( poy )
- s =e LA aes == r. ==
BtCRTS) | ‘ueadIy “SUTOA eUluIery: "SUIOA eure] a

*MOT[AL WO pat spezod AOMOP OY} PUT SUIPA SU IE] SB 1049CUL SUTINO][OO pot yy ‘syueld cy ‘Ww

‘Eadky x L adhy

eNOS adv 7,

COTTONS

INDIAN

STUDIES 1N

146

‘g[Qe} OAOGE OY} UL PEpN[OUL JOU a1¥ 4Nq patandd0 SES8O.10 SNOTAGO MOF B “poes [eANgwU MOI A[JAV

uol7e10edx

8
I hE ie Pitas eo I eee Lol eal eae a dnoid youve ur onvy
cg cr | a fe. | 69 | 9¢ | gr | gc |. ec | st } 98 | a 9¢
ee) ee Mes ee a : = |e ea ce
{
<“ ia + «< “ | at role te Oo =< “_ +
-m@qes | So} & | o | 25 Gc ei | NaleC al ices cF se e| & ==) © mx |Sce| Bw Ee)
Helse | see | Sh lo>hienre oy alee ed al ae eee iced oe OMOT
a he om | > | irae <s . wae ct = a .
Ss Aa Sei Pad sig hence hae ae se
ene || me lee ee Pe lie eerie ol
ueip| ¢ | |B |e | & | 8 Bo | Be Be ee eg eo a a ee aserOg ou
ee a a Bo) BS Be leit Sa ae ee ee enn
I € 0 ¢ | 0 cl 0 0 I oe sjuaied sv uayey,
}
"u9aA) "uaa. *SUIO | 5 “SUID A = ODVIO MT
s s
© =
9.() € e€ 66 ;
LT 6L 98 Lt o1jey
“MOT[OA : ; . , ; -
alta MOT[OA MO|[OA B[ed UO pax MOT[PA UO pat pu pay | IOMOTT Sy
“MO[OA UO pat speqgod YAIM AaMOY PUL SUTOA SB Avz Sv 19}]VOI SUTANO[OD pat YIM SPURL
oe
b ie g addy xor eddy,
OF : reddy xg eddy, 'y

‘or adAy x € adky

AX Wav L

LEAKE

Taste XVI.

AND RAM PRASAD

Seasonal variation in date of flowering.

Types,

Monopodial Type

Sympodial

COeNMoewOMm

1907 and 1908. Sown in pots and transplanted.

1909 to 1911.

Taste XVII.

146

83
8U
gt)
78
93
96

1907,

1908.

207
210
110
110
114

96
106
117
115

Sown in field.

1909.

|

1910.

Not Sown

145

Not Sown

1460

1911.

94 97
90 69
0S 74
82 8
9+ 61
96 75
11 S4
74 52
Pe 56
° a7
ay i

The interrelation between the length of the vegetative period and the type of branching.
Type 3 x Type4. Field Series.

Number
of days.

Below 65
66

Average
period,

OROWWOOINAIWH

YPM fea | Fr aS as

ad
NwWwW DH Iw NS

eee Slt the Este

S4

ae
i=

\
moe ee | |

40) 50 60 70 50 90
aC 1 =: e = =
2 = == : == a
2 at = zs = Be
7 1 = = —
12 1 1 = —
16 7 6 —
16 9 6 2 — —
13 2 1 — — =
15 4 6 5 2 —
26 13 11 4 2 1
22 12 11 5 2 1
16 13 7 13 1 -
15 10 10 6 1 —
8 4 7 t = =
12 13 8 3 2 1
3 18 18 10 5 —
Bib 23 31 16 4 2
28 a2 20 20 3 2
34 25 25 9 1 3
5) 8 10 4 1 2
ul 9 11 5 1 —
12 10 19 12 3 ~—
12 9 18 9 3 I
8 13 10 8 1 1
4 5 12 7 2 —
4 1 3 1 — —
== 2 = 1 med =
1 2 6 5) 1 —
2 4 2 3 =
— 2 = 4 1 —
1 1 4 2 1 se
1 — — 2 — —
— — _ 1
= = 1 = =
_ 1 l — _ 1
1 — — 1

o
aD

104

branching,
— a
2-day 5-day

period, period.

15 20

25 1s

16 ————

19

19 19

20

ae

21 =

25

27 26

32

33 36

39

41)

38 39

41

43 44

47

47

46 49

30) ————

52

56

HS)

56 By)

SY ==

60 53

55

61

61 61

By)

45 59

66 66

TABLE

14

Type 3°;

= | Wiles Sa att at gray puts «apie seie De IM rN reer tea rs te m=O se NO tet cetesonee CO om: emt cml ce
aa | cette ne) tM a, ae vas Coe . . . - 4
a sense. Ain lar are SOU te ot een ian! Ole Ol pe Cle) ose + Moe TNS Oe ee ee
gs a ee OO . : ions
a ae 2 ———— — ——-
= 4 A QO’ 1s : rm: ND re Nim srt: Hise bce Set ts Nem MNRKo cs cain
x2 . mm cr i Eo eeed ie Een dk eS ok eS od) NNN Ole > oie FIN RK CIA eS
so ‘ Sete set sett THD OUI HOT 2 OMe ete 5 re IAN CSR NOM RK GOANAMN Aas AaAsi
3 Pt ONO eT ced reo ee Fs SOON eet fot st re OM ede RK MS cme Hor tee
. < = ~ = —
gy wet em OU Ns ets cet mei tt 8k tee et ot ON ee fa | — OD OD wt we et et SH ee nN
= “ : : : B pats 5 nee 2) halle cc cee s
Se ee SS a a a a _
=
=} FO a OCI Te is sie ME on oe oe Oo Or ee Tacit en Ouest NO pet 108 aie ore
= ee : : a ; SiS os variant agen oe ler itn cme :
a ee cs
2 eo Ol i eo. Sk tk ee mei icy a Ty) es
S eae ae chy tnl cee ee or R ; a on pt SORIA Re Ser rang) ni Me lie Ue ae
SS So a ee es. —— = : —
= es OC Scie cme cet msi SCA 5 SiC 8G a gs Siered sedi 5) pedi) 0) TN) eet oP wed 08 34 6) eek oe ee
—= sy mya: ery ° Cj av btw: Sayre, oat at hee ole “ ai As c : ee ree ine
| —* -. = : - ———————————
Ss eel Nae Koos Hao RR fc ES (ser iia lie ear — Pern aint eth meena feng ees Bi | OE ep Ge ey
LL a a SS SS Sa a ———
a
Bu ae. asain: CMMI Ln Kn! Mm Dae cool) Deere Wty avis crinemCne mis ire ei Pea aie vm fe oy Se Pit vay nh eyo ae
S Pee a GST at deo Gain Taare ent Bet Qe ret cS eee, te ee i i
= 5 : hes ; Ae et aoe : cea Sani Rae aia anaes a oe k,l ry Oe i :
———— eee
LD Co has letra mee tie Ot Beem mae spay eats — halicha BCrAE at eb I a oe | cist SS abies ~ 1
ror) ha ; ee HEC a tee qceart conae : Sn oe eee ae :
— Ret ESE ee Meme cheer ay Gee sae a aoa tee =e Gnd Ct ee eee sr ek Shc cae sa) Ri ela vans eg ieee he =“ qeet Soi eee 3
or) i i a a ce Sacer oe i ar nea anal ie merit irre en Sse oy yr Sy RS ey
ex Car Coed Wry) be ae ea ml oes SPP er elt! Sr Ors et fo. UT enyesk SSM ois asta One kal tares ce Loy
pa Tie rae Th Sei a al a CRO greet iced mn inate is mire aye Smnty leuien Mee A IC) A SH I AHS tha, =
> Se telnet 2) tat a de Ce eee NC ee Hee ok Walt or aera ay TNS Coa on Ps st BF
x Sie Avart oat eh RS crus rt tc ee 3 nt MG dees Mee te tin cm IS sy ee NS ee metres Pi brea icy Etats Lome age pacer 4
——— era eee a ee ere ee ee a ee ee.
g |e ice Pata ll ae me a re ra eee Re ORD ORI a trey US
Do tekst eaten Void se Re rise toy (ai pho ee fe Siete “Poamhg mtgt CUMS mney ys iesns UD? soils) ESI NG Deri oy ie Si ot NR 2 ca Se te SS Ssh Be RS ee eee ee
> . . . . . . . . . . . sO ac “> a) ie ie eel ae . eee en es eee Se Te ek oY ee en Oe es mk Ce ee Pe
en : RE Ri ah rai ees ANS CROP EC tet MS Gee + oe Vilien een Se et AS See ee cee
F ee ae RCO ae Oe ee ees ee ee ee Mt et Area RAC So Mii TN met POS ea ne
‘OS AOloq Ste eae ol ate Tee ricer ie Sagi ner eter or dc) ¢ Pao b Por TS CPR gies See ere sec
SSSR SSS eSB VSS SS IVBVSSOSRMSIASoO SS TE FEM SMIOSCHRAANNSPSAM SE SOO +H
ADOHNOWMMNMOHDIORHANSSOHOWOINS P-BSSSnrknsssSsszesercs NOSSHOANDS = FHOS— HMI
MWiHRMOTOO-AHA-OSKGSCOKSOSCHOCOCRSEOCOCSORLE SH HOG COMP KKCMOMR SHAK Se t- Ot SSSR Se eis

| |
126 128] 130.

_

——
—_

ae

132 | 134/136 135

!
|
|
|

140 142 | 144

|

146 148 | 150

|
|
152 154 | 156} 158 160

|
|
|
|

Above 160.

_
_-o

=

tc
ow

toby wom te Oe

l1—Do-<

CO 4 00 O11 Or ¢

—- 6
ay
- 3
aN.
+ 6

7 a |

1+

—

eee henctte9 perme

ae :
Peo Rpwy

Ke
a A

Wwioou=

—-
—

Lh) rr |

| Offspring.

148 "TABLE

|

Below S80.

=
ee:
a a eet

| | !
|
| a is lalate
80 82 81 86 | $8] 90] 92 94 | 96 | 98 100 | 102] 104/166] 108/110 112/114] 116] 118 129 122 124

2 Ell on en
ibe eer

lo

_

Mromomae: s :
«

pt bt poet pes S

oy fel fee Se
.]
—wepRts:

— os!

Lom woul es ume!
» Clee t

=
tom!
a en noe

— i
Gots
“Ibo Le

m—-1t5

or

_—

nd)
whom
Fmto

2
435 | Ree ae ae a ea Ped 1 wee 74 ¥ a3 l a cane 1 rer
2 EERO MBS 8 De © cca TMS Vi a VP pC eM SP We, Ce
701 | 1 1 1 1 2
)
| 5

Lom bolo bu

7
<<
ee oe)

Re Kolo we loos

=
&
el
—
to_—
_
—

_
> 1
=

te ed

=
roy
-S
—

—
-_
—
Os mm we
—_

XV II1—contd.

| |
| Eas
126 |128 [130 |132 124 136 138 140 142/144 | 146 | 148] 150] 152 | 154/156 158) 160 *

plants.

I
a

=

= oy

oo

———$<$<$<<$<$<$——$—————

IW we

SS

Above 160,
Number
Parents.
Offspring.

|

l
-—
wWWwWoOnDooe
tt et et et
et et et et et
PMRSDHOLOLIAEAMD

» 118
118

lor lol
t
_
~
s)

—
to
eNO
—_
~
=
rey

—
— LS
_—
=
et
— et ee
OOo ce

a

_

w
el

-_
or
2)
nn
|
we)
wo
a)
eet et ee
Woon

wom
to

—
—_

+ | f
WCWlOMWMNODOSNAHMNwWwo ws

—

-_

=)

wet

2
ONnwwouwnIss

mth

1)
ar
a
~
S

mtom lo
woe

to
to
mt
-
pay

99

—"

“184
_
Ts
to

a
Loe ete eee
NODOIKH ROTI RPOQwe
=
¥
~

mit elo
="
~_
_
us
—)
|

|
measly | 6 129

one

—i— a
Cred

—

ho

sr
lompotrrcbebrorse

to
fad ped et
_
-_
RSriw——
—w
-_
lo
i

be

—_

u
t

NS yw
WNmOOlLS
i"
re
yr
|
— bo

low
mae to tole
in)
a
to
te
He OO te CO NS = BS Ot 01 GS
all apelin ee ee ee ee
LOND Ho NY bY HOLS
em He be SS Ye Gs Ge So So 8 8s CO

Rm lo to lo le ww se
—_
©
2)
l

149 TABLE

|

|
S0 | 82) 8# | 86 | 88 | 90 | 92 94 96 | 98 | 100/ 102

| |
104) 106 108} 110 | 112) 114 | 116 | 118 | 120 122 | 194

| |

Below 80,

_
tlm oO
_

~!

to

~)

_

-_
a

me eee to eh

“—~

~)

ty
Ye i

_—

i
trois
=a

as)
S
we
i
_—

—
~)
=)
.
_
—
—
CO Wee — ye an re
weve
«

—
a ee ee

tow:

sie ew =" : :
—!
=
-—etoRic-

_
_
—

au)
ae a

ero ticto

oid
25
w
-~
_
~—
=I
—
me lotr

XV IIl1l—contd.

126 | 128) 130/132 134 136 | 138 | 140 | 142

to

lo lobo

He eb

ne

el et i ON dl ee dl ol

e we lou Or lone wto lo = lo

— ee

to ~ coe lo US me os bo tor ie Kwe

loOwLoP LOLe

mde re ILI eI

wee eS

{
}
}

144 146

|

1148150, 152) 154 | 156

158 160

Above 160

|
|
|
|

Number of

plants,

—

eS hoe

—to—

to

we

bo

to

to

i — o a)

bo

—_

ee oe

_

we

toe

— i

_

——

— ee ee

lo

toe

as te SD

to

— a)

lo

es:

Parents.

hha

a lel ice el al |
o—
SSW RPUSSO BEAD OH DIR Bem IOWOWW WIA

|

! +
~~

: ods
aa el Sod
C

ea
v= lon

~le!

| Offspring.

|

128

BSeEBES

_— et eet
wwwwrnm
—— i — 7 — }

TABLE

Ez ee ee
104} 106} 108 110 112 114 116 | 118 | 120} 122 | 124

80 | 82 88 88 a0)]\02

}

100 | 102

94 | 96 | 98

Below 80.

_
me bolo

—
ee Re

:
-_
lor bo

vw
3
rm

-_—

to
to

mor
we

a
By

>
tb
©

wv

A
to

on
bo
uN

_
a)
lclbo—

~~

8

te abo

on
to
-

NG al ub ie
ee 1 OL Wore
coUuenanwwn
:
eee ee
="
—"
~ e re
_ to
= wie
Dd —
p= La [eS cll oo
= ee eS SS ee ee
_ Ree we ws _

XVI 11—coneld.

126 | 128 | 130 | 132.
|

me bob!
lo

ND cu

o
WR RK ONS =
Ww

—s)

a
eee

es bo
bo us
De we: ee — Pee
_—

-
Wek eee.
tlowe

ee OL
i

~~
~
bo

134

-

wigs ww

_

NWN a ee Sh

—

~~

to

=——"

| ; |
136/138 140 142 144 146 148 150 | 152 154

} |

to
~

eh Sd
a

=

—y

m bo

bor bo
toes

eK ES OS

lo

a)
Lad
~
we

—_
bo
—y
bo
w

wee

eto

to bo

ae

ro
weWwe

_ =
eee
me bom bo bo
=
— eee

156

158

SSS; ——__

160

tole

Above 160.

to

we

pd et et TS,

_—
Ow vi

wou,

v=

ue —w

Cnr ntovsl—w

wea
too Orle

_
_~
Sc

ee ee
w vw Ce
Oonwwu

-wwneee
Sie O10 Ye Ut te

| +
wot tT

LU tS US RL Om ee

mio

CcunnCi

tc

—— =

_

Seton Nwo

tlowocet

=nNwe

_

ee
AnNvxreiowtls

me
Se Ne

We Se Se

me WWHW HW KW ole le lot

‘sm

a

we

i

|

5,
i

}
. E
ae
ee
z

Abbas

J
%

a

-
—

g
D4

es

a
.
—

r#

‘g OdAy, X p CGAY

| 8

13

"€ aA, x Gy eUAy,

Se) bcd ”-
~_
~ —
-_
Dc] : © : : =
. _ . .

“| vunyp x [ea ....»&d . § “Gn * § eat 9 eum

Taste XIX.

4 | 06 sn 210

Parent Type 4 | eal tel tn 5/16] 15/13] 6] 9| 5| 3] 2) 5 allalallta Nie ME elle 5 ; =
(07) (1907). | |

Parent Type 3 | eee | as | fa : Albee ‘

(1907)

Fi(197) o UV) 7p 5] Vy) 3) apm) 2 3] 3) 1] 4] 2 1
Parent Type
(1908),

‘Type 4 = Types

Parent Type ~ | =| es k e 1 2
(1005)

Fy (1808) . [ee 1] 2
Parent Type 5 6} 6) 2) 7jas} | 1!) a
(1907).

©
o
2
o
-
ey
a
om
»

0

Parent Type 4 | 1 2)i)2 1] 1) 2
(1907).

Fi (1907)

Parent Typo 8] oe 1| 2 ela} a}a}a
(108).

os

Type 0 = Typo 3.

Parent ‘Type 3 |
(1908).

Fy (1908),

(1600). a. i -

Paront Type 7 2) 6) 2) 2) 1) 1 1 :

Parent Type
(1909),

2
°

Fi (1900) —|l— | 2

Parent Type

=
©
&

nits! ol vj} si ela = : i

Type? = Tre 3.

Parent Type 3 T No
(agit),

Fs (1011) - 1| 7} 10] 29 | a6
Parent Type
(1907),

t grown.

29 | 96 | a2 | 99 | 53] 48) 17 | aa) 9) | 25] 16 | 45 | ad | 25 | 13] 18

ey
e
aa
=
Se
z
=

| 8) 6

Paroot ‘Type 3 c Ay
ig} is | ss 1 1/2 1

F; (1907)

' te " ‘ + : a
Parent Type 9 , A
(1908), | | .

1
Parent Type A
tw), 77" =

Ps (1905) | 4
Parent
‘isa, 7!"

S
c
o
-
5

Type? = Type 3.

16/17) 11} 18} 7} 10] 6] S} 8

Ss
on
e
©
os

Pareot ew
‘iam. "7 : | “

Py (1005)
Pare
Toy, ee 1°

Type 10 = Type

Parent
‘om. Type 3%

¥y (1009)

vir! ., F!.

i\Banda

BI

5

82 83° 84

a
° ,
° 2

82 83 84 85 86

COTTON DISTRIBUTION IN U.P.

REFERENCE

% OF AREA KHARIF

Aipewessou... ..
Above eA ie
Above 10).

Above 5.

Below 5.

‘«
\o
>
\'
(or
9
0) 0 \
Basti Gorakhpur d
@) /
Azimgarh
ae ee
Ghazipur 4 ie
0
Ben ares
BENGAL
23 0-5 oe
Mirzapur be ere
*e
ie
4

o
On
®
O)

28

ie)
+. 0

a 86
:
CoTToN DISTRIBUTION IN U.P. >>!

REFERENCE

——% or AREA KHARIF ——

Above
Above

Above

Above

Moradabad) faayNGr Below
\a o Se
3 > JPilibhit Z}

)

RAJPUTANA

)
Sultanpur

ay Azimgarh
Part ee eS

Jaunpur
vA

IVIFAT, ©.

84

88 92 96

MEAN TEMPERATURE OF DAY}

(20 Years Average)

JULY

Mean Temperature of IndianLand Area, s3°sF ahr .

Ss T ] B BK T
coe ma
fet’ NS Brahmapwira
(3 MS
X ae SS Pt ai) Ae
G - (NH is a Te ‘ oa of)
ee War jee ing, eGantok B25. ee seus
ra fe,
a Motihari gies P71 Ae" i, BS
ee § * 85Barh ae ° Cogn Bely ; Br
Pou 5 Mbzatfaeour Dina} a aes ‘i Oy
. 2, : Ay >) oi °M “ip os, i =
ares Bha gaily 2/09 og"? _ Silchay / ¥
oe" OP ae Of fry hy NayaDu hi r ey es 7 [s se
4 Dalto erhamporé , ee ~
LO is “aribagh o a Ki dat
anc { am =
bbulpore pe at a3

° Chaib
Fendra ‘a A on Z Bo, y

i.

Od) Dalpur

)) a

Balasore

Cuttacl/} False Paint

bg
BAY

BENGAL

88 92 96

32

88

Chitrale — Gilghat*
Killa Darash «

Kabul*

» Deralsray
Khan

“Abu age e

“Deesa Pench

Veraval

BOMBAY ()

ARABIAN

LA

Ratnagiri

Nag'pur iE ipup

92
male ere ares)
MEAN TEMPERATURE OF DAY—}*°

(20 Years Average)

JULY

Mean Temperature of Indian Land Area, a3'sFahr

Brahmaputra

Seah tn
+Cooch

Y Siett
eer *Shillong ;
a ce {

rs

. ieee
Seofi

Cutta/) False Point

os

Tesi BAY

izofsapatam BENGAL
canada
5

DJ Masulipatam
2 aN

Basse

Diamond

a

92

SEASONAL RAINFALL
JUNE TO OCTOBER

Si oe °
Hazaribagh * Seq 2.
Ranchi_s Ramganj Krish
Puruli = Burd DO
? nkura
$84" Mid
Bala

iZagaputam BENGAL

Cocanada

Fy Masuliputtam

84 88 90

68

MAF. &.

| Chitral Gilghate am eh Ss

Killa Darash* ae Skardu
r
Minimars

“Dras

/
RL
Ody« We
¥ A ’ :

A f C4
§j
wan
\\ »Mussooree_
< Nehra Dun ~.

ule

TAN

28

J
<I
x,
a
—
<
ia)

=

BOMBAYY
A RABIAV

SEA

75 \P

Ratnagiri}

100

72

8 ees Pa Bha
L~ De’ ‘
aNje Ath “ey
50 Hazaribagh :
ui

92 96

SEASONAL RAINFALL
JUNE TO OCTOBER

Brahmaputra

*Muzaffalpur
Bhapra Pike e gina,

amganj Kri

ee Bankur

eee
~~ Sod

PLATE 1.

G. OBTU SIFOLIUM.—TYPE 1.

PLATE II.

G. HERBACEUM.—TYPE 2.

PEATE: Ii],

G. ARBOREUM.—TYPE 3.

PLATE IV.

TYPE 4.

V

p
At!

PLATE VI.

TYEEXS:

PLATE VII.

TYPE 6a.

PLATE VIII.

DYWPE Gk

PLATE IX.

TYRES.

PEATE XS

TYPE-S.

PLATE XI.

TYPE 10.

PLATE XII.

oe —

oF :

ee

‘acs

",

e iy, 2 Py . ;

> ", <=

"~

>

G. SANGUINEUM.—TYPE 11.

PEATE: XE

© adAL

“AIX ILW1d

Wray

ae

PY AdAL

"'€ dadAL

PLATE XVI.

20) IYPEcsexoiy Pees:

6 AdAL

"€ AdAL

TIAX FLV 1d | —

‘6 AdAL <X € AdAL JO 2y

THAX aLV1d

PLATE XIX.

+ MONOPODIAL TYPE.

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DV Noe" Ve
EN Now, I
V, No. I,
VeeNo; oT:
Ve Nose rn.
Va) No: 2EV.
VeuNO Ms
WieENGe? =a,
Vie No: IE.
VE No: LEle
WilsNo; EV;
Wiz Noi Ve
VI, No. VI.
VI,sNo; VIT.
I, No. I;
fg Ne is
i Nos LET
ia Noe. EV:
I, No. Ve
No? VE
I, No. VII.
I, No. VIII.
i NOs clcx,
RYN. 7X:
II, No. I;
LT; Note LE
Ii; No: TI
ENO. LV;

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A Sclerotial Disease of Rice, by F. J. F. SHaw, BSc. (Lond.), A.B.C.S,,
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Studies in Indian Tobaccos. No. 3.—The Inheritance of Characters in
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Studies in Indian Cottons, Part I—The Vegetative Characters, by H. M.
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Some New Sugarcane Diseases, by HK. J. BUTLER, M.B., P.L.S.; and
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Preliminary Note on the Classification of Rice in the Central Provinces,
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The Composition of the Oil Seeds of India, by J. W. LeaTHER, Ph.D., F.1.¢,
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Experiments onthe Availability of Phosphates and Potash in Soils, by
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SLUR INR ONE

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SILT; Noe LY
ELE No. Vi.

Vor TV, No. V:

Vol. IV, No. VI.
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A contribution to the Knowledge of the Black Cotton Soils of India, by W.
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The Date-Sugar Industry in Bengal. An Investigation into its Chemistry
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Evaporation from a plain Water Surface, by J. WALTER LEATHER, Ph.D.,
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Experimental Error in Sampling Sugarcane, by J. W. LEATHER, V.D.,
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The Coccide attacking the Tea Plant in India and Ceylon, by EK. EH. GREEN,
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The *' Psylla” \)isease of Indigo, by A. J. GRov®, M.Sc., and C. C, GHOsH,
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Vol.
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Camel with recommendations for Systematic treatment, by A. S. LEESE,
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No. Il. Rinderpest: Further Investigations on Questions connected with the Eco-
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An Outbreak of Cotton Pests in the Punjab, 1905, by H. M. Lerroy, M.A., #.E.S.
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Report on Cocoanut Palm Disease in Travancore, by EK. J. BUTLER, M.B., F.LS.
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Memorandum regarding Leading Eucalypts suitable for India, by F. BoorH-
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RED ROT OF ne
B. J, BUTLER, M.B., F.L.S
imperial. Myeologist

AND

ABDUL HAFIZ KHAN -

Assistant to the vs mperial Mycologist

AGRICULTURAL RESEARCH INSTITUTE, PUSA
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October 1915, BOTANICAL SERIES, Vor... VI; No; 5:

MEMOIRS OF THE
DEPARTMENT OF AGRICULTURE
IN INDIA

RED ROT OF SUGARCANE

BY

EK. J. BUTLER, M.B., F.L.S
Imperial Mycologist

AND

ABDUL HAFIZ KHAN

Assistant to the Imperial Mycologist

AGRICULTURAL RESEARCH INSTITUTE, PUSA

PUBLISHED FOR

THE IMPERIAL DEPARTMENT OF AGRICULTURE IN INDIA

BY
THACEER, SPINK & CO., CALCUTTA
W. THACKER & CO,, 2, Creep Lanz, LONDON

PRINTED RY
. THACKER, SPINK & CO., CALCUTTA. :

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“PW YORE
\NICAL
(VEN

RED ROT OF SUGARCANE.

BY
Ra BUTLER: MOB. Ls:,

Imperial Mycologqist,

AND
ABDUL HAFIZ KHAN,
Assistant to the Imperial Mycologust.

The most serious disease to which sugarcane is subject in India
is undoubtedly that known as “ red rot,” caused by the fungus Col-
letotrichum falcatum Went. In 1906, a preliminary account of its
characters and the damage caused by it in Bengal, especially that
part which is now Bihar, was published in these Memoirs.’ The
most important conclusions there come to were that the disease
ordinarily results in Northern India from the use of infected canes as
“seed” and that the most hopeful method of checking it was by
careful selection of the setts at the time of planting.

In a subsequent paper” the advantage of this practice was
emphasised, some striking illustrations being given. Further ex-
perience has only increased the evidence of the value of sett selection
which, while not an infallible preventive, is ordinarily instrumental
in greatly diminishing the incidence of the disease. It is worth
considering this evidence in detail, since infection from diseased seed
has recently been denied by American® and West Indian’ writers.

1 Butler, E. J. Fungus diseases of sugarcane in Bengal. Mem. Dept. of Agric. in India,
Bot. Ser. I, No. 3, 1906.

2 Jb. The selection of sugareane cuttings. Agric. Journ. of India, IH, 1907, p. 195.

3 Edgerton, C. W. The red rot of sugarcane. Louisiana Agric. Exper. Sta. Bull, 133,
HOEY, pelt.

‘ Red rot fungus and the sugarcane in the West Indies. Agricultural News, XI, Nos,
286-7-8, 1913.
1

152 RED ROT OF SUGARCANE,

SETT SELECTION.

In 1906, alternate rows of diseased and healthy setts of a
Madras cane, known as Yerra, were planted at Pusa, on February
26th-27th. Prior to planting, the presence of Colletotrichwm falcatum
had been determined in a large proportion of the canes from which
the diseased setts were cut. By the 6th week after planting a
decided difference was noticed in the two lots of seedlings. Those
from diseased canes were withermg in considerable numbers.
Colletotrichum falcatum was found, in practically every case
examined, in the young shoots, usually in the mycelial condition
but in several instances producing spores at the basal nodes. In
this, asin other cases where it was necessary to determine the
identity of the organisms present in diseased cane, considerable
use was made of the method of incubating aseptically removed
slabs, described on page 8 of the Memoir above referred to. White
ants, which are a frequent cause of injury to cane seedlings and
which complicate the diagnosis in many cases of fungal attack,
were absent from this particular crop, probably because the field in
which it was grown was liable to flooding. Sphawronema adiposum,
a fungus which is able to attack cut setts but not uninjured cane,
and a parasite belonging to the genus Cephalosporium, which will
be described in a subsequent paper, were found in a few cases.

The field in which this crop was grown had not been under cane
in recent years and no other cane had been grown the previous year
within about half a mile. There was, therefore, no reasonable risk
of external infection and certainly no possibility of such an infection
as would lead to the death of many plants in the trenches planted
with setts from diseased cane,while the alternate, strictly comparable,
trenches with healthy seed escaped. The photograph reproduced
as Plate XII in the Agricultural Journal of India for 1907, shows
the appearance of this plot on May 17th, and could hardly be more
decisive. Under the microscope it was easy to trace the fungus
from the setts up into the young shoots, and throughout April and
May continued infection of the young shoots from their point of

BUTLER AND HAFIZ. 153

origin occurred. On May 30th the number of sound shoots was
counted in the trenches numbered 3 to 18, of which the odd numbers
were from healthy seed and the even from diseased, the result being
679 and 117 respectively. The crop was lost by a severe flood in
August, so that the final result cannot be given.

In 1907, the experiment was repeated in a field not subject to
flooding, ten trenches being sown on March 7th with the varieties
Striped Mauritius and Red Mauritius. By the end of April the
results were as striking as in the previous year, but in May
many shoots withered in the trenches planted with healthy seed
of the Red Mauritius variety. This variety was taken from a
diseased field and was so generally infected that the ratoons nearly
all died out. It is probable that many of the apparently healthy
canes contained Colletotrichum, for, as will be shown below, while
reddening of the pith is a sure indication of disease, unless the canes
have been mechanically imjured, absence of reddening does not
always imply freedom from it.

In 1908, the cane selected was again Red Mauritius, which was
planted on March 6th. Germination was good in all the trenches.
The condition of four rows, Nos. 12 to 15, on May 30th, is graphically
shown below, the small circles each representing a sound shoot.
Rows 12 and 14 were from diseased seed, rows 13 and 15 from healthy.

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Besides these comparative experiments, the main crop of cane
grown on the Pusa Farm has been yearly supervised, so that setts
showing red marks in the pith are not planted. The result has been
that, excepting the season 1907-08, which will be separately con-

154 RED ROT OF SUGARCANE.

sidered as the cane was weakened by an attack of sugarcane-fly,
no serious outbreak of red rot has occurred since 1905-06. The
disease is always present and was fairly bad in Yerra in 1905-06
and in Red Mauritius in 1906-07, in both cases accumulating in the
following Ist ratoons so as to destroy most of the crop. In 1908-09
it was difficult to find enough diseased cane to supply cultures for
experimental work. In other years we have had 5 to 10°, diseased.
These results are probably better than in any other estate in Bihar,
though recently several estates have adopted sett selection. In one
such case, that referred to on page 198 of the Agricultural Journal
of India for 1907, a very severe attack of red rot occurred in 1905.
Sett selection has since been carried out as a routine practice and
the Manager reported recently that he has now no disease in his crop.

At the Samalkota Sugar Station in Madras, sett selection has
been regularly carried out for the past ten years, at the instance of
Dr. C. A. Barber, now Government Sugarcane Expert, Madras.
Dr. Barber was. we believe, the first to advocate this method of
fighting red rot.’ In the beginning it was combined with
picking the setts mm a strong mixture of lime water and
carbolic, with a view to checking moth-borer and the Queens-
land Acarus “rust”, but this was subsequently not considered
necessary. In 1906, Dr. Barber stated that it was not possible
by the mere rejection of red-marked setts to root out the disease’.
In spite of all care all but two of the local kinds were found
vradually to become worse, until they had to be replaced by new
seed from outside. Once these had been discarded, however, better
results were obtained. Thus the 1907 report states that selection
had a satisfactory effect, disease becoming less every year among
the best varieties. In 1908, a severe storm at the end of September
was followed by withering of a good many canes.’ Both Dr. Barber
and the senior writer saw the crop in the following February and

1 Sugareane in the Godavari and Ganjam Districts. Dept. of Land Records and Agri-
culture, Madras, Bull., Vol. Il, No. 43, 1901, p. 188.

2 Barber, C, A. Scientific Report of the Samalkota Agricultural Station for the year
ending 3lst March, 1906, p. 25.

BUTLER AND HAFIZ. 155
agreed in considering it remarkably free from red rot, and in holding
the injury to the roots caused by the storm responsible for most of
the damage. We also found evidence of the existence of a root
disease not previously known, which will be described in a subsequent
paper. In 1909-10, red rot was still present on the farm but to a far
less extent than heretofore. In 1910-11, very little disease was noted
either on the farm or in the nursery, though it was still fairly preva-
lent outside the farm. In 1911-12, red rot was present only to a
small extent, though found in nearly all the varieties. Irom these
records it is clear that red rot has not been stamped out by sett
selection. It is equally clear that the disease is much less prevalent
than in the early years of the existence of the farm. The farm
was started as a result of a disastrous outbreak of red rot in the
Godavari Delta, which threatened the extinction of cane cultivation
in that area. It 1s fair to assume that the local varieties, which were
at first grown, were very largely from diseased stock. So long
as these varieties were retained, sett selection did not give satis-
factory results. When they were replaced by other, comparatively
healthy varieties, sett selection was effective in keeping the disease
under control.

It is now necessary to consider why sett selection has proved
ineffective in checking disease when the seed was taken from a
severely diseased crop.

In selecting healthy setts for planting under field conditions
reliance must be placed on the absence of obvious reddening of the
pith, visible at the cut ends. Disease was severe in the Pusa crop
of 1907-08 and the opportunity was taken of testing how far this
method could be relied on. On November 21st, 1907, 6 canes were
selected which, on cutting into lengths, were found to have reddened
internodes above and below, but in the middle to be free from obvious
reddening. In 3 of these, careful examination with a lens revealed
one or two fine reddened points, corresponding with the cut ends of
small bundles ; the other 3 appeared quite free from discoloration.
Slabs were cut out aseptically and incubated, and the presence of
Colletotrichum falcatum was demonstrated in one of the six. On

156 RED ROT OF SUGARCANE.

March 17th, 1908, the experiment was repeated with 24 canes
slightly reddened at the base but apparently clean higher up.
Slabs were cut from above the limit of the discoloration. Of these
15 showed no marks even with a lens, while 9 had minute red points.
The presence of Colletotrichum was demonstrated in 3 of the former
and 1 of the latter. Therefore, in an attack of the severity des-
cribed, nearly 17 per cent. of apparently clean slabs, taken from
canes slightly affected with red rot, were shown to contain the
fungus within their tissues. Under such circumstances it would
have been necessary to discard all canes, any part of which was
discoloured ; new infections were occurring right up to harvest time
and some would probably have escaped even such rigorous selection.
It would be cheaper and more satisfactory under ordinary estate
conditions to discard the whole crop and import healthy seed.
In Pusa, since the amount of seed required was small and questions
of cost and trouble did not arise, it was found possible to retain
some of the more valuable varieties and the 1908-09 crop had little
disease. The amount discarded was, however, very great and would
have been ruinous under estate conditions.

All that can safely be stated, so far, is that planting setts from
obviously diseased canes leads to a considerable development of
disease in the resulting crop and that, provided the variety is fairly
free from disease to start with, sett selection keeps red rot within
reasonable limits, unless some untoward circumstance, such as_ the
epidemic of cane-fly at Pusa in 1907-08, intervenes.

THE INFECTION OF SOUND SETTS.

If there were no other method of perpetuating red rot than by
the use of diseased seed, one could, of course, stamp it out, even in
view of the above facts. This brings us to the consideration of the
parasitism of Colletotrichum falcatum and of the means at its dis-
posal for obtaining an entry into previously healthy cane. We can
then more readily discuss the further measures for its control and
the influence of external conditions. such as the attack of cane-fly
at Pusa in 1907-08. on the prevalence of the disease.

BUTLER AND HAFIZ. 157

As was pointed out formerly,’ this fungus is not, in Northern
India, provided with as suitable a mechanism for spore distribution
as in the case with most parasitic fungi. As long as the cane is
growing, there is comparatively little risk of air-borne contamination
from the stems of diseased plants in the crop. Dead and rotting
canes are, however, frequently well provided with spores. Knor-
mous numbers are often found in the pith cavities of old canes.
These may contaminate the soil or get into the irrigation water.
They may thus reach the newly planted setts. Several experiments
have been carried out to ascertain whether infection of sound setts
may take place in this manner.

In 1908, a short trench of Purple Mauritius cane was planted
on March 7th, the setts in one-half being previously dipped into a
suspension of Colletotrichum spores, from a pure culture, in distilled
water. On April 30th there were 13 living shoots belonging to 12
setts in the non-inoculated half and 6 belonging to 6 setts in the
inoculated. .

Similar experiments on a larger scale, in 1909, gave conflicting
results, both inoculated and control canes showing a number of
withered shoots after two months. On May 14th there were 167
healthy shoots and 27 withered in the control trench and 174 healthy
and 40 withered in the inoculated. White ants were bad in both,
but there was also evidence that Colletotrichwm had reached the con-
trol trench, probably on the feet of the farm labourers, who walked
up and down from trench to trench during the irrigation of the crop.

In 1910, the experiment of the previous year was repeated, the
cane being sown on March 12th. Germination was slightly better
in the control than in the inoculated trench and on May 3rd there
were 325 shoots in the former and 304 in the latter. The control
trench, which immediately adjoined the inoculated one, developed
red rot as in the previous year, probably from infection during
irrigation. A similar trench some 30 yards away, but supplied by a
separate distributary, was, therefore, also selected for comparison.
On June 21st there were only 255 healthy shoots left in the

| Butler, E. J., loc. cit., 1906, p. 16,

158 RED ROT OF SUGARCANE.

inoculated trench, while there were 340 in the control trench and 784
in the trench further away.

A form of Colletotrichum falcatum which is truly parasitic
on the leaves of sugarcane, was described and figured on page 13 and
plate III, fig. 9 of the previous Memoir, and has also been found in
Louisiana.’ The ability of this form to cause typical red rot when
introduced through setts has been demonstrated, as the following
experiments show. As these experiments have a definite bearing
on the question of the propagation of the disease through the setts
they are included here, though the leaf form of the fungus will be
more fully considered below.

On December 5th, 1907, 21 canes of Red Mauritius were
inoculated from a pure culture of the leaf parasite shaken up m
distilled water. The inoculations were done by removing a cylinder
of cane at a lower internode with a small sterile cork borer, inserting
some of the culture in the wound and replacing the cylinder after
cutting a piece off the end with a flamed knife, so as to leave a cavity.
The stem was then bound with fine sterile gutta-percha sheeting. On
March 6th, 1908, the inoculated canes were cut and examined. One
had been damaged by a jackal and was discarded. The others were
outwardly sound. With 18 of these a trench was planted on March
7th, the canes being cut into the usual setts each with three “ eyes.”
In cutting it was observed that obvious reddening of the pith occurred
in from 1 to 3 internodes above the seat of inoculation. The remain-
ing two canes were examined microscopically and the presence of Col-
letotrichum demonstrated. A similar trench was planted alongside,
with sound setts from the same plot, to serve asa control. The condi-
tion on May 30th is shown graphically below. Red rot was severe
in the inoculated trench and practically absent in the other.

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BUTLER AND HAPIZ. 159

On March 7th, 1908, a short trench of Purple Mauritius was
planted, half with setts dipped ina pure suspension of the leaf form
of Colletotrichum, the other half not inoculated. On April 30th, 62
healthy shoots belonging to 50 setts were found in the latter and
30 belonging to 26 setts in the former. Germination had been
approximately equal in the two halves.

On March 12th, 1910, half a trench was similarly planted with
setts of Ashy Mauritius, dipped in a pure suspension of the leaf
form of Colletotrichum. Germination was better in the inoculated
than in the controlhalf, and on May 3rd there were 182 shoots in
the former and 130 in the latter. On June 21st there were 77 healthy
and 80 withering shoots in the inoculated half-trench, while the
control half-trench had 156 healthy and 55 withering. As already
stated the controls this year developed red rot, probably from
infection during irrigation. A full trench of the same cane near by
but supplied by a separate distributary had, as above mentioned,
on this date 784 healthy shoots.

In all these cases the presence of Colletotrichum was demon-
strated in several of the withering shoots, and they establish fully
that true red rot can arise from infection from both the stem and the
leaf forms of the fungus through the planted setts. Not only is the
disease perpetuated by planting previously diseased setts, but
healthy setts can be infected at the time of planting, if reached by
the fungus, and, no doubt, subsequent infection from below ground
can also occur. It is well known from previous work that the fungus
can enter at wounds exposing the pith, such as the cut ends of the
setts, and, as will be shown below, infection through the roots also
readily occurs. The course of the infection up into the stem can be
traced in many cases and direct connection between the mycelium
in the sett and that in the new shoot established. Raciborski!
has very correctly described the passage of the disease from the
planted sett up into the young shoot.

' Raciborski, M. De Bestrijding van het rood-snot. Archief v.d., Java-Suikerindustrie,

V, 1897, p. 1133.

160 RED ROT OF SUGARCANE.

THE INFECTION OF GROWING CANES.

It is generally stated by workers outside India that red rot
frequently arises from wound infection of the stem of the cane,
after it has developed far enough to be exposed to the attacks of
stem borer, that is usually in the second half of its growth period.
Some observers even hold that this is the only way in which the
disease can arise. The results of numerous inoculations, indeed,
fully prove that cane can be artificially infected through wounds
similar to those caused by insects. But, as was definitely stated by
Prinsen Geerligs' in 1898, wound infection will not. sufticiently
explain every case of attack and we hope to show that in Northern
India it is of secondary importance. :

Went, who first described the disease, obtained successful
inoculations by puncturing the rind with a fine needle and inserting
conidia of Colletotrichum.’ The infection was, however, localised
and after 20 days was chiefly confined to the inoculated internode,
traces only being found in the two higher up. Attempts to inoculate
the unwounded rind failed, except when very young internodes
were selected. Went concludes that natural infection occurs
chiefly through the holes made by boring insects, but that the place
of insertion of the leaf sheath at the node is also permeable.
Howard,’ ten years later, described the results of imoculations
with the same fungus in the West Indies. When wound inocula-
tions were made on vigorously growing canes about 6 months old,
the fungus was found to have infected one or two internodes only,
after two months. In fully grown cane, during the ripening period,
however, infection was much more complete, up to 18 inches of the
pith being invaded in less than a month, in one series. Inoculations
at the leaf bases were successful in some cases but failed in others.
Lewton-Brain‘ did a limited number of inoculations in Hawaii in

1 Archief, VI, 1898, p. 450.
2 Went, F. A. F. ©. Het rood-snot. Mededeelingen Proefstation “ West-Java,” Archief

y. d. Java-Suikerindustrie, I, 1893, p. 265.

3 Howard, A. On some diseases of the sugarcane in the West Indies. Ann. of Bot.
XVII, 1903, p. 373. ;

4 Lewton-Brain, L. Red Rot of the Sugarcane stem. Exper. Sta, of the Hawaiian Sugar

Planters’ Assoc., Div. of Pathology and Physiology, Bull. 8, 1908.

BUTLER AND HAFIZ. 16]

1906-07. Wound inoculations on the stem of Yellow Caledonia
(White Tanna) canes were made. After two months the inoculated
internode was found to be infected and there appeared to be imdi-
cations that the disease was spreading through the nodes to the
internodes above and below. Ten months later, no further progress
had been made. Presumably the inoculations were made on young
plants. Infection through borer holes is considered by this writer
to be practically the only way in which fresh attacks arise, but the
propagation of the disease by planting diseased setts is accepted.
Edgerton' reports a very large series of inoculations in Louisiana.
He states that the disease spreads from the point of inoculation up
and down through the cane for from two to five jomts during the
season, but is not visible externally. Sometimes, however, if the
stalk is inoculated very young, the growth of the fungus is so rapid
that the whole stalk is killed, but this is not usually the case. In-
fection through borer burrows is stated to be by far the commonest
cause of the disease. Infection through the leaf bases and the root-
let buds at the nodes is considered possible but was not proven.
Infection through the planted setts is denied. Selection of setts 1s
advocated, not because they can carry red rot but because the
resulting crop should be superior if only healthy seed. is used.

Quite recently the results of inoculation experiments by South
and Dunlop in St. Kitts and Barbados, are described in the “* Agri-
cultural News’ (Vol. XII, Nos. 286-7-8, 1913). In the St. Kitts
experiments wound inoculations at the nodes and internodes caused
limited infection, which ceased to develop after about the first
month. In the more susceptible canes the fungus spread quickly
throughout the entire internode, but did not penetrate the joints.
The cane was strongly growing White Transparent, seven months old.
Inoculations on the leaf scars and between the leaf sheaths and the
stem failed. Attempts to infect cuttings gave more complicated
results. All the imoculated cuttings were reddened throughout
after 83 days, while only about half the controls showed reddening

1 Edgerton, C. W., loc. cit., p. 5.

162 RED ROT OF SUGARCANE.

in every node and this was less intense asa rule. The latter cuttings
had generally a white strand along the side from which the shoots
arose. Of 30 inoculated cuttings 8 growing points were found
to become diseased ; a similar condition was observed in two controls.
In one inoculated shoot Colletotrichum was identified. The control
shoots were more numerous, weighed more and were generally more
healthy in appearance than those from inoculated cuttings. In
Barbados, 40 setts each of White Transparent, Bourbon, B 147 and
B 376 were inoculated before planting, a parallel set being grown
as controls. The inoculations took, but growth within the setts
was slight, the discoloration being only visible for about 2 by 4
inch round the wound after 6 weeks. There was no definite sign of
penetration into the buds at this period. After three months there
were 7 withered shoots in the controls and 15 in the imoculated
cane. Colletotrichum could not be found in the diseased growing
tips. The healthy shoots were cut off from the moculated setts by
the formation of a woody partition. The authors conclude that
the fungus is a facultative wound parasite, infecting chiefly through
wounds such as borer holes, and not carried into the young shoots
from infected setts.

Since it is fully established that stem wounds allow of arti-
ficial infection, experiments were carried out with a view to ascer-
tain how far this is a common origin of the disease in nature. Prac-
tically the only wounds which open a passage direct to the pith
are those due to the various stem borers so commonly found in cane.
These wounds were tested for Colletotrichum on several occasions.

In November, 1907, when red rot was prevalent at Pusa, six
canes with borer holes were examined by cutting out aseptically a
slab to include the reddened area always found in the immediate
vicinity of the hole. On incubation none gave Colletotrichum.
In the same month the following year the experiment was repeated
with 9 bored canes. The result was again negative. In January,
1909, 12 bored canes were similarly examined at Samalkota, with the
same result. In addition we have examined by sectioning the neigh-
bourhood of the holes in bored canes on many occasions when the

BUTLER AND HAFIZ. 163

crop was suffermg more or less severely from red rot and have not
been able to satisfy ourselves in any case that the disease had
originated at the borer hole and not elsewhere. Yet we are
satisfied that there are other methods of infection than from below
ground and these may now be considered,

Went and Howard both refer to infection through the scar
left when the leaf sheaths break away from the stem. In December.
1907, 20 Red Mauritius canes, almost fully grown, were inoculated
at the scar left by pulling off a leaf about the centre of the stem.
The leaves were old and came away readily. The inoculated
portion was kept moist for about 24 hours by covering with moist
sterile cotton wool. After three months the canes were examined.
One was damaged by a jackal and was discarded. In 3, acervuli
with spores of Colletotrichum falcatum were found on the surface of
the scar. In 11, there was no reddening of the tissues and no sign
of penetration. In5, there was slight reddening at or near the node.
No hyphe, however, could be found and on incubating slabs, cut
so as to include the reddened parts, no Colletotrichum developed.
Even in the 3 cases where the fungus had fructified on the spot,
no penetration occurred.

At the cane node there are two other points of discontinuity
inthe rind, where the shoot bud (the “ eye”) comes through
and where the eyes of the adventitious roots occur. These were
found to admit the fungus readily. In April, 1912, 12 buds of
Samsara cane were injured by rubbing with the fingers and
inoculated with spores and mycelium kept moist by cotton wool
as before. One was examined after three days and the hyphe were
found to have penetrated the bud and to be growing vigorously.
After eleven days another was examined. The mycelium was still
confined to the bud, which was ‘much reddened. A week later
the rest were examined. All had taken the infection well and
in several the‘hyphe had already entered the main stem. Injury
to the eye’ buds, especially as the cane approaches maturity,
is unfortunately only too common, and the fungus can readily
penetrate if such buds become contaminated, The uninjured

164 RED ROT OF SUGARCANE.

bud is less readily infected. Inoculations made by placing on the
bud scales, in a moist atmosphere, fresh acervuli with spores, from
a pure culture, showed little progress at the end of a month.
There was a slight reddening of the scale, especially along the
margin and hyphe were numerous in the reddened part. The
underlying bud layers were only faintly discoloured and very few
hyphe had entered them. The deeper layers of the bud were quite
free. The buds were swelling at the time of inoculation and the
progress of the infection was so extremely slow that it is doubtful
if the young shoot could be reached before the outer layers had
withered or lost contact with it. This is in harmony with our
general experience that uninjured young shoots are not found
attacked by red rot, unless the parent stem is also infected.

The adventitious root eyes are much more easily infected.
Twelve Samsara canes were inoculated, in the same manner as in the
last experiment, on April 16th, 1912, the root origins beg promi-
nent but quite sound and uninjured. On the 24th, 9 were examined
and were all found infected, the hyphe being well established and
growing in towards the pith of the main stem. ‘The experiment
was repeated on May 22nd, 1913. Four perfectly healthy canes of a
thin variety which had a good development of young, clean, ad-
ventitious roots, varying from about one-eighth of an inch to one
inch in length, were inoculated in the laboratory in the manner
described above. The culture used was five days old and 16 roots
in all were inoculated. One root was sectioned after a week and
found to have taken infection well. The penetration of the hyphe
into the tissues was clearly visible and is shown in fig. 3. Reddening
had extended down the root and penetrated about 15 mm. into the
main stem. Characteristic hyphe of Colletotrichum were found in
the reddened part of the stem. - Two days later 3 more roots were
examined, and the same conditions found. On this day the rind at
the base of the root was found slightly discoloured and the dis-
coloration extended during the following days, until it was clearly
visible externally in all the inoculations. The normal dark green
colour of the rind changed to a dirty mottled red, which spread

BUTLER AND HAFIZ. 165

in vertically elongated streaks and, at the end of the 2nd week,
entirely surrounded the node and had extended for an inch or two
above and below. After twenty days several inoculated roots were
examined at their origin from the stem and large quantities of hyphe
were found passing from the root to the stem, not only along the
vascular tissues but also laterally into the stem parenchyma in all
directions. Fig. 5 shows the conditions at this stage. The stems
were split longitudinally a month after the imoculations and were
found entirely infected, the characteristic pith discoloration, with
transversely elongated white blotches, being well developed.

Further inoculations were made on June 3rd, 1913, on the feeding
roots of well-established cane plants growing in large culture pots
in soil. The soil was carefully removed until the roots were exposed
and these were inoculated by sprinkling with a suspension of spores
from a pure culture, care being taken not to injure the roots. The
soil was then replaced. After sixteen days two of the ioculated
roots were examined by sectioning. Both showed a small area of
reddening 2 or 3 mm. behind the growmg point. Penetration
was found to have occurred here (Fig. 4) and the hyphe were
extending freely in the tissues of the root.

Out of many hundreds of canes affected with red rot examined
during the past ten years, we have met with a limited number of
cases where natural infection had occurred through some part of the
stem above ground and where the base of the cane was unaffected.
In February, 1910, a Khari cane was examined and was found
to show definite symptoms of red rot in the 5th and 6th internodes
from the base, the lowest internodes being quite free from reddening
or hyphe. At the node between the two infected internodes there
was a broken shoot, distinctly reddened in its interior. Character-
istic hyphe of Colletotrichum were found in this shoot and could be
traced from the broken surface through the bundles into the pith
ofthe stem. There were no other injuries, and infection had doubt-
less occurred through the broken shoot. A second cane of the same
variety showed a similar case of infection through a broken shoot

at the 5th node. Higher up several nodes showed slight infection,
2

166 RED ROT OF SUGARCANE.

which was traced in every case to an infected adventitious rootlet
eye. At the 20th node there was another broken shoot, through
which infection had occurred. In all these cases, sections taken so
as to include the shoot or root and the main stem, enabled the
hyphe to be clearly traced from one to the other. In another case
infection occurred through root eyes at the 15th node and also in
the 8th internode through a crack in the rind. This is one of the
few cases where infection has been found arising from a definite
stem wound. The cane was also affected with smut (Ustilago
Sacchar?).

From the above experiments it appears that wounds caused
by boring insects, while undoubtedly capable of admitting the
parasite should it reach them, are not, in practice, responsible for
many cases of red rot in India. Old leaf scars are not readily pene-
trated, but since infection through the leaf bases has been obtained
by Howard, this probably depends on the degree to which abscission
has progressed at the time of inoculation. Under normal conditions
the leaf scars are not exposed until the leaf has completely withered
and, as our inoculations show, such scars are not readily infected.
During the process of wrapping, which is common in parts of India,
less completely withered leaves are sometimes torn away from the
stem, and the scars left in these cases are probably a source of
danger. Ina few cases the cracks which form in the internodes
of some varieties probably admit the fungus. One such case is
recorded above. But the commonest points of entry in new
infections of the above ground stem, in India, are undoubtedly
the shoot and root eyes at the nodes.

THE SOURCE OF INFECTION IN NEW ATTACKS.

We now come to another aspect of the subject and that is the
source of infection in those cases in which new attacks occur in the
cane stem. Practically all observers are agreed on the comparative
rarity of the sporing stage on the surface of diseased cane stems,
until these have dried up more or less completely. When we con-
sider the extraordinarily abundant production of spores in most

BUTLER AND HAFIZ. 167

fungi which depend on the wind for their dissemination and the chan-
ces of the individual spore alighting on a susceptible part of the cane
stem, this point becomes of significance. But, as was pointed out in
1906, there is another part of the cane on which a fungus agreeing
in morphological characters with Colletotrichum falcatum is frequently
found and produces spores in greater quantity and more exposed
to the wind than the stem form. This is the midrib of the leaf.
Earlier writers have reported the occurrence of the fungus as a
saprophyte on old, withered leaves of sugarcane.’ That it also
occurs not uncommonly as a leaf parasite seems to have escaped
the notice of most observers, though Edgerton” refers to it. Hx-
periments were carried out at Pusa to determine if the leaf form
could infect the stem and conversely.

Three of these experiments have been described above (page 158)
where it was shown, firstly, that wound inoculation with a pure
culture of the midrib form caused visible infection of from one to
three internodes after three months and, when the inoculated canes
were planted, red rot developed in them with severity, and, secondly,
that inoculation of the setts, immediately before planting, causes
just as severe disease as when the stem form of the fungus is used.
In another case a pure culture of the leaf fungus was used to in-
oculate Striped Mauritius cane, 11 inoculations bemg made to-
wards the base and ten towards the apex of the stem. Ina little
over two months, 7 of the former were examined and showed
74+34+2+1+0+4+4+4 internodes affected. In 6 of the canes
inoculated towards the apex, 3+1+0+1+1+2 mternodes were
found diseased after the same period. These experiments show
that the leaf fungus can attack the stem and the cut setts, the
symptoms produced being those of typical red rot.

The parasitism of the midrib form was next tested on leaves.
In the first experiment spores from a pure culture were sown in
drops of water on the upper surface of the uninjured midrib of
growing canes. Out of 6 inoculations, none succeeded. The ex-

’ Went, F. A. F. C. Notes on Sugarcane Diseases, Ann. of Bot, X, 1896, p. 588.
2 Edgerton, loc. cit., p. 4.

168 RED ROT OF SUGARCANE.

periment was repeated on 4 shoots in the laboratory, kept under
bell jars to prevent the inoculated spot from drying rapidly. These
also failed. Ina third series, similar to the last, 6 moculations were
made without result. A month later, however, out of 12 similar
inoculations, 5 succeeded. When the midrib was wounded, much
better results were obtained. In the first trial 5 leaves were inocu-
lated in plants growing in tubs in the laboratory, the upper epider-
mus being first removed by scraping with a sterile knife and the spot,
after inoculation, bemg covered with damp sterile cotton wool to
keep moist. All succeeded well, the characteristic red discoloration
being well developed by the 9thday. In another series, 4 inoculations
were made after injuring the epidermis of the midrib by touching
it with a hot knife blade for 2 or 3 seconds. All took the infection
severely. Four more were inoculated on another occasion, after
scraping off the epidermis, and again all took.

Experiments were next made to test the parasitism of the stem
form of Colletotrichum falcatum on the leaves. In the first experi-
ment, spores from a pure culture were sown in drops of water on the
upper surface of the uninjured midrib of canes growing ina tub
in the laboratory. Of 15 inoculations, none succeeded. The ex-
periment was again tried and of 7 inoculations, all succeeded. In
a third series, out of 13 inoculations, 5 succeeded. When the midrib
was wounded before inoculation the results were as follows. In
the first trial 5 inoculations were made after scraping off the epi-
dermis. The inoculated spot was covered with a pad of damp
sterile cotton wool. None succeeded, the fungus growing by choice
into the cotton wool. A similar experiment at a later date was
made on 3 shoots standing in water, no cotton wool being used but
the shoots being covered by bell jars. All succeeded. In a third
experiment, the epidermis was injured by touching with a hot knife
blade and all of 4 inoculations succeeded.

Though both leaf and stem forms, are capable of penetrating
uninjured leaves, infection occurs much more readily when the
leaf is wounded. The microscopic details of penetration will be
described below. In nature, it has been observed that Colletotri-

BUTLER AND HAFIZ. 169

chum is common around the hole which a minute boring insect
frequently makes in the midrib. Salmon" has found in similar
experiments with the mildews (Hrysiphacew) that “ green fly ”
(Aphis) has the same effect as a wound, in weakening the resistance
of the plant cells to infection.

From these experiments it is apparent that there is no essential
difference in the ability of the forms of Colletotrichum, found on the
living midrib of the leaf and on the stem, to attack stems and leaves
of sugarcane. Taken in conjunction with their morphological
similarity, they must be held to be the same fungus. The species
appears to be confined to sugarcane. The only other Colletotrichum
resembling C. falcatum found widely distributed in India, is C.
Lineola Corda, which attacks the leaves of jowar (Andropogon
Sorghum) frequently. Morphologically the two species are closely
allied, but the jowar fungus does not attack cane leaves. Out of
16 inoculations, half on unwounded spots on the midrib, half after
scraping off the epidermis, none succeeded. Kdgerton” failed to get
symptoms of red rot by inoculating sugarcane stems with this
species and also with the allied C. cereale. It is probable that
many of the new attacks of red rot on the above-ground part of
the cane stem, arise as a result of infection by spores blown
from the diseased midribs of cane leaves. We do not see how it
will ever be possible to avoid this, but it only gives greater force
to the arguments in favour of concentrating attention on the
elimination of diseased setts at the time of planting.

The actual penetration of the fungus into the tissues was studied
in the leaf inoculations. Including both leaf and stem forms,
altogether 63 inoculations were made on the uninjured midrib, of
which 17 succeeded and 46 failed ; while when the epidermis was
wounded 20 out of 25 inoculations succeeded. In the successful
cases where the epidermis was uninjured, penetration usually oc-
curred not directly into the midrib but by superficial growth of the
fungus until the large motor cells lying in groups on either side

99

1 Salmon, E. 8. Cultural experiments with “ Biologic forms” of the LErysiphacee.
Phil. Trans. Royal Society, Ser. B, Vol. 197, 1904, p. 112.

2 Edgerton, C. W., loc. cit. p. 7.

170 RED ROT OF SUGARCANE.

of the midrib were reached ; these were then penetrated (Fig. 1).
As is already known, the germ-tubes of the spores readily form
appressoria (described under the name of “ gemme”” by Went
and of chlamydospores by most other writers). These are thick-
walled, durable cells, capable of surviving detachment from the
mycelium, It is probable that they serve a double purpose
of close adhesion to the surface of the host plant and of accumu-
lation of enzymic energy to secure penetration of its walls. In
Colletotrichum falcatum the infection hypha seems to arise as
a rule from an appressorium (Figs. 1 and 2). Entry through
stomata was not observed, the infection hypha passing directly
across the outer wall of an epidermal cell, or, in some cases,
down between the side walls of two cells. After entry, the
hyphz may at once branch freely and fill the large motor cells
with a mass of mycelium, or may penetrate deeply into the leaf,
passing from cell to cell with ease in the large-celled parenchyma
between the bundles, but not readily entering the latter. In
some cases the sclerenchyma was penetrated, but usually, when
the leaf had not been injured before inoculation, the hyphe
remained, for the first week at least, confined to the thin-walled
cells. In the cases where the leaf was first wounded, conditions
were somewhat different. There was a superficial growth as
before, extending beyond the limits of the wound. But penetra-
tion was not now confined to the thin-walled motor cells but
occurred freely into the epidermis of the midrib just beyond
the wounded part (Fig. 2), and the mycelium ramified through
the sclerenchyma as readily as through the parenchyma. In some
cases, the layers of underlying sclerenchyma left after removing
the epidermis, were an effective barrier and penetration only
occurred beyond the midrib into the thinner tissues between the
bundles, but in others, presumably when the injury was more
severe, the thick-walled tissues showed much mycelium. All the
invaded tissues developed a bright red colour.

In less than a week fructification may occur on the infected
spot. The hyphee first collect in masses in the epidermis, through

BUTLER AND HAFIZ. 171

which they then break as stromatic bodies, on which the character-
istic spores and sterile hairs are formed.

RELATIVE SUSCEPTIBILITY OF THE TOP AND BOTTOM OF
THE CANE.

Experiments were recorded in the previous Memoir’ to show
that much of the damage caused by red rot is due to inversion of
the cane sugar. This appears to be because glucose is more readily
assimilated by the fungus, growth in solutions in which the sugar
was provided as glucose, being invariably better, at least at first,
than when cane sugar was supplied. Flasks were prepared with
solutions containing 10 per cent. cane sugar and glucose respectively
together with peptone and sodium chloride, and inoculated with a
loop of a suspension of spores, from a pure culture, in distilled water.
The growth in the glucose flasks early took the lead and maintained
its superiority for some weeks. More recently, Lewton-Brain* has
dealt with the same question in considerable detail. He found that
the inverting action of the fungus was considerable, but that, as
stated in the previous Memoir, the actual consumption of sugar was
small. This consumption, he found, fell entirely on the levulose.
Thus in one experiment, though 75 per cent. of the sucrose was
inverted, not one-twentieth part of this was actually consumed by
the fungus and this appeared to be all levulose. ‘The inversion was
proved to be due to the presence of invertase, which was found both

in the mycelium and also in the solution in which the fungus was
grown.

Since it is known that the upper portion of the cane is richer
in glucose, though poorer in total sugars, than the lower’, experi-
ments were made to compare the growth of the fungus in the top
and bottom portions. In the first experiment, 3 sound Striped
Mauritius canes were cut and brought to the laboratory. They were
each inoculated at a lower and an upper internode by removing

1 Butler, E. J., loc. cit., 1906, p. 7.
2 Lewton-Brain, L., loc. cit., 1908, p. 32.
3 Leather, J. W. Chemical composition of sugarcane and raw sugars. Agric, Ledger,

No. 3 of 1897, p. 13.

172 RED ROT OF SUGARCANE.

a cylinder aseptically with a small cork borer and inserting a small
quantity of a pure culture of Colletotrichum. After eight days they
were examined. The lower inoculations were found to have infec-
ted 2+2+1 internodes respectively, the upper 1+2+4. The ex-
periment was repeated with Samsara canes, 6 being inoculated in
the same way. After seven days one was examined and was found
to have 2 internodes infected at the base and 1 at the top. Two
days later another was examined, 5 and 2 internodes respectively
being found infected. Two days later the rest were examined. In
one the infected portions had united in the middle. In the other
three, 11+10+7 internodes were found infected at the base and
3+5+2 at the top. The experiment on page 167, where inocula-
tious with the leaf form of the fungus were made at the top and
bottom of growing canes, should also be compared. As the prac-
tical pot at issue was to determine if any recommendations could
be made for planting one part of the cane rather than another,
where red rot is prevalent, the natural inversion that goes on after
cutting was not taken into account, since it must equally go on in
planted setts before germination. The experiments do not suggest
that tops, though richer in glucose, can be more rapidly invaded by
Colletotrichum than the rest of the cane and there appears to be no
objection to their use from this point of view. The second experi-
ment and that given on page 167, indeed suggest that the contrary
is the case. Tops are also less likely to contain the fungus, when it
has originated from below at a late stage in the growth of the cane.

CONTROL OF THE DISEASE,

The control of red rot was stated by the earlier investigators
to be likely to be very difficult, owing to its position in the interior
of the cane, the frequent absence of definite symptoms by which it
might be detected in the growing crop and the practical impossibility
of preventing wounds which would give an entry to the fungus.
But of recent vears little has been heard of the disease in Java,
where it was first described, and it may be concluded that it has not
proved so serious an enemy as was once feared.

Qo

BUTLER AND HAFIZ. Wie

In India it is in many places the greatest obstacle to successful
cane cultivation. In Madras. Bombay and Bengal the area under
thick cane has in certain districts periodically shrunk as a_ result
of an accumulation of red rot in the crop, to expand again only when
the diseased cane has been replaced from outside. Red rot has often
been the limiting factor to the successful cultivation of heavy yield-
ing canes.

The experiments given above are, we think, sufficient grounds
for holding that this should not be the case; that, granted that a
start is made with a healthy stock, it should be possible to keep
the disease under control with no more than an occasional severe
outbreak due to a specially unfavourable season.

The first requirement is to start with a healthy stock. In those
districts, such as the Godavari Delta and some parts of Bombay
Presidency,’ where the local canes have become widely infected,
new healthy seed must be brought in from outside. The very
successful history of the Samalkota Sugar Station in the Godavari
Delta, shows what excellent results may be obtained by this measure,
under efficient supervision. It is highly probable that the little
that has been heard of red rot in Java, in recent years, has been due
to the efforts made to obtain good cane for planting, from special
seed nurseries, combined with the growth of seedling canes which
will be referred to below. As the Samalkota results are available
in the Annual Reports of the Station, the methods adopted need not
be more fully detailed. The past history of the cane must be
taken into consideration. There is good evidence to show how
dangerous it is to grow a variety from stock known to have been
seriously infected, even though the crop may do well for the first
few years. Large estates or groups of estates should be self-con-
tained in the matter of seed supply and should possess a nursery
or testing garden for the trial of new varieties under the best condi-
tions. For the ordinary native cultivation, Government Farms
should be utilized tor the same purpose. In this way a supply of

' Kulkarni, G. 8. Preliminary study of the red rot of sugarcane in the Bombay
Presidency. Dept. of Agric., Bombay, Bull. No. 44, 1911, pp. 5-6.

174 RED ROT OF SUGARCANE.

healthy seed can be assured and new varieties can be introduced
into cultivation as required.

Systematic and thorough selection of the setts used for planting
must then be done each year or the new varieties will not maintain
their freedom from disease for long. The methods to adopt are
described in a previous paper' which should be referred to, and
present no difficulties in practice. We consider that there is no
single operation in the cultivation of thick canes in most parts of
India of greater importance than this. It is not to be anticipated
that the disease can be got rid of by a single selection nor, indeed,
usually got rid of in its entirety by annual selection ; the most that
is claimed is that it can be kept within reasonable limits in normal
years and with good cultivation. The object of the selection 1s to
prevent an accumulation of red rot to such an extent as materially
to reduce profits and render it difficult to obtain a sufficient supply
of sound seed for the coming season.

Of lesser importance, but still worth doing in most cases, 1s the
regular removal of all withering clumps during the growing and
ripening season. Such clumps, if left, dry up and produce spores,
sometimes in considerable quantity. Infection of even perfectly
sound cane through the aerial root eyes and through injured buds
has been shown to occur and though our experience in Northern
India has been that such infection is not common, it is perhaps more
frequent in Madras, where the disease appears to be more virulent
and rapid in its onset than with us. There is, also, the danger of
infection through the soil, especially in irrigated cane, by means
of the shed spores.

Judging from Godavari experience it is important to give cane a
long rotation. How far this results from the peculiar circumstances
of cane cultivation in heavy paddy soils, with very frequent irriga-
tion, is not clear. Inthe Godavari Delta, the old practice was eight
or nine years’ rest from cane after a two years’ cane crop (one year
plant cane and one year ratoon). More recently, it has been reduced

1 Butler, E. J., loc. cit., 1907.

BUTLER AND HAFIZ. 175

to six years, probably with bad results.'| At Samalkota a two years’
“dry” rotation (not followed by paddy) and a three years’ “ wet ”’
rotation (followed by paddy) were tried, but found unsatisfactory.
On the other hand, at Pusa, cane one year in five has been satisfactory
and the non-irrigated cane in Bihar usually gets a much shorter
rotation. The fungus appears to die out rapidly in moist soil, but
cultures exposed to the air and kept moderately dry retain their
vitality for at least five months. In Hawaii, Lewton-Brain found
that plate cultures allowed to dry out invariably gave no sign of
vitality after three months. But if cane is present, there seems to be
little doubt that Colletotrichum lives and spreads through the soil
and, in the young irrigated crop, passes from trench to trench, either
with the seepage of irrigation water, on the feet and implements of
coolies or (though less certainly) by direct growth through the soil.
It has been shown that the roots are readily infected and we have
lost several series of comparative experiments at Pusa through
ground infection of control trenches. Fortunately the radius of
spread does not appear to be large and if the measures detailed
above are carried out, little injury should be caused in this manner.
In spite of all these precautions, serious attacks of red rot,
from circumstances not ordinarily under control, may occur from
time to time. We have met with two such cases. One was the out-
break at Pusa in 1907-08, which was, without doubt, the result of an
epidemic of cane-fly (Pyrilla aberrans) on the cane that year.
Leaf-hoppers are well known in other countries to be associated with
bad attacks of fungus diseases as, for instance, in Hawati, where
“vind disease ’” follows with great intensity the epidemics of leaf-
hoppers. It is probable that the action of these insects is chiefly
to reduce the vitality of the cane and render it increasingly sus-

1 Wood, R. C. Scientific Report for the Samaikota Agricultural Station for 1908-09.
Government Press, Madras, 1909.

2 The cause cf this disease is not yet quite clear. Howard in the West Indies and
Went in Java held that it was identical with red rot, the fungus, Melanconium Sacchari,
previously believed to be its cause, being merely a follower of Colletotrichum falcatum.
Subsequent workers, such as Lewton-Brain, Cobb and Edgerton in the West Indies, Hawaii
and Louisiana, have reverted to the previous view, though quite recently South and Dunlop
have failed to establish the parasitism of Melanconium Sacchari. In the East, this fungus has
not been recorded as a parasite.

176 RED ROT OF SUGARCANE.

ceptible to infection from aerial spores, which were formed i con-
siderable quantity on the leaves during the outbreak at Pusa. The
other case was a severe attack of red rot following extensive flooding
of the cane fields, in some estates in Bihar several years ago. The
effect of unfavourable external conditions such as this on the onset
of the disease is discussed more fully below.

SUSCEPTIBILITY OF VARIETIES OF CANE.

Little light has been thrown by these investigations on the
question of the relative susceptibility of different varieties of cane
to red rot. One fact that 1s obvious to any observer of the disease
in India is that the thin varieties of cane are, on the whole, less sus-
ceptible than the thick. Some of the Indian thin canes are so
widely divergent from the thick races, that writers in Java have
suggested that they may have originated from different species of
Saccharum. If so, one may perhaps anticipate that the relative
immunity of the thin kinds will prove to be a deep seated “ germ ~
character. With regard to the thick canes, certain observers.
and in particular Dr. C. A. Barber, hold that the temporary or
apparent immunity of certain thick varieties can be broken down
by bad cultivation. He describes’ how, in the Godavari Delta,
successive canes have held favour during the past forty years, each
in turn growing luxuriantly and bringing wealth, but after a few
vears becoming diseased. The constitution of each cane had been
broken in turn by the ever-present fungus, until all the plants of that
kind in the district were infected. Again he states’ that of the
varieties of cane brought from other countries for trial at Samalkota,
none were really immune and it is probable that ultimately all will
succumb in turn when placed under the adverse conditions of the
local agriculture. In the same paper’ he mentions as a curious fact
that the Hospet cane varies greatly in its liability to disease in the
different regions where it is found. A similar case occurred with the

1 Kobus, J. D. Overzicht van het verloop der importatieplannen van vreemde rietsoorten
op een eiland buiten Java: Archief v. d. Java-Suikerindustrie, IIT, 1894, p. 662.

2 Barber, C. A. The Samalkota Sugarcane Farm. Agric. Journ. of India I, 1906, p. 45.

8 Jb. Seedling canes in India. Agric. Journ. of India, VII, 1912, p. 324.

# Ib. loc. cit. p. 329.

BUTLER AND HAFIZ. 177

Bombay Pundia cane which, when introduced at Samalkota, went
out from disease in the first season, though the parent stock
is little subject to red rot. Kulkarni! also notes the gradual
deterioration of thick canes in parts of Bombay,.and West Indian
literature is full of references to the breaking down in health of the
Bourbon cane, once widely grown, and this breaking down seems
to have gone on more rapidly in some localities than in others.
These experiences with sugarcane are by no means unique
amongst plants ordinarily propagated in a vegetative manner, that
is by tubers, cuttings and the like. In Ireland the “ Champion”
potato was largely cultivated for many years, on account of its
resistance to potato blight, but it has lost its resisting powers and
been replaced by newer “seedling” varieties. Several similar
cases are known, and the phenomenon is of considerable scientific
interest.” Such progressive deterioration is in many cases
apparently innate and has been likened to senescence, being capable
of being checked when a new generation is started by sexual!
reproduction (e.g., by raising from seed). But it can be hastened
by exposure to unfavourable conditions or, on the other hand, be
postponed by profound change in the environment. Barber states”
that the surest way to induce red rot in cane to make its appearance
is to plant the canes in a water-logged site. Harrison, in British
Guiana, considers that the susceptibility of certain kinds of plants
for instance, the Bourbon cane, to fungus attacks is due in part at
least to defective soil conditions.’ Such statements can be multi-
plied and must, we think, indicate a real phenomenon. In the
opposite direction are such cases as that recorded by Calkins,’ where
a constantly changing culture medium was found to have the effect

1 Kulkarni, G. 8. Preliminary study of the red rot of sugarcane in the Bombay Presidency.
Department of Agriculture, Bombay, Bull. No. 44, 1911.

2c, f. Hartog, M. Problems of Life and Reproduction. Progressive Science Series,
LOLS p. L9-

3 Barber, C. A. The Samalkota Sugarcane Farm. Agric. Journ. of India, I, 1906,
p. 45.

* Harrison, J. B. Varieties of Sugarcane and Manurial experiments in British Guiana.
West Indian Bulletin, LX, 1909, p. 36.

® Calkins, Gary N. Protozoology, 1910, p, 109.

178 RED ROT OF SUGARCANE.

of prolonging the life of the race in Protozoa, bred from a single
individual and not permitted to conjugate; when asexual pro-
pagation only is allowed and the environment kept constant, the
race soon degenerates and dies out.!

It is not unlikely, therefore that. there are two types of relative
immunity, a genetic, such as is shown by the thin canes so widely
cultivated in India, and a fluctuating, and that the latter is much
more exposed to the influence of external conditions than the former.
Indian thick canes, which have been subjected to vegetative propa-
gation without a break for many generations, seem to show no
evidence of genetic immunity. Hence frequent change of climate,
good cultivation and good hygienic conditions generally, seem to
be of great importance in preserving them from epidemics of disease.

The possession, by India, of a large range of relatively immune
(to red rot) thin canes, of hardy habit and great tillering powers,
though less productive than the thick canes of other countries,
may prove an asset of the greatest value. The growth of seedling
canes has been recently undertaken at the Coimbatore cane-breeding
station under the control of Dr. Barber. If it is found possible by
hybridization to combine the resistance of some of the thin canes to
red rot, with the yielding qualities of a thick cane, a great step for-
ward in enabling India to grow enough sugar for her own consump-
tion and perhaps even to compete successfully with the sugar
exporting countries, will have been taken. It is a happy augury
that amongst the best of the canes now grown in Java (the most
formidable competitor in sugar production that India has to meet)
are the progeny of crosses between an Indian thin cane, the
“Chunnee” obtained from Shahjahanpur (where, Dr. Barber
informs us, it is locally called Chin), and the Cheribon thick cane
grown in Java.

Pusa,
June 30th, 1913.

1 Since this was written a very illuminating discussion of the deterioration of
sugarcane varieties after long-continued vegetative propagation, written by Harrison, Stock-
dale and Ward (West Indian Bulletin, XIII, 1913, p. 177), has been received, ’

PLATE

iM iin
eS NN Mi EQOA OE)

TANK
/ Mi
WH |
IH HIN

I

Infection of suga !

r cane leaves an
Colletotrichum falcatum,

d roots by the red rot fungus,

Z rey ea

Fig.

Fig.

DESCRIPTION OF THE PLATE.

Penetration of a hypha, arising from an appressorium of the stem form of
Colletotrichum falcatum, into a motor cell of the unwounded leaf of
sugarcane. The midrib commences immediately on the right of the
figure. X 350.

Penetration of a hypha, arising from an appressorium of the stem form of
Colletotrichum falcatum, ito a cell of the thick-walled part of the epi-
dermis at the edge of the midrib of the sugarcane leaf. The neighbour-
ing central part of the midrib had been wounded by scraping off the
outer layers with a sterile knife. X 500.

Penetration of an adventitious root on the stem of sugarcane by hyphe
of Colletotrichum falcatum, through the uninjured epidermis. X 500.

Penetration of an underground root of sugarcane by hyphe of Colletotri-
chum falcatum, through the uninjured epidermis. X 300.

Passage of the mycelium of Colletotrichum falcatum froma young inocu-
lated adventitious root, of the same series from which fig. 3 was drawn,
back to the main stem of the sugarcane. The hyphe enter the vascular
tissue and the parenchyma of the stem and spread in all directions,

X about 50.

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PENG. LUT
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VN Ole vel
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iT No. AI.
DIS No 11.
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II; No. VII.

II, No. VIII.

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iLT,:No, A.

TL, No. II.

III, No. III.

Lit, No. LV.

Tit, No: Vv.

Lil, No. Vi,

BOTANICAL SERIES.

Studies in Root Parasitism. The Haustorium of Santalum album.—
Part I.—Early Stages by C. A. BARBER, M.A., F.L.S. Price, Re. 1.

Part I1.—The Structure of the Mature Haustorium and the Inter-relations
between Host and Parasite by C. A. BARBER, M.A., F.L.S. Price, Rs. 3.
(Out of print.) :

Indian Wheat Rusts by E. J. BUTLER, M.B., F.L.S., and J. M. HAYMAN,
D.vV.s. Price, Rs. 3. (Out of print.)

Fungus Diseases of Sugarcane in Bengal by E. J. BUTLER, M.B., F.L.S.
Price, Rs. 3.

Gossypium obtusifolium, Roxburgh, by I. H. BURKILL, M.A. Price, Re. 1.

An Account of the Genus Pythium and some Chytridiacee by E. J. BUTLER,
M.B., F.L.S. Price, Rs. 4-8.

Cephaleuros virescens, Kunze ; The Red Rust of Tea by HAROLD H. MANN,
pD.Ssc.; and C. M. HUTCHINSON, B.A. Price, Rs. 4. (Out of print.)

Some Diseases of Cereals caused by Sclerospora graminicola by E. J.
BUTLER, M.B., F.U.S. Price, Re. 1-8.

The Indian Cottons by G. A. GAMMIE, F.L.S. Price, Rs. 7-8. (Out of print.)

Note on a Toxic Substance excreted by the Roots of Plants by F.
FLETCHER, M.A., B.Sc. Price, Re. 1-8.

Studies in Root Parasitism. III..—The Haustorium of Olax scandens by
C. A. BARBER, M.A., F.L.S. Price, Rs. 2-8.

Studies in Root Parasitism. IV.—The Haustorium of Cansjera Rheedii by
C. A. BARBER, M.A., F.LS. Price, Rs. 2-8. (Out of print.)

Some Experimentsin the Hybridising of Indian Cottons by P. F. Fyson,
B.A., F.L.S. Price, Re. 1-8. (Out of print.)

The Varietal Characters of Indian Wheats by ALBERT HOWARD, M.A.,
A.R.C.8., F.L.S.; and GABRIELLE L. C. HOWARD, M.A. Price, Re. 1.
(Out of print.)

The Mulberry Disease caused by Corynewm mori, Nom., in Kashmir, with
Notes on other Mulberry Diseases, by E. J. BUTLER, M.B., F.L.S. Price,
Re. 1-8. (Out of print.)

The Wilt Disease of Pigeon-Pea and the Parasitism of Neocosmospora
vasinfecta, Smith, by E. J. BUTLER, M.B., F.L.S. Price, Rs. 5.

Studies in Indian Tobaccos. No. I.—The Types of Nicotiana rustica,
L., Yellow Flowered Tobacco by ALBERT HOWARD, M A, A.R.C.S., F.L.S. }
and GABRIELLE L. C. HOWARD, M.A. Price, Rs. 4.

Studies in Indian Tobaccos. No. 11.—The Types of Nicotiana tabacum,
L., by ALBERT HOWARD, M.A., A.R.C.S., F.L.S.; and GABRIELLE L. C.
HOWARD, M.A. Price, Rs. 9.

Studies in Indian Fibre Plants. No. I.—On two varieties of Sann,
Crotalaria juncea, L., by ALBERT HOWARD, M.A., A.R.C.S., F.L.S.; and
GABRIELLE L. C. HOWARD, M.A. Price, Re. 1.

The Influence of Environment on the Milling and Baking Qualities of
Wheat in India. No. I.—The Experiments of 1907-08 and 1908-09. By
ALBERT HOWARD, M.A., A.R.C.S., F-L.8.; H. M. LEAKE, M.A., F.L.S.; and
GABRIELLE L. C. HOWARD, M.A. Price, Re. 1-8.

The Bud-Rot of Palms in India by E. J. BUTLER, M.B., F.LS.
Price, Rs. 2.

The Economic Significance of Natural Cross-fertilization in India by ALBERT
HOWARD, M.A., A.R.C.S., F.L.S.; GABRIELLE L. C. HOWARD, M.A.; and
ABDUR RAHMAN KHAN. Price, Rs. 4-8.

iViolemvi, Nos lk
Wole, LV, No. 11:
Wols sbVeoNo: LET.
Vol. IV, No. IV.
Vol. IV, No. V.
Wol-me Ve NOs Wl.
WiolsoewV.. Nos I.
Wolk aVeyNO. LL.
NOlrae Vit NOs DE.
Vols iV-No: LV.
NiOlsme Ven Nios. NV.
WOlneVie No. 1.
Vol: Vi, No. If.
Vol. VI, No. III.
Vol. VI, No. IV.
WolenVil-No: V.
Vol. VI, No. VI.

Vol. I, No. if
Vol. I, No. II,
Voll; No: SLD:
Vol. I, No. IV.
Violen INO. Vs
Viol. i, Now VI-
Wolva ts No.. WEE:
Vol. T, No. VIII.
WOle LINO. ul xc
iol iINOss ok.

BOTANICAL SERIES—contd.

The Millets of the Genus Setavia in the Bombay Presidency and Sind,
G. A. GAMMIE, F.L.S., Imperial Cotton Specialist. Price, Re. 1.

Studies in Indian Fibre Plants. No. 2.—On Some New Varieties of Hibiscus
cannabinus, L., and Hibiscus Sabdariffa, L., by ALBERT Howarb,
M.A., A.R.C.S., F.L.S.; and GABRIELLE L. C. HOWARD, M.A. Price, Rs. 3.

Notes on the Incidence and Effect of Sterility and Cross-fertilization in
the Indian Cottons, by H. M. LEAKE, M.A. (Cantab.), F.L.S.; and Ram
PRASAD. Price, Re. 1.

Note on the Inheritance of Red Colour and the regularity of self-fertiliz-
ation in Corchorus capsularis, the common Jute Plant, by I. H.
BURKILL, M.A.; and R. 8S. FINLOW, B.Sc., F.C.S. Price, Re. 1.

Observations on Certain Extra-Indian Asiatic Cottons, by H. M. Leaks,
M.A., F.L.S.;and RAM PRASAD, Asst. to the Economic Botanist, U. P.
Price, Re. 13:

The Morphology and Parasitism of Rhizoctonia, by F. J. F. SHAw, B.Sc.,
A.R.C.S., F.L.S. Price, Rs. 2.

The Inheritance of some Characters in Wheat, I, by A. HOWARD, M.A.,
A.R.C-S., F.L.S. ; and GABRIELLE L. C. HOWARD, M.A. Price, Re. 1.

The Influence of the Environment on the Milling and Baking Qualities
of Indian Wheats. No. 2.—The Experiments of 1909-10 and 1910-11, by
A. HOWARD, M.A., A.R.C.8., F.L.S.; H. M. LEAKE, M.A., F.L.S.; and
GABRIELLE L. C. HOWARD, M.A. Price, Re. 1.

The Varieties of Soy Beans found in Bengal, Bihar and Orissa and their
Commercial possibilities, by E. J. WOODHOUSE, M.A.; and C. S. TAYLOR,
B.A. Price, Rs. 2:

On Phytophthora parasitica nov. spec. A new Disease of the Castor Oil
Plant, by J. F. DAstour, BsSc., First Asst. to the Imperial Mycologist.
Price, Rs. 2.

Studies in Peronosporacee, by E. J. BUTLER, M.B., F. L. S., Imperial
Mycologist ;and G.S. KULKARNI, L.Ag., Mycological ‘Assistant, Bombay.
Price, Rs. 2,

Notes on Pollination and ce Rerulieation in the Common Rice
Plant, Oryza sativa, Linn., by G. P. HECTOR, M.A-, B.Sc. Price, Re. 1

A Sclerotial Disease of Rice, by F. J. F. SHaw, B.Sc. (Lond.), A4.R.C.S.,
F.L.S. -Price, Re. 1.

Inheritance of Characters in Nicotiana tabacum, by GABRIELLE L. C-
HOWARD, M.A. (Jn the press.)

Studies in Indian Cottons, Part I—The Vegetative Characters by H. M
LEAKE, M-A., F.L-S.;and RAM PRasabD. (In the press.)

The Red Rot of pias by E. J. BUTLER, M.B., F.L.S.; and A. HaFIz
KHAN. Price, Re. 1

by

Some New chen Diseases, by E. J. BUTLER, M.B., F.L.S.; and
A. HAFIZ KHAN. (In the press.)
CHEMICAL SERIES.
The Composition of Indian Rain and Dew, by J. WALrEK LEATHER,

PSDs, Est. C9 serices hes 1:

The Composition of the Oil Seeds of India, by J. W. LEATHER, Ph.D.,
Price, Re. 1. (Out of print.)

The Pot-Culture House at the Agricultural Research
J. W. LEATHER, Ph.D., F.1.c. Price, Rs. 3.

Experiments on the Availability of Phosphates and Potash in Soils, by
J. W. LEATHER, Ph.D., F.1.C. Price, Re. 1-8.

The Construction of Drain Gauges at Pusa, by M. H. ARNOTT, M.INS'.C.E.,

F.1.C.

Institute, Pusa, by

with a Preface by J. W. LEATHER, Ph.D., F.1.c. Price, Rs. 3. (Out of
print.)

The Loss of Water from Soil during Dry Weather, by J. WALTER
LEATHER, Ph.D., F.1.C. Price, Rs. 2. (Out of print.)

The System Water, Calcium Carbonate, Carbonic Acid, by J. WALTER
LEATHER, Ph.D., F.1.C. ; and JATINDRA NATH SEN, M.A., F.C.S.

Price, Re. 1.

Water Requirements of Crops in India, by |J.
FEC: -Lrice, Rs. 3:

The Nature of the Colour of Black Cotton Soil,
(Lond.), F.C.S., M.S.E.A.C. Pricé, Re.

Water Requirements of Crops in India—II, by J. Moe as LEATHER?
Ph.D., F.1.C., Imperial Agricultural Chemist. Price, Rs. 2-

WALTER LEATHER, Phi.Db.,

by H. E. ANNETT, B.Sc

Vol. I, No.
Vol. II, No.
Vol. II, No.
Vol. IT, No.
Vol. II, No.
WMolslt; No;
Vol. III, No.
Vol. III, No.
Vol. ILI, No.
Vol. III, No.
Vol. I, No.
Vol TNO:
Vol. I, No.
Vol. IT, No.
Vol. J, No.
Vol. I, No.
Vol. II, No.
Vol. II, No.
Vol. II, No.
Vol. II, No.
Vol. II, No.
Vol. II, No.
Vol. IT, No.
Vol. II, No. VIII.
Vol. II, No.
Vol. II, No.
Vol. III.

RY.

We

Ill.
Ve

X.

CHEMICAL SERIE S—conid.

The Composition of the Milk of some Breeds of Indian Cows and Buffaloes
and its Variations, Part I, by A. A. MeraGIrt, B.Sc. (Lond.), F.C.s. ;
and H. H. MANN, D.Sc. Price, Re. 1-8.

Records of Drainage in India, by J. WALTER LEATHER, Ph.D., F.I.C.
Price, Re. 1

The Rab System of Rice Cultivation in Western India, by H. H. Mann,
p.sc.; N.-V. JOSHI, B.A., B.Sc., L.Ag.; and N. V. KANITKAR, B.Ag.
Price, le. 1.

The Composition of the Milk of some Breeds of Indian Cows and Buffaloes
and its Variations, Part IT, by A. A. MEGGITT, B.Sc. ; and H. H. MANN,
p.se. Price, Re. 1-8.

A contribution to the knowledge of the Black Cotton Soils of India, by
W. H. HARRISON, M.Sc. ; and M. R. RAMSWAMY SIVAN, B.A. Price,
Re. 1. :

The Date Sugar Industry in Bengal. An investigation into its Chemistry
and Agriculture, by H. E. ANNETT, B.Sc., F.C.S., M.S.E.A.C., assisted
by G. K. LELE, L.ag., and BHAILAL M. AMIN, B.A. Price, Rs. 3.

Evaporation from a plain water surface, by J. WALTER LEATHER, Ph.D.,
F.I.C., F.C.S. Price, Re.1.

Studies in the Chemistry and Physiology of the Leaves of the Betel-vine
(Piper Betle) and of the Commercial Bleaching of Betel-vine Leaves,
by H. H. Mann, v.Sce.; D. L. SAHASRABUDDHE, B.Sc., L-Ag.; and
V. G. PATWARDHAN, B.Ag. Price, Re. 1-8.

The Gases of Swamp Rice Soils, by W. H. HARRISON, M.Sc., and P. A.
SUBRAMANIA AIYER, B.A. Price, Re. 1-3.

Experimental Error in Sampling Sugarcane, by J. W. LEATHER, V.D.,
Ph.p., F.1.C. (In the press.)

ENTOMOLOGICAL SERIES.

The Bombay Locust, by H. M. LEFRoy, M.A., F.E.S., F-Z-S. Price, Rs. 2-8.

The more Important Insects injurious to Indian Agriculture, by H.
M. LEFROY, M.A., F.E.S., F.Z.S. (Out of print.)

The Indian Surface Caterpillars of the Genus Agrotis, by H. M. LEFrRoy,
M.A., F.E.S., F.Z.S.; and C. C. GHOSH, B.A. Price, Re. 1-8. (Out of print.)

Individual and Seasonal Variations in Helopeltis theivora, Waterhouse,
with description of anew species of Helopeltis, by HAROLD H. Mann,
p.se. Price, Re. 1-8.

The Coccide attacking the Tea Plant in India and Ceylon, by E. E. GREEN,
F.E.S. ; and HAROLD H. MANN, D.Sc. Price, Re. 1. (Out of print.)

The Mustard Sawfly, by H. M. LEFROY, M.A., F.E.S., F.Z.S.; and C. C.
GHOSH, B.A. Price, Re. 1. (Out of print.)

The Rice Bug, by H. M. LEFROY, M.A., F.E.S., F.Z.S. Price, Re. 1.

Remarks on Indian Scale Insects (Coccid@), by E. E. GREEN, F.E.S., F-Z.S.
Price, Re. 1-8.

The Red Cotton Bug, by H. M. LEFROY, M.A., F.E.S., F.Z.S. Price, Re. 1.
(Out of print.)

The Castor Semi-Looper, by H. M. LEFROY, M.A., F.E.S., F.Z.8. Price, Rs. 2.

The Tobacco Caterpillar, by H. M. LEFROY, M.A., F.E.S., F.Z.8. Price, Re. 1-8,

The Cotton Leaf-Roller, by H. M. LEFROY, M.A., F.E.S., F.Z.S. Price, Re. 1-8.
(Out of print.)

Notes on Indian Scale Insects (Coccide), by H. MAXWELL-LEFROY, M.A.,
F.E.S., F.Z.8. Price, Re. 1-5.

Life-Histories of Indian Insects (Coleoptera 1), by H. MAXWELL-LEFROY,
M.A., F:¥.8., ¥.Z.8:. Price, Rs. 2.

Life-Histories of Indian Insects—II. Some Aquatic Rhynchota and
Coleoptera, by D. NoWROJEK, B.A., Assistant to the Imperial Ento-
mologist. Price, Re. 1-5.

Life-Histories of Indian Insects—III. The Rhinoceros Beetle (Oryctes rhi-
noceros) and the Red or Palm Weevil (Rhynchophorus ferrugineus), by
C. C. GHosH, B.A., Assistant tothe Imperial Entomologist. Price, Rs. 2.

The Food of Birds in India, by C. W. Mason, M.S.E.A.C., late Supy.
Entomologist, Imperial Dept. of Agriculture in India. Edited by
H. MAXWELL-LEFROY, M.A., F-E.S., F.Z.S., Imperial Entomologist.
Price, Rs. 7-8.

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ENTOMOLOGICAL SERIES—(contd.)

~~ - Vol. IV, No. I. Eri Silk by H. M. Lerroy, M.A,, F.E.S., .Z.8., Imperial Entomolo-
CSR ames gist ; = rth, C. GHOSH, B.A., Assistant to the [Imperial Entomologist,
j ce, Rs.

ae Vol. 1V, No. Il. Tetrigine (Acridine) in the Agri. Research Institute, Pisa, Bihar, with

descriptions of new species by J. L. Hancock, ¥.E.8. Price, Re. 1.

Vol. 1V, No. Ill. The Big Brown Cricket (Brachytrypes achatinus, Stoll), by C. OC. GHosu,

; B.A., Asst. to the Imperial Entomologist. Price, Re. 1. :
7 Vol. IV, No. 1V. Life Histories of Indian Insects (Hymenoptera), by G. R. Dorra, Si

A _ B.A., Asst. to the Imperial Entomologist. Price, Rs, 2.

BsSK Vol. IV, No. VY. Inquiry into.the Insecticidal Action of some Mineral and other Compounds
it on Caterpillars, by H. MAxwE.LL-LEFROY, M.A., F.E.8., F.Z8., Imperial
¥ ‘ Entomologist ; and R.S. FINLOW, B.Sc., F.c.8., Fibre Expert to the Govt.
x ae of Bengal. Price, Re. 1-8.

Ps . BACTERIOLOGICAL SERIES.

eee ol, 1k -No. I, Studies in the Bacteridlogical Analysis of Indian Soils, No. 1, By C. M.
= ee HUTCHINSON, B.A., Imperial Agricultural Bacteriologist. Price, Ks 2-8.

Vol. I, No. II. Rangpur Tobacco Wilt, by C. M. HuTcHINSON, B.A. Price, Rs. 2.

VETERINARY SERIES.

Ld ‘Vol. I, No. I, Anaphylaxis in the larger animals, by Major J. D, E. HOLMEs, D.8c., M.A.,
L.0.V.D., Imperial Bacteriologist, Muktesar. Price, Rs. 2.

: Yol, I, No. If. Salvarsan in the treatment of Surra in horses, dogs and rabbits, by
fs Major J. D. E. HOLMES, D&c., M.A., 10.V,D., Imperial Bacteriologist,
7 Muoktesar, Price, Re. 1-4.

Vol. 3, No. Lil. Some more successful experiments in the treatment of Surra in the camel
: with recommendations for systematic treatment, by A. S. LEgsE,
M.R.C.V.8. Price, Re. 1.

Vol. I, No, IV. On the Immune Bodies occurring in anti-rinderpest serum and on the
variations occurring in the serum proteins of animals during immuni-
zation and hyper-immunization by Dr. P. HARTLEY, D.Se. (In the press.)

i Vol,-Il, No. I, Some cases of Surra treated in the field and’in the laboratory during the
ose ee 1911, by Major J. D. E. HOLMes, M.A., D.S¢., LO.V.D.
ey rice, Re, 1.

BULLETINS ISSUED BY THE AGRICULTURAL RESEARCH
INSTITUTE, PUSA.,

3 No. 1, Notes on Cotton in Bihar in 1904, by H. M. Lerroy, M,A., F.E.8., F,Z.8., Imperial

1 Entomologist. Price, As. 4 or 6d.

Kad: No. 2. An Outbreak of Cotton Pests in the Punjab, 1905, by-H. M. Lerroy, M.a,, F.E.8,

pee , F.Z.8., Imperial Entomologist. Price, As. 4 or 6d.

The Extension of Jute Cultivation in India, by R. 8S. FINLow, B.8c., F.¢.8., Jute
Specialist to the Government of Eastern Bengal and Assam. Price, As, 12 or
1s, 2d. (Out of print.)

First Report on thesFrnit Experiments at Pusa, by A. HOWARD, M.A. -(Cantab.),
A.R.C.8. (Lond.), F.L.8., Imperial Economic Botanist. Price, As. 6 or 6d.

Report on Trials of the South African Locust Fungus in India, by E. J. Burier,
M.B., F-L.S., Imperial Mycologist ; and H. M. LEFROY, M.A., F.E.8., F.Z.8.,. Imperial
Entomologist. Price, As, 2 or 3d, ~

The Ticks Infesting Domesticated. Animals in India, by C. WARBURTON, M.A,,
Zoologist to the Royal Agricultural Society of England. Price, As. 4 or 6d. ~

A Preliminary Account of the Biting Flies of India, by H, M. LEFROY, M.A., F.E.S.,
¥.Z,8., Imperial Entomologist... Price, Re. lor 1s. 6d. (Out of print.)

Official and Recommended Methods for use in Chemical Laboratories of the Depart-
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Agricultural Chemist, Price, As. 4 or 6d. ;

Report on Cocoanut Palm Disease in Travancore, by E, J. BUTLER, ™M.B.,
F.L.8., Imperial Mycologist, Price, As. 6 or 6d. -

Treatment and Observation of Crop Pests on the Pusa Farm, by H. M. Leroy,
M.A., F.E.8,, F.Z.8., Imperial Entomologist ; and C. S, Misra, B.A. Price, As. 6

or 7d. ,
On Flax Dodder, by A. Howakpb, M.A., A.R.C.S,, F:L.8., Imperial Economic 5 Sy
Botanist.. Price, As, 4 or 6d. ) F eh
The Making and Care of Lawns in India, by A. HOWARD, M.A., A.R.C.8., F.L.S,, a,
Imperial Economic Botanist. Price, As. 4 or6d. (Out of print.)
_ Sugarcane at the Partabgarh Experiment Station, by G. CLARKE, F.1.c., Agricultura:
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Dn - ; ~ . At: —
= ~t . oi ¥

RCM

‘BULLETINS ISSUED BY THE AGRICULTURAL RESEA

ete | INSTITUTE, PUSA—(conid.) Cea
ne ee: No. 14, The Milling and Baking Qualities of Indian’ Wheats, by A. Howakp, Ma:

Rh ree A.R.C.S., F.L.S. ; and GABRIELLE 4. C, Howard, M.A, Price, As. der6d. 0
fae os ~ No. 15. Note on the Extension of Cultivation of Fibre Plantsin India. Price, As. 6 or 8d, — 3
3 feats No. 16. Second Report on the Fruit Experiments at Pisa, by A. HOWARD, M.A. (Cantab,), ens

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No. 17. The Milling and Baking Qualities of Indian Wheats, No.2, by A. Howarb, M,A.
(Cantab.), A.k.C.8.(Lona,), F-.8., Imperial Economic Botanist; and GABRIELLE ~
L. ©. HowarpD, M.A., Associate and late Fellow of Newnham College, Cambridge. —~
: . Price, As. 6 or 8d. Pe Sore Sere A ee
: No. 18. Report on the Outbreak of Blister-Blight on Tea in the Darjeeling District in
aaa 1908-09, by W. MCRAE, M.A., B.Sc. Price, Re. 1 or Is. 6d. MOS A Sees
Se NEE pe, No. 19, List of Names used in India for Common Insects, compiled in the Laboratory of the —~
ra ae Imperial Entomologist, Pisa. Price, As. 12 or ts. 2d, Kier ee
Moa ete No. 20. -Memcrandum on Indian Wheat-for the British Market, by Sir JaMEs WILSON, K.C.S.1
“EL es Price, As, 4 or 6d. 5. fo day eee
Sai Breit No. 21, Memorandum regarding Leading Eucalypts snitable for India; by F. BoorH-TucKgER, —
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No. 23.. Insecticides—Mixtures and Recipes for use against Insects in the Field, the Orchard, ©
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Entomologist. Second Kdition, Revised and Enlarged by T. B&INBRIGGE FLETCHER,
Offg. Imperial Entomologist... Price, As. 12 or Is, 2d. : Bee
The Indian Saltpetre Indastry, by J .W. LEATHER, Ph.D., F.I.C., Imperial Agri-
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Imperial Agricultural Chemist. Price, As. 8 or $d, — - : eae
Report on the Flax Experiments conducted at Dooriah in 1910-1911, by E, M, VANDE- —
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Note on the Present Position “of Cotton Investigation in India, by BERNARD ~~
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Experiments on the Cultivation of Sugarcane at the Partabgarh Experimenta
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HUSSAIN, B.A. Price, As, 5’or 6d. Apes ee as
The Cultivation of Lac in the Plains-of India. . By-C. 8. Misra, B.A., First‘Asst,.to the
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Directions for the Cultivation of Eri Silk. ~ Price, As. 3 or 4d. S Sa ee
Report on the Flax Experiments-at Dooriah during the year 1911-12, by -E.. ML’
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Botanical Series. Vol. VI, No. 6.

Memoirs of the
Department Of Aariculture

in India

SOME Sy ge DISEASES

BY

-E. J. BUTLER, M.B., F.L.S
Imperial Mycologist

AND

ABDUL HAFIZ KHAN

Assistant to the Imperial Mycologist

takes) Si be er ee erates
gE RE NS Oa SETS. tle

MY

ae

‘AGRICULTURAL RESEARCH INSTITUTE, PUSA
PUBLISHED FOR

Ae: IMPERIAL DEPARTMENT OF AGRICULTURE IN- INDIA

BY
THACKER, SPINK & CO,, CALCUTTA

3 ce SW, THACKER & CO., 2, Crrzp Layz, LONDON

i Price, Rs. 2. 2

December, 1918. BoranicaL Serigs VoL. VI, Ne. 6

MEMOIRS OF THE
DEPARTMENT OF AGRICULTURE
IN INDIA

SOME NEW SUGARCANE DISEASES
ae
Ed, BUTLER, WB: F.L.S
Imperial Mycologist

AND

ABDUL HAFIZ KHAN

Assistant to the Imperial Mycologist

AGRICULTURAL RESEARGH INSTITUTE, PUSA
PUBLISHED FOR
THE IMPERIAL DEPARTMENT OF AGRICULTURE IN INDIA
BY
THACKER, SPINK & CO., CALCUTTA
W. THACKER & CO., 2, Creep Lane, LONDON

CALCUTTA

PRINTED BY THACKER, SPINK AND oO.

PEATE I:

Fig, 3.

Fig. J,

SOME NEW suG ARCANE

BY

J. BUIXURR, MBE

fingsriak Uyeclogsst,
SZ

ARUDBS TAGS RUAN
“DESCRIPTION OF PLATE x
he

Appearance of split eane\Vinfected with Cephalosporium Sacchart.

: _Natural size. /
{Cex Viades Ty RNY NaS W, G7 <7}; yi, .
_ Appearance of w EN cane infected with Hendersonina Sacchari. xf. |

dj ink aS uiRne Cases ieeiotherne tm, Sacchari, Nevural size...
eaused by Colletotric AWN: folectum NOR A. SHO V0. Gr,
the name red rot might equally well be - \Y~

duces a eeatia es op of ~ Cane —

& oe ROAR,

: atch cases of severe erates of ee Otay, Weetiod Foy ©
i planting as cause the death of the young PG ts, 7 De Wbwinei
> more fully referred to below, little is noticed until the ay &
about half grown. At this period affected canes lag i meow wad
nee single stools, or patches of varying size, may soon be
observed scattered through the fields in which the disease ic
re orevalent. From this on until the time of harvest, witheving of
“individual canes, Pls of whole stools, occurs. The leaves dry
wp, as supplied with water, followed by the sters,
iuare if the cane be _ longa n-

A Chavae-

ol.

Bes

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SSE : S f SS Shey re m4

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SOME NEW SUGARCANE DISEASES

BY
BK. J. BUTLER, M.B., F.1.S.
Imperial Mycologist,
AND
ABDUL HAFIZ KHAN

Assistant to the Imperial Mycologist.

b= Wir

(Cephalosporium Sacchari Butl. n. sp.)

In the course of investigations of the red rot of sugareane,
caused by Colletotrichum falcatum Went, a second disease, to which
the name red rot might equally well be applied, since it pro-
duces distinct reddening of the cane pith, has frequently been
encountered. With a little practice the two diseases can be dis-
tinguished and they are caused by entirely distinct organisms,

Asin red rot, the earlier symptoms are elusive. Except
in such cases of severe infection of the cuttings selected for
planting as cause the death of the young shoots, in the manner
more fully referred to below, little is noticed until the cane is
about half grown. At this period affected canes lag in growth and
stunted, single stools, or patches of varying size, may soon be
observed scattered through the fields in which the disease is
prevalent. From this on until the time of harvest, withering of
individual canes, or even of whole stools, occurs. The leaves dry
up, as if insufficiently supplied with water, followed by the stems,
which become light and hollow. If the cane be split longitudin-
ally when the leaves are first observed to wither, a charac-

182 NEW SUGARCANE DISEASES

teristic discoloration of the pith, as shownin PI. I, Fig. 1, may
be observed, A comparison of this figure with a coloured drawing
of true red rot®* will show the difference in appearance of the two
diseases. Instead of bright red patches and streaks, broken by
transversely expanded white areas, there is a diffuse purple or
dirty-red colouration, in which brighter red, vertical lines mark
the position of the bundles. The tendeney of the colour to
become muddy at an early period is its most strongly marked
character and serves to distinguish the disease from any other
known to us. In old cases the red almost disappears, being
replaced by an earthy brown. The pith dries up more rapidly
than when attacked by Colletotrichum and becomes hollow,
Within the hollow portion, a fluffy grey growth of mould is often
found.

Microscopic examination shows that the stem is infested
throughout the reddened portion, which may be confined to a few
joints or extend to the whole length of the cane, by a fungus,
whose hyphe ramify through the cells in all directions, penetrat-
ing the fibro-vascular bundles as freely as the large-celled paren-
chyma of the pith (PI. II, Figs. 2and 3). Most of the hyphx
are fine, not exceeding 3 » in diameter in recently infected parts,
and either uniform or tapering gradually. They pass from cell to
cell through the pits in the walls, when pits are present (Fig. 2),
but can also penetrate unbroken walls and seem to have no pre-
ference for one tissue more than another. Even when old, they
remain hyaline. Occasionally they have been found bearing conidia
in the vessels (Fig. 5) and possibly do so also inthe parenchyma
of the pith, as the condition shown in Fig. 4 appears to repre-
sent an early stage of spore-formation. The conidia are small,
hyaline, unseptate, oval, and usually appear to be borne singly
on short lateral branches of the mycelium. They resemble
closely the microconidia frequently found in the vessels of plants
affected with the Fusarium wilts,

* See Memoirs, Dept. of Agric, in India, I, No, 3, 1906, Pl. I, or Agric, Journ, of India, IT,
1907, PLeA,

BUTLER AND HAFIZ 183

In the hollow which invariably forms in the pith of affected
internodes at a late stage in the disease, there is usually a
copious greyish-white, fluffy growth of hyphe, similar to those
found in the tissues, and bearing great numbers of conidia,
resembling those observed in the vessels, though often of some-
what larger size. The characters of the fungus can be studied
from this growth or, more satisfactorily, from pure cultures, and
they show it to be a member of the form-genus Cephalosporium,
several species of which are known to be lower stages in the
development of /7ypocreacee.

The hyphe are hyaline, slender (about 3 to 5 « in diameter),
richly branched, sparingly septate when young, not varying
abruptly in diameter, though swollen segments are found in old
cultures. Sometimes the individual filaments unite together to
form coremial strands, usually only two or three hyphe taking
part in the formation. An early stage of such a strand is shown
in Fig. 6a. Chlamydospores have not been observed either in
culture or in diseased canes.

The conidia are borne on short, simple or branched, lateral
hyphe and also terminally on the ultimate branches of the
mycelium. They measure 4 to 12 (usually 5 to 8 u) by 2 to
3 4, when formed, but increase in size prior to germination. Their
shape varies from shortly oval to ovoid or long elliptical. Occa-
sionally they are curved or with one side flattened. They are
almost invariably unicellular when formed, but some become
septate prior to germination, the septa being 1 to 3 m number
(Fig. 11). Spores and mycelium are hyaline.

The conidiophores are usually of definite shape, measuring
from 6 to 30% by 3 to 4m at the thickest part, swollen slightly
in the middle or lower third, narrowing below where they arise
from the mycelium and tapering above, but with an obtuse apex
on which the spores are borne (Fig. 7). They may be scattered
along the hypha or arise in bunches near together (Fig. 6).
Branching is not uncommon, being irregular or cnee whorled or
forked. Each branch usually bears conidia at the apex, but if
kept very moist the branches may grow out into long hyphe.

184 NEW SUGARCANE DISEASES

The first conidium is borne at the tip of the conidiophore and,
when full grown, is pushed to one side by the formation of a
second spore at the same point, and so on until a number have
been produced. Sometimes, instead of forming a second spore,
the apex continues growth as a hypha, which may again bear
conidiophores (Fig. 13a). Unlike most members of the genus,
the successively formed conidia are not held together in a mucilag-
inous drop, but adhere merely by surface attraction and are very
easily dislodged. If kept free from currents of air in a moist
chamber, heads of 5 or more can readily be found. They adhere
more firmly at the base than elsewhere and if carefully mounted,
after treatment with acetic acid, such appearances as those figured
in Fig. 7a can be observed. At first sight this suggests that the
spores arise near together and at the same time, as in the genus
Haplotrichum, but more careful examination shows that this is
not the case, only one spore being formed at a time and the whole
apex of the conidiophore taking part in its formation. The apex
is pushed out by growth from within the hypha to form the
spore, and if examined just after the latter has fallen and before a
second appears, the condition shown in Fig. 9 is found. In moist-
chamber cultures spores formed on the aerial mycelium appear
as shown in Fig. 8, lying in asingle layer with their long axes
parallel. Spore-production is extraordinarily copious, almost
every hypha being abundantly provided with conidiophores
throughout the greater part of its length.

Germination occurs within 24 hours in many cases, the large
septate and small unseptate conidia behaving in much the same
manner (Fig. 11). The germ-tube is usually single and_ ter-
minal, but the septate spores may give out a tube from each end.
Union of two or more germ-tubes, belonging to different spores,
is common (Fig. 12), up to five spores having been found united
in this manner. One or two of the germ-tubes grow out into
strong hyphz from the united mass, the arrangement being prob-
ably merely a nutritional adaptation.

In culture on nutrient agar the mycelium remains white and
filamentous and no formation of stromata or of other types of

BUTLER AND HAFIZ 185

spores has been observed. The species does not agree with any
previously described and has been named Cephalosporium Sac-
char, the diagnosis being as follows :—

Cephalosporium Sacchar: Butl. n. sp. Effusum, candidum ;
hyphis repentibus, parce septatis, 3-5 « diam, ; conidiophoris
continuis, simplicibus furcatis vel subverticillato ramosis, sursum
obtusis, medio vel versus basim incrassatis, 6-30 u long. 3-4 » crass. :
conidiis ex apice ramulorum pluribus continuis exsilientibus et in
capitula collectis sed facillime secedentibus, hyalinis, ovoideis vel
oblongo-ellipsoideis, continuis, 4-12 W 2-3 u.

In culmis Sacchari officinarum in India or

Cultures of the fungus can be readily obtained in the same
way as with Colletotrichum falcatum, by cutting out aseptically
a portion of the pith, in the early stage of the disease, and incubat-
ing in a sterile Petri dish or potato tube. The growth is frequent-
ly pure from the commencement, but if not, can be readily purified
by sub-culturing from the aerial mycelium. The following in-
oculations were carried out with pure cultures from nutrient agar,
in which spore-production was copious.

On December 13th, 1908, 20 strong sound Red Mauritius
canes were inoculated ina stem wound with a suspension of spores
and mycelium in distilled water, from a culture two weeks old.
The inoculations were made in the lower half of the cane by
removing a cylinder of pith with a small sterile cork borer, insert-
ing a drop of the suspension, and replacing the cylinder after
cutting off a little of the end, so as to leave a small cavity. The
wound was then bound with sterile gutta-percha sheeting.

The canes were examined after 3 months. In one case only
had the whole cane dried up, the infection extending above the
wound to the top of the cane, and below for 3 internodes. In the
others infection had progressed through from 4 to 6 internodes.
The inoculated internode had become hollow in every case, those
above and below being also hollow in some cases. The symptoms
were typical and the fungus was found throughout the discoloured
parts,

186 NEW SUGARCANE. DISEASES

On the same day 15 canes were inoculated at leaf scars, the
sear being covered after inoculation with a wad of damp sterile
cotton-wool, bound with gutta-percha sheeting. The same
material was used as in the last experiment.

These inoculations also succeeded, distinct reddened bundles
being found after 3 months, extending downward from the scar
through the next internode. The progress of the infection was
less than in the last case and could not be traced beyond the
one internode. The hyphe were found chiefly in the red-
dened bundles and there was not much lateral spread. It was
noticed that shoots arising from the inoculated nodes were not
infected.

The fungus cannot penetrate the unbroken rind of the
internodes. Sixteen inoculations were made to test this, a drop
of the suspension being placed on the rind, and covered with
damp sterile cotton-wool for 48 hours. In no case was there
the slightest sign of infection.

As in Colletotrichum falcatum, however, the adventitious
root eyes at the nodes can be inoculated successfully. Ten
inoculations were made on the uninjured eyes, in exactly the
same way as in the last experiment, except that the cotton-wool
was removed after 36 hours. The first sign of success was
noticed after 4 days; one of the eyes being slightly discoloured.
On sectioning, the hyphe were found to have penetrated through
the epidermis and to have reached almost to the centre of the
bud. Three days later all the inoculations had clearly succeeded,
the eyes being reddened and full of hyphe. After 10 or 11 days
the eyes were quite killed and the hyphx were passing into the
main stem.

To see if the cane can be infected through the setts at the
time of planting, half a trench of Striped Mauritius was planted
on March 7th, 1907, after dipping the cut setts in a suspension
of spores and mycelium in distilled water. The remaining half
trench served as control. Germination was approximately equal in
the two halves, but 22 shoots withered in the inoculated half up
to June 17th, whereas there were only a few deaths in the other

BUTLER AND HAFIZ 187

half, apparently due to white-ants. The Cephalosporium was
recovered from 3 of the withered shoots examined.

As this fungus has been several times found associated with
Coiletotrichum falcatum, experiments were made similar to the
last, but with mixed suspensions of the two fungi, in 1907. The
results were striking. Germination was good in both halves of the
trench, but 45 shoots withered in the inoculated half in the first
three months, while there were very few failures in the rest of
the trench. Colletotiichum was recovered from 2 of the withered
shoots and Cephalosporium from 8, out of 14 examined. The
experiment was repeated in 1908 with much the same result.

These experiments show that the fungus can enter the cane
through wounds, through the uninjured root eyes at the nodes,
and through the planted setts. In true red rot the writers have
shown* that the latter is the most frequent method of infection
in Northern India. In the present case the evidence is not so
complete, but it has been found that wound infection is far more
common than with Colletotrichum. The wounds examined have
been all borer holes, which are much the most frequent wounds
in standing cane. Altogether 27 have been examined by incu-
bating an aseptically removed slab from just beyond the margin
of the hole and, of these, 15 have given the fungus. Out of 9
examined in the 1908 crop when nearly fully grown, 8 were
infected. This was the year when the most widespread attack
of Cephalosporium Sacchari was observed at Pusa.

The disease is found over a large part of India. It has been
observed at Surat, Poona, Samalkota and throughout North
Kastern India. Asa rule it seems to accompany red rot, quite
a considerable percentage of the cases examined being due to
mixed infection. Inthe 1908 attack at Pusa, however, there
was practically no true red rot and a good deal of damage was
done by an unmixed attack of Cephalosporiuwm. Similarly, in
1912, several cases of pure infection were encountered, and pure
cultures were obtained direct from the mycelial growth in the

* Butler, E. J., & A. Hafiz Khan, Red Rot of Sugarcane, Mem. Dept. of Agric. in India,
Bot, Ser., VI, No. 5, 1913.

188 NEW SUGARCANE DISEASES

hollows of the pith. Asa rule, so far as has been observed, the
infection is not virulent ; spread within the cane is gradual and
communication from one plant to another slow. A large number
of borer holes become infected late in the season, but the parasite
usually seems to remain confined to a few internodes. As bored
setts are usually discarded at planting time, infection by this
channel is not often transmitted to the following crop.. The
fungus has not been found on the leaves or on the surface of the
stem, in the field. In the laboratory, however, a case of sponta-
neous infection of the leaf with a fungus exactly resembling that
under consideration occurred, and this led to further examination.
Inoculations were made with cultures obtained from the growth
on the leaf and also with cultures from diseased cane stems.
Both gave similar results. Six inoculations were made with each
of the two cultures. In two cases the inoculated leaf withered
too rapidly to allow of definite observations. The remaining ten
were quite successful, Pl. LI, Fig. 1, showing the condition after
three days. Penetration had occurred chiefly through the large
thin-walled motor cells.: We have not been successful in finding
cases of leaf infection in the field, but as leaf spots are extremely
common on sugarcane such a search is not easy. The spot caused
by Cephalosporium Sacchait in artificial inoculations resembles
that caused by IZelminthosporium Sacchari, described in a later
section of this paper, and all the field cases examined belonged to
this latter fungus.

Rotten canes must set free large quantities of spores and the
source of the infection of cane wounds may come from these, from
the leaves and, if the species passes part of its life as a soil sapro-
phyte, as is not improbable, possibly also from the soil. The genus
is so common in cultivated soils, and this species has so little to
distinguish it from some of the habitual soil dwellers, that its
recognition would be difticult. It lives very readily as a sapro-
phyte, however, growing luxuriantly on most of the common
culture media and remaining alive in culture for about 3 months,
so that it is probably present in the soil of cane fields. The
inoculation experiments indicate that infection of the setts at the

PLATE Il.

—S
Bh)
Kom

DO

CEPHALOSPORIUM SACCHARI.

BUTLER AND HAFIZ 189

time of planting, causes a good many of the young shoots to
wither within the first 3 months. As this early withering
has not been very frequently observed in natural attacks, we
conclude that infection of the growing cane, through wounds and
through the root buds at the nodes, is the more common origin
of the disease.

The control of the disease should evidently be on much the
same lines as in ture red rot*. As, however, wound infection
is far more common, the importance of removing diseased clumps
before they have time to rot and set free spores is much greater.
As a rule the disease is not a severe one and though our
experience with it is limited as yet, it is probable that it is
incapable of doing permanent damage so long as the measures
advocated against red rot, which we consider to be essential
to the successful growing of thick cane in Northern India,

are carried out.

Descrivrion or Prats II.
(Cephalosporium Saccharv Butl.)

Fig. 1. Leaf of sugarcane inoculated with Cephalosporium Sacchari, showing
fungus in the motor cells and emerging to the surface. x 640,

Ww

Hyphz in parenchyma of stem and passing from cell to cell through the
pits in the walls. x 190.
,, 9 Hyphe in fibro-vascular bundles. x 190.
» +. Hyphaina cell of the parenchyma with indications of spore formation,
x 320.
» 9 Spore formation on hyphe in a vessel of the stem. x 320,
6. Mycelium bearing conidia, from a pure culture on nutrient agar. At wu the
commencement of the formation of a coremial strand is seen. x 320,
,, 7% Conidiophores from a pure culture, after treatment with acetic acid. At «
the adherence of the conidia at their bases is indicated, x 640.
8. Successive stages in conidial formation, drawn from a continuous observa-
tion, x 640,
» 9. Conidiophore immediately after a spore has fallen and before a new one has
commenced to appear, showing the hollowed apex. x 640.
,, 10. Conidia from nutrient agar culture, 6 days old, x 640,

* Butler, E. J., & A, Hafiz Khan, Joc, cit.

Fig. 11
Fem 5
oo ale

NEW SUGARCANE DISEASES

Germination of the conidia, showing especially the septation which some-
times precedes germination. x 320.

Union of the germ-tubes of the conidia. x 320.

A spore-bearing hypha continuously observed, showing the extent of growth
and spore formation in 22 hours. At @ conidiophores have continued

apical growth after forming a single conidium, ‘The septa have been

omitted.

II.—Cottar Ror.
(Hendersonina Sacchari, Butl. n. g., n. sp.)

In the last section a disease was described which has fre-
quently been mistaken for red rot. A second disease is sometimes
confused with red rot in India, and the following description may
lead to its recognition and thus secure further information regard-
ing its distribution and the amount of injury it causes the crop,
points on which our observatious are incomplete.

Towards the end of 1908, a number of canes withered at the
Samalkota Sugarcane Station, following on a cyclone at the end of
September, which did much harm to the crop. A recrudescence
of red rot was feared, as many of the canes were reddened at the
base. We examined the crop in January, 1909, and found ex-
tremely little true red rot. Some of the damage was apparently
not caused by disease but by injury to the root system asa result
of the storm. A good many cases were seen, however, in which
the symptoms suggested the action of a definite parasite at the
base of the plant, and a fungus was isolated from the tissues in
these cases which proved capable of reproducing the disease.

In the following season the disease reappeared and was more
fully studied, but for the last three years there have been no
further records of its occurrence on this farm. Recently it has
been found at Jorhat Farm in Assam, and it is possible that it
also exists in the Central Provinces, where we have observed a
somewhat similar condition, but without being able to make a
detailed examination. It has not been recorded elsewhere, and
the parasite differs so widely from any previously known that it
has been necessary to make it the type of a new genus. The
varieties of cane so far observed to have been attacked are Red
Mauritius, Striped Mauritius and B. 208.

192 NEW SUGARCANE DISEASES

The symptoms outwardly resemble those of red rot, so far
as the withering of the top is concerned. The top leaves wither
back from the tip along the edges, the midrib remaining green
later than the rest of the leaf. The larger leaves below the crown
appear to suffer first, those at the apex remaining unaffected for
some time. When the leaves have fully withered, the cane is
found to be much lighter than normal. On splitting, the upper
part is usually pithy and dry in the centre, or even with a central
cavity along each internode, around which the pith is dry, white
and flaky. Lower down the pith may be still juicy but has a
curious translucent watery appearance ; still lower the centre por-
tion may be brown, while red streaks or patches may often be
seen, especially at the nodes (Pl. I, Fig. 2). At the base, where
the feeding roots arise, the red colour predominates and is
especially visible at the nodes. In old cases the lower internodes
also dry up and may develop a central cavity, surrounded by red or
brown pith. The roots arising from the basal nodes are usually
blackened and rotten, and the appearance suggests that the
disease enters the base of the stem from the roots.

In the Jorhat cases the buds at the lower nodes had sprouted
and subsequently withered. The reddening was specially well
developed at the nodes from which these shoots arose. It is not
yet certain how far this sprouting of lateral buds is a symptom of
the disease, as it appears possible that the Jorhat attack is com-
plicated by the occurrence of a non-parasitic, seve/-like degenera-
tion of the cane, which requires further investigation.

Microscopic examination shows that the roots and base of the
stem are invaded by a fungus. The mycelium is confined to the
definitely reddened parts, being absent from the translucent upper
portion of the cane. In the Jorhat cases the fungus was found
collected chiefly at the nodes, the intervening internodes being
_ almost free from hyphie until a late stage. The discoloured roots
always contain a considerable quantity of mycelium. In the
early stages the hyphe run between the cells in the inter-
cellular spaces. Here they are usually extremely fine, though
thick ones sometimes occur. Later on, branches from the inter-

BUTLER AND HAFIZ,. 193

cellular mycelium penetrate the walls to enter the cells. All the
tissues are invaded, bundles as well as parenchyma (PI. IV, Fig. 1).
In the cells, and especially in the vessels, very thick hyphe may
occur, sometimes almost alone, sometimes intermingled with the
fine filaments. Careful observation shows that both kinds belong
to the same mycelium and can be found in direct continuation
with one another. Sometimesa haustorium-like, branched mass,
arising from thick hyphee in an intercellular space, almost fills a
cell. At first septation is rather scanty, but in the older hyphex
the septa lie very close together, the segments thus formed being
often broader than they are long. No trace of spore-formation
was found either in the tissues or on the surface of the diseased
eanes before their death.

Cultures of the fungus were obtained in the usual way,
from the surface of the cane pith cut with a red-hot knife. The
fungus grew well on ordinary nutrient agar, forming a dense
superficial growth. The following description is from pure
cultures on this medium.

The mycelium is white or faintly tinged yellow. The
hyphe (PI. IV, Figs. 2 & 3) are very variable in size. The main
branches are very thick, sometimes up to 15 in diameter, at
first hyaline and sparingly septate, then pale yellowish and
closely septate. Branching is copious and often rectangular.
Very noticeable is the tendency of the older branches to give off
extremely fine, hyaline, thin-walled hyphe, the difference in
diameter being so great as to suggest a distinct mycelium, until
carefully examined. The thin hyphe are often irregularly
swollen or even nodular and measure sometimes as little as 1 » in
diameter. Anastomosis of neighbouring branches is not uncom.
mon (PI. IV, Fig. 3a). Intermediate stages, consisting of hyphz
6 to 8 in diameter, freely septate when old and often irregularly
swollen, are common. The thicker branches break up readily
into chlamydospores (PI. TV, Figs. 4 & 5), which may be termi-
nal or interealar, and are usually arranged in short chains.
They consist of thick-walled cells filled with reserve material,
round, oblong or long elliptical in shape, variable in size, the

194 NEW SUGARCANE DISEASES

largest up to 33 u in diameter, at first hyaline but later slightly
coloured. They frequently separate from the mycelium when
mature, and may be found lying singly or in short chains inter-
mingled with the hyphe. ‘They germinate in water by an
outgrowth of the endospore. The nodular swellings on the
smaller hyphze seem to be of the same category, but do not
usually become detached ; they often give off fine branches after
reaching maturity. In many cultures no other spore form was
found. In several, however, a pycnidial stage developed, and this
stage was subsequently found on the surface of old canes which
had been killed by the disease.

When pycnidia were formed, they began to appear in about
3 weeks after sowing the tubes, as fine blackish dots immersed in
the white mycelium. These grew to form prominent black
nodules, which, under the microscope, were found to consist of
stromatic tissue with one or more sporiferous loculi entirely
immersed in the stroma.

The stromata are leathery, roundish-conical, about 1 to
2 mm. in diameter, and consist of an outer portion of brown,
fibrous tissue, composed of closely woven hyphe, and an inner,
bounding the loculi, of dark brown pseudo-parenchyma, many
layers deep. Sometimes bands of the fibrous tissue separate the
loculi, each cavity then having its separate investment of pseudo-
parenchyma (PI. III, Fig. 1), in others the whole centre portion
of the stroma is cellular and the cavities have no distinct walls,
except the lighter coloured layers from which the spore stalks
arise. The cells of the pseudo-parenchyma are distinct, angular,
isodiametric in the deeper layers, but becoming more elongated as
the hymenium is approached, with unthickened but deeply pig-
mented walls and oily contents, and measuring up to 10m in
diameter. The colour gradually becomes lighter towards the
spore-bearing surface.

The pyenidial loculi are deeply sunken in the stroma, very
irregular in shape, and often communicate with one another by
narrow passages. Every stage may be found between stromata
with a single loculus, through such cases as shown in PI. III,

Prare Ill.

Hendersonina Sacchari.

Photo.-Engraved & printed at the Offices of the Survey of India, Calcutta, 1913.

BUTLER AND HAFIZ 195

Fig. 1, which we take to represent the union of several unilocular
stromata, to truly multilocular pseudo-parenchymatous masses.
The cavity is usually narrow, due to cushion-like projections of the
wall into the lumen. In small unilocular stromata, the cushion
arises usually at the base, the cavity being convex outwards.
In larger stromata cushions may also project from the sides, lead-
ing to very irregular formations. In transverse section through
the middle of the stroma (Pl. III, Fig. 3), the cavities are quite
irregular in position, but near the apex the arrangement may be
roughly circular, due probably to several loculi converging to a
common opening. Luaterally situated loculi may, however, open
by separate orifices. In longitudinal section (Pl. III, Fig. 2),
there is no uniformity of disposition or size of the cavities. The
mouth is formed late and is usually not prominent.

The spores are of two kinds, both borne on exactly similar
sporophores. The latter arise from the innermost layer of the
pseudo-parenchyma, which is not sharply marked off as a
hymenium but consists simply of small, rather elongated cells,
light yellow in colour (Pl. 1V, Fig. 6). From these one or more
hyaline sporophores arise and project into the cavity. Usually
the sporophore is branched, the branches arising chiefly near the
base and each terminating ina single spore (Pl. IV, Figs. 7 & 8).
In some stromata unbranched sporophores predominate (PI. IV,
Fig. 9). Septa occur sparingly and the ultimate branches are
always slender and unseptate. Hach sporophore appears always
to bear only one kind of spore, but as they are closely crowded
at the base, and both spore forms are found side by side in the
same cavity, it is difficult to be certain on this point. Both
kinds of spores appear to arise simultaneously,

The most highly differentiated spore form (PI. V, Fig. 1),
consists of brown, elongated cells, rounded at the ends,
unicellular or with one or two transverse septa, usually straight
but sometimes curved, the curvature in bicellular spores being
occasionally sigmoid. They measure 15-24 \/ 3°75 to 5m. In
some cultures unicellular spores predominated, in others bicellular.
Three-celled spores are the least numerous. Germination occurs

196 NEW SUGARCANE DISEASES

in less than 24 hours by an outgrowth from one or both ends, the
germ-tube remaining unseptate until it has reached a consider-
able length (PI. V, Figs. 4 & 5).

The other spore form (Pl. V, Fig. 2), consists of hyaline
filamentous cells, usually without septa but with many oil drops,
straight or irregularly curved, variable in length and breadth, the
longest being often very narrow, sometimes broader at the base
and tapering to a narrow apex, sometimes quite uniform in
diameter. They measure 20 to 60 by ‘6 to 24. Germination
was not observed.

An intermediate form of spore (Pl. V, Fig. 3) is sometimes
found, consisting of pale yellow, elongated, 1-septate cells, borne
on sporophores of the type above described and measuring 18 to
30 by 2 to 3°75 mu.

After the pyenidial stage had been obtained in pure cultures,
it was found in two cases on old canes attacked by the disease,
one being an inoculated cane at Pusa, the other a diseased cane
from Jorhat. The stromata develop in the internodes, under the
epidermis, or one or two layers deeper in, and are smaller than
those obtained in culture. The outer fibrous layer is absent or
reduced, the mass of the stroma being parenchymatous. The
base is broad and extends for some distance all round, as a narrow
stromatic layer between the tissue cells. In the centre, the
epidermis is raised up and ruptured by the roughly conical, deep
portion of the fruit body, which is hollowed out into one or
several cavities. Unilocular stromata are common, the cavity
being irregular or sometimes imperfectly divided by incomplete
walls. The mouth is usually single and develops late. Both
spore forms occur in the loculi.

There is much greater uniformity in the spore characters in
any one culture of the fungus, than would appear from the above
description, which is based on the examination of a large series.
Thus, in some cases, the first spore form was almost entirely
unseptate and small and the second broad and distinct ; in others
bicellular spores of the first type were chiefly found, intermingled
with which were a few hyaline, attennated spores of the second

BUTLER AND HAFIZ 197

type ; in others again the narrowly filiform second type predomi.
nated, while there. were numerous “ intermediate” spores and
only a few typical bicellular brown spores ; finally in some cases,
especially in the earlier cultures, all the types above described,
except the “intermediate,” were found, though 3-celled spores
were never common.

Hence we have had some difficulty in deciding whether the
fungus should be considered to belong to the sections Phwodidyme
or Pheophragnue of the Spherioidacee. But the characters as a
whole agree better with those of the latter section, of which
Hendersonia is the main type, than with the former, which
consists mainly of Diplodia and its allies, and the 3-celled brown
spores should, we think, be considered as the most highly differ-
entiated and, therefore, most important for systematic purposes,
The genus differs from the rest of the Pheophragmie chiefly
in the characters of the stroma and the possession of two distinct
types of spore in the same loculus. The latter peculiarity
approaches it to the genus Phomopsis amongst the Hyalospora,
and it is of great interest as supporting the view that the
hyaline, filamentous bodies of the latter genus are true spores,
and not merely disjuncted basidia as has been maintained by
some observers. There cannot be the smallest doubt that these
bodies in the present fungus are spores, for they are borne on
true sporophores, in an exactly similar fashion to the spores of
the first type. Another fungus in which two spore forms are
found in the same pyenidium is Fusicoccum viticolum Red*. -
which Shear} has shown to be the imperfect form of Cryptos-
porella viticola Shear. Shear distinguishes the two spore forms
as pycnospores and scolecospores, and believes the latter to be
true spores, not basidia as held by Reddick. The pyenidia
themselves are Immersed in a stroma, which is evidently of much
the same type as that of the sugarcane fungus.{ It should be

* Reddick, D. Necrosis of the grape vine, Cornell Univ. Agr. Expt. Stat. Bull, 263, 1909.
t Shear, C. L. The ascogenous form of the fungus causing dead-arm of the grape. Phyto-

pathology, I, 1911, p. 116.
¢ Gregory, C.J. A rot of grapes caused by Cryptosporella viticola, Phytopathology,

ILI, 1913, p. 20, fig. 1.

198 NEW SUGARCANE DISEASES

noted that several species described as Phomopsis are multilocular
stromatic*, though Diedicke, who has most fully studied the
genus, places it amongst the simple forms, and does not seem to
include any species with a multilocular stroma. Another genus
which may be compared is Hndothiella Sace., where the bodies
described as filiform basidia or pseudo-paraphyses are proRaly
scolecospores.

The following is the diagnosis :—

Hendersonina Butl. nov. gen. Spheropsidacearum—Stro-
mata innato-erumpentia, atra, coriacea, parenchymatice contexta.
Pyenidia (loculi) immersa, inzequalia, ostiolis spe confluenti-
bus. Basidia ramosa. Sporule in basidiis acrogenx, alters
fuliginese, ellipticee vel elongate, recte vel curvule, continue
vel 1-2 septate, alteree hyaline, filiformes, recte vel flexuose,
continue,

Hendersonina Sacchari Butl. n. sp. Stromatibus cortice
innatis demum erumpentibus, subgloboso-conicis, 1-2 mm. diam.,
atris, intus 1-pluri-locularibus ; loculis irregularibus, subinde
incompletis vel inter se communicantibus, ostiolis seepe confluen-
tibus; contextu brunneo, minute parenchymatico ; basidiis ramoso-
fasciculatis, hyalinis ; sporulis dimorphis, aliis fuligineis, rectis vel
curvulis, ellipsoideis vel elongatis, utrinque obtusis, continuis vel
1-2 septatis, 15-24 YW 3°75-5 w, aliis hyalinis, filiformis, rectis vel
flexuosis, pluriguttulatis, 20-60 YW °6-2 u

In culmis Sacchari officonarum in India, or

As this fungus is possibly the lower stage of a Pyrenomycete,
attempts were made to obtain an ascigerous stage in culture on
organic media. The following were tried :—-Potato, carrot, onion,
plantain, Colocasia corm, fruit of Carica Papaya, fruit of
Psidium Guajava, boiled rice, bread paste and sugarcane pith.
The last medium was the only one, besides nutrient agar, in
which pyenidial stromata were developed and none gave rise to

* c, f. Harter, L. L. & Ethel C. Field. Diaporthe, the ascogenous form of sweet potato
dry rot. Phytopathology, 11,1912, p, 121, Roberts, J. W- A new fungus onthe apple. 7h.,
p. 263,

BUTLER AND HAFIZ 199

an ascigerous stage. After a week the onion cultures were the
best, followed closely by carrot, potato, Carica, Colocasia, Psidium
and rice. On bread also the growth was good, while plantain
and sugarcane were much inferior, especially the last. The
mycelium was loose and had begun to lose its original white
colour in many of the tubes, the upper part of the aerial
mycelium (which exceeded 2 inches in height in some of the
cultures) turning grey, except in the plantain, rice and sugarcane
tubes, while the growth near the surface of the medium remained
white in all. A submerged growth had developed in the water
in the bottom of the (potato) tubes and this became light purple
or pinkish in the carrot, onion and Carica tubes. The medium
was discoloured dirty grey in the onion tubes, eye-grey to purple
in those of rice and slaty-blue in those of bread. Five days
later a blackish crust had developed on the surface of the water
in the potato, carrot, onion, Carica and Psidium tubes, the
submerged mycelium was light cinnamon colour in the onion and
Psidium tubes, brick red in the plantain, and unchanged in the
others. The colour had diffused into the water. The aerial
growth was turning brown on the surface in the carrot, onion,
plantain, Colocasia, Carica, Psidium and bread tubes, with a
little pink in places in those of Colocasia. The medium was
blackened in the potato, carrot and Carica tubes, while the upper
part of the rice was dark coloured and a bluish line separated
this from the unaltered lower portion; the same bluish colour
developed in the upper part of the bread paste. Seventeen days
later there was little change, the growth in the potato and onion
tubes was darker, that in the carrot, plantain, Carica and
Psidium tubes had become pinkish in places, the submerged
mycelium was chocolate-brown in the plantain, and chestnut in
one of the sugarcane tubes, and a few dark dots were noticed in
the denser parts of the mycelium of several tubes. These did not
develop further and were ultimately found to be merely
condensations of sterile mycelium.

The following inoculations were carried out with pure
cultures, obtained as above described,

200 NEW SUGARCANE DISEASES

In October, 1909, 35 canes in a plot of strong healthy Red
Mauritius, about 7 months old, were inoculated at Samalkota by
removing a cylinder, about half an inch in length, with a small,
flamed cork borer and inserting a small tuft of mycelium. The
cylinder was replaced after cutting a little off the end, and the
wounded stem bound with sterile gutta-percha sheeting. Seven
weeks later, two of the inoculated canes commenced to wither.
In January, 1910, another inoculated cane withered and was sent
to Pusa for examination. The condition of the stem was similar
to that in the canes from which the fungus was first obtained.
The inoculated internode was bright red. The reddening extended
upwards for 3 internodes, the 8rd having only one bright red
bundle. Higher up, the translucent, watery condition of the
pith, already described, was found. Downwards, the reddening
extended for 2 internodes. Hyphze were numerous in the redden-
ed parts and pure cultures of the fungus used in inoculating were
obtained readily. In February, four more of the withering inocu-
lated canes were received, as well as two non-inoculated canes
arising from the same clumps, which seemed to have become infee-
ted secondarily. The inoculated canes had typical symptoms. In
two, the infection had extended downwards to the base of the stool,
the upper part being less affected. In the other two, distinct red-
dening occurred for 2 or 3 internodes above the wound and the
pith was translucent and hollow in the centre for some 10 inter-
nodes higher. The hyphe were, as usual, confined to the reddened
portion, where they were plentiful and easily obtained in pure
culture, giving rise to the same fungus used in inoculating, The
two non-inoculated canes had become infected from below ground,
apparently through the roots, and were withering asa result of the
attack. The fungus was recovered from these canes also. Karly
in March the plot was inspected by one of us (K. J. B.). A number
of the inoculated canes had withered and the disease had extended
in some cases to other canes in the same clumps. The rest of the
plot was quite free from disease, except where the second series of
inoculations, to be described below, was located. The symptoms
were typical and left no doubt of the success of the inoculations.

BUTLER AND HAFIZ 201

On the same date as the last, a second series of inoculations
was carried out in the same plot, by removing the soil from
around the base of four clumps of cane and watering the exposed
roots with a suspension of the mycelium beaten up in water.
These also took well, and when seen early in March each clump
had one or more withered canes. No other canes were withering
in the rest of the row to which the clumps belonged nor, so far as
could be observed, in any part of the plot, except the row in
which the first series of inoculations had been made. One cane
from each clump was split and found to have typical symptoms of
the disease, but there was no opportunity for microscopic exami-
nation. The symptoms were, however, so definite that there
was no doubt that this series was quite as successful as the first.

As the disease does not occur at Pusa, it was not considered
advisable to carry out any inoculations in the field. Some were,
however, made on plants growing in tubs, at a distance from the
farm crop. The number of canes inoculated through stem wounds
was six, growing in four tubs, one tub being kept as a control.
The inoculations were made in November 1909, exactly in the same
manner as those of the first series at Samalkota. In the following
March none of the inoculated canes had withered and two were
removed for examination. The inoculated internode was found
diseased in both, the mycelium being confined to the one internode
in one case but extending to the next higher upin the other. The
other plants were kept in the tubs until the following year, by
which time they were still alive but had ceased growth, except for a
few feeble side shoots. In May 1911, they were cut and examin-
ed. The inoculated canes were much reddened near the wound,
the reddening extending for from 2 to 4 internodes each way.
Above the reddened part, the characteristic translucent appearance
and internodal hollowing of the pith was found. The red parts
were full of the hyphe of the parasite and, in ene case, pyenidial
stromata had developed in the internode next below that inocu-
iated, The roots were not affected and no spread to other shoots
in the same stool had oceurred. The parasitism was, therefore,
much less marked than at Samalkota, though the spread of the

202 NEW SUGARCANE DISEASES

mycelium in the living tissues indicates a certain degree of para-
sitic activity.

In three other tubs, the main canes of which had been injured
and in which a “ ratoon” crop was growing, inoculations similar
to those of the second series at Samalkota, were made in Decem-
ber 1909, the exposed roots being watered with a suspension of
the fungus. No outward sign of disease appeared, and the follow-
ing April the stools were uprooted, as also that in the control
tub. All were found perfectly healthy and no trace of the parasite
could be detected in the tissues.

It seems probable, therefore, that the disease is restricted in
its distribution by climatic or other unknown conditions, as so
many of the fungus blights of crops in India are. How much
damage is caused by it is unknown. Both at Samalkota and
Jorhat it was enough to cause uneasiness, but in the latter case it
is still uncertain whether there are not two diseases at work. If
it should be found that the condition observed at Jorhat, in the
Central Provinces, and elsewhere, of excessive tillering, combined
with degeneration of the plant, sometimes to such an extent that
no true canes at all are formed, the whole plant remaining grass-
like, is caused by this parasite, then it will have to be reckoned
as a serious disease. Meanwhile, further investigations are in
progress and it is hoped that the publication of the present des-
cription will lead to the recognition of the disease, where it occurs,
and fuller observations on its field characters. It is obviously
impossible, at present, to suggest methods for its check beyond
the ordinary precautions urged on previous occasions for routine
observance in sugarcane cultivation in India.

Descrivtion oF Puares III, IV anv V.
(Hendersonina Sacchari, Butl. )
Piate III,
Fig. 1. Section of a sclerotium of Headersonina Sacchari, probably formed by the
union of several unilocular stromata. Mach cavity is surrounded by a
separate pseudo-parenchymatous wall, between which the structure is

filamentous. A mass of spores of both types is extruding from the
mouth of the left-hand cavity.

PLATE Iv,

K. D.Das.Cp.

HENDERSONINA SACCHARI,

Puate V.
alg,

a
gS
4

~ “oe
=~ > | - PAL
yY =, a
~ @s red
xo Ry <= Y
A s v
>
S \
° » >
¥ > <> =<
P

eos 2
%09

Hendersonina Sacchari.

emi ee 5 ee. ke ek ak nn Al nas of tha Survac et Tudis, Calcutts, 1918.

Fig.

BUTLER AND HAFIZ 2038

2, Ripe sclerotium in longitudinal section, showing the irregular loculi, immersed
in a pseudo-parenchymatous stroma, The cavities were filled with spores,
which are not shown,

3. Another ripe sclerotium in transverse section, At the base, this sclerotium
showed only two large loculi. Near the apex there were seven cavities,
arranged roughly in a ring and mostly opening by a common mouth.

Pirate IV,

_1. Transverse section of part of the pith of a cane, infected by Hendersonina

Sacchari, The hyphe grow from the intercellular spaces into the cell
cavities, Intermediate and fine hyphex are present, but not the very
thick kind. X 190,

2. Very thick hypha from the surface growth obtained on incubating an in-
oculated cane. Branches of intermediate diameter are numerous. These
in turn will give off very fine threads. Septation has not yet occurred,
X 350;

. 3. Mycelium from a culture, showing fine and intermediate hyphe and chlamy-

Fig.

dospores, ‘The fine branches are irregularly swollen, and at «a, two have
anastomosed by a side branch, Later stage of growth than Fig. 2.
X 350,

4, Chlamydospores separated from the mycelium and germinating, X 500,

». Formation of chlamydospores. w early stage, intercalar ; 4 ditto, terminal ;
ca chain separating from the mycelium (one is germinating), d old
chlamydospores from a chain, that on the right has lost part of its
exospore, X 350,

(. Part of the wall of a pycnidium, showing the abrupt transition from the
sclerotial cell layers to the basidial layer,

‘. Basidia with filamentous spores (scolecospores). X 625,

8, Basidia with the broad type of spore (pyenospore). X 625,

). Simple types of basidium, the 3 on the left with hyaline scolecospores of a
broader type than usual, that on the right with a young pycnospore,
X 628,

Pruate V.

1. Pycnospores of Hendersonina Sacchari, 520:

2. Scolecospores of ditto. Some stromata contain only the narrow type,
uniform in diameter shown at @ a, others the more irregular forms,
X 520,

. 3. Intermediate type of spore between the pyenospore and scolecospore types,

X 520,
+, Germination of the pycnospores, early stages (24 hours after sowing).
X 520.
Ditto, later stages (48 hours after sowing). X 520.

or

I11.—Hetminriosporiose
(Helminthosporium Sacchari Butl. n. sp.).

A species of this well-known genus, many members of which
are parasitic on Graminew, is common on the leaves of sugarcane
at Pusa. The infected leaves first show small red spots, which
spread rapidly, chiefly in a longitudinal direction and, especially
towards the tip of the leaf, may run together to form long streaks.
The centre of the spot soon changes to a dirty straw colour, around
which the margin remains red for a time and then changes to dark
brown. The spots occur equally on the midrib, where they may
be confused with those caused by the leaf form of Colletotrichum
falcatum,* and on the thinner part of the leaf. When numerous,
they cause death of the leaf tissues beyond the limits of the spots;
the tip of the leaf often withers completely and there may be
long withered strips down the margins.

The mycelium of the parasite is found in the leaf cells of the
spotted portion and also collects in small stromatic masses on the
surface of the spot. The hyphe are brown at the surface and in
the outer cell layers, but hyaline deeper in. They pass from cell
to cell through narrow cracks in the walls (Pl. VI, Fig. 2), which
are especially noticeable in the thick-walled sclerenchyma which
overlies the bundles, but within the cells are swollen so as almost
to fill the cavity in the smaller cells. In the epidermis they fre-
quently form small stromatic masses. The cells appear to be
killed in advance of the growth of the fungus, as although the
hyphe are numerous in the dead cells, it 1s rare to find penetra-
tion of a still living cell.

As soon as the centre of the spot begins to turn straw-
coloured, fructification occurs by the growth of sporophores from

* Butler, K.J. & A, Hafiz Khan. Red Rot of Sugarcane. Mem, Dept, of Agric. in
India, Bot. Ser., VI.!No. 5, 1913.

BUTLER AND HAFIZ 205

the stromata, both those within the epidermal cells and those on
the surface of the leaf.

The sporophores are stout, erect, rather rigid hyphie, which
arise from the peripheral cells of the stromata (Pl. VI, Fig. 4).
They are usually unbranched, 3 to 10 septate, dark greenish-brown
below, paler above and several times bent or ‘‘ geniculate.” Spores
are produced at each bend and at the apex, the lowest being the
first formed and the bent condition being due to the spores being
always apical at first and being then pushed to one side by con-
tinued growth of the sporophore from just below the insertion of
the spore. The sporophores are 100 to 190 » long, by 5°5 to 75 u
broad.

The spores (Pl. VI, Fig. 5) are borne singly and readily fall
off They are cylindrical or long elliptical in shape, with very
thick walls, and divided into from 4 to 11 compartments by broad
partitions. The colour varies from olive green to brown and
the size from 35 to 60 long, by 8'5 to 12 u broad.

Germination occurs rapidly (within 3 hours in some cases)
hy the protrusion of a germ-tube from each end of the spore. At
the same time the internal partitions sometimes break down in
the centre (Pl. VI, Fig. 6). Part of the mycelium formed from a
single spore culture in a hanging drop, is shown in the text figure :

ED Das Gp

206 NEW SUGARCANE DISEASES

drawn 48 hours after sowing. The fungus can be readily culti-
vated on most ordinary media, but the spores formed in culture
are usually smaller than those from the host plant. The
following inoculations were made with pure cultures on nutrient
agar,

In January 1909, 8 leaves of growing sugarcane were iocu-
lated on the upper surface with a suspension of spores and. my-
celium, a drop of which was placed on each leaf and covered with
damp, sterile cotton-wool. The leaf was not injured in any way.
After 6 days, 3 of the inoculations were found to have succeeded,
a red spot developing at the inoculated point and eventually giv-
ing rise to a typical infection. The other 5 and the controls
showed no change. <A second series was made at the same time
on young healthy shoots brought into the laboratory. On these
6 inoculations were made, 3 being kept damp with cotton-wool
and 3 by covering with bell-jars. The latter showed evident
signs of infection in 2 days, but only 1 of the former succeeded
and the spot was not visible until the fourth day. A month
later 9 more inoculations were made on healthy shoots kept
under bell-jars, all of which succeeded. One showed reddening
in 24 hours, 4 more were visible in 3 days, and all were well
marked in 5 days. A little later, 21 more inoculations were made
in a sunilar manner to those of the last series. All succeeded
well.

The microscopic examination of the inoculated leaves showed
that penetration occurred directly into the epidermal cells
(Pl. VI, Fig. 3). The infecting hypha swells up slightly in
close contact with the cuticle and a narrow tube arises from the
swollen portion and pierces the wall. As soon as the cell cavity
is entered, the hypha swells up again and becomes freely septate,
often forming a small mass of pseudo-parenchymatous cells.
From this, branches pass to the deeper cells, and others approach
the surface to form new stromata. The pressure exerted by these
epidermal stromata is sometimes enough to rupture the outer
wall before reproduction begins. The fructifications are usually
confined to the quite dead central part of the spot.

BUTLER AND HAFIZ 207

The species does not seem to agree with any of those pre-
viously described. About 30 are known on various grasses, but
none on Saccharum. The diagnosis is as follows :—

Helminthosporium Sacchart Butl. n. sp. Maculis amphig-
enis, elongatis, initio rubris, dein avellaneis vel stramineis ac
ferrugineo-marginatis, 3-25 W 2-6 mm; cespitulis minutis, atris ;
hyphis fertilibus erectis, simplicibus, 3-i0—septatis, geniculatis,
olivaceo-brunneis, apice pallidioribus, 100-190 \Y 5°5-7°5 «; coni-
diis acrogenis, cylindraceis vel oblongo-ellipticis, utrinque
rotundatis, 3-10-septatis, crassissime tunicatis, olivaceo-brunneis,
35-60 VW 85-12 um.

In folis Sacchare officonarum in India, or

A fungus which, from the published descriptions, bears a
considerable resemblance to the above, is known in Java and
Hawaii under the name of Cercospora Sacchari, Br. de Haan.
It was first described by Breda de Haan in Java,* as the cause
of a leaf disease to which the name ‘‘eye spot” was given. A
further account was subsequently published by Dickhoft and
Heint. In Hawaii it has been briefly described by Cobby.
From the figures published in Wakker and Went’s well-known
text-vook of sugarcane diseases\, it appears probable that this
fungus is really a Helminthosporvwm and not a Cercospora. A
comparison of the two fungi has not been possible and could alone
settle the question of their identity.

Little requires to be said regarding the treatment as, so far,
this disease has not been found to damage the crop materially.
If it became severe, it would be advisable to strip off and destroy
the affected leaves in the early stages of the attack (we have
found the first spots in June, at Pusa). There area good many

* Breda de Haan, J. Van, Rood Rot en andere Ziekten in het Suikerrict, Meded. van het
Proefstation West Java, XVI, 1892.

+ Dickhoff, W. G. & 8. A. Arendsen Hein. Kenige Waarnemingen omtrent de Oogylekken-
ziekte, Archief Java Suikerindustrie, 1X, 1901, p. 865,

{ Cobb, N. A. Fungus Maladies of the Sugarcane. Exper. Stat, of the Hawaiian Sugar
Planters’ Association, Division of Pathology and Physiology, Bull, No. 6, 1909, p. 36.

§ Wakker, J. H & F.A F.C. Went. De Ziekten van het Suikerrict op Java. Leiden.
1898, Pl. XXI, figs, 1—5,

208 NEW SUGARCANE DISEASES

leaf diseases of sugarcane, but they cause little loss as a rule and,
even in more advanced countries than India, do not seem to

require treatment.

Pusa,
August 5th, 1913.

Descriprion oF Puate VI.

(Helminthosporvum Sacchari But.)

Section through leaf of sugarcane, showing hyphe of //elminthosporium
Sacchari in the outer cells. X 350,
2. Part of a similar section, to show passage of the hyphe from cell to cell,
X 500.

_ 3. Infection of upper surface of leaf, from an inoculation, showing penetration
of the outer wall of the epidermis and formation of small stromatic
masses in the epidermal cells. X 950,

4, Conidiophores with young conidia, X 350.

Conidia. The three upper immature, the three lower mature. X 500

6, Germination of the conidia. X 500.

Reduced conidiophore and conidium, from a pure culture, X 350,

8. Base of conidiophore, arising from within the epidermis and emerging across

the cell wall, not through a stoma. X 500.

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Botanical Series, Vol. VI, No. 7
v4

———— Remoirs of the
Be Department Of Agriculture
eo in India

- PRELIMINARY NOTE ON THE CLASSIFIGATION OF
RICE IN THE GENTRAL PROVINCES

BY

, > R. J. D2 GRAHAM, M.A;, B.Sc

Economic Botanist, Central Provinces

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ge habits RESEARCH INSTITUTE, PUSA

PUBLISHED FOR

Pe Tat IMPERIAL DEPARTMENT OF AGRICULTURE IN INDIA

re BY d
a ee SPINK & CO.; CALCUTTA

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a Price, ‘Rs. 1-8

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December 191-5. BoraNiCaL SERIES. Vou. VI, No. 7

MEMOIRS OF THE
DEPARTMENT OF AGRICULTURE
IN INDIA

PRELIMINARY NOTE ON THE CLASSIFICATION
OF RICE IN THE CENTRAL PROVINCES

BY
R. J. D. GRAHAM, M.A., B.Se

Economic Botanist, Central Provinces

LIBRARY

NEW YORK

FANICAL
GAKUVEN,

AGRICULTURAL RESEARCH INSTITUTE, PUSA

PUBLISHED FOR

THE IMPERTAL DEPARTMENT OF AGRICULTURE IN INDIA
BY
THACKER, SPINK & CO., CALCUTTA
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PRINTED BY
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PREFACE.

THE material made use of in the preparation of this Memoir has
been gathered from observations made at the Nagpur Farm, and
the information supplied by the District Officers of the Central
Provinces and Berar. To these officers I desire to tender my
sincere thanks for their unfailing kindness in collecting information
and specimens. Without their assistance it would have been well
nigh impossible to bring together a satisfactory collection of rices.

The paragraph on ‘ Rice Soils’ was written by Mr. F. J.
Plymen, Agricultural Chemist, C. P. To him my best thanks are
due for permission to include the note in the present publication.

R. J. D. GRAHAM,
NAGPUR, Economic Botamst, C. P.

July 1913.

Mast TY S

PRELIMINARY NOTE ON THE CLASSIFICATION OF RICE
IN THE CENTRAL PROVINCES

BY
R. J. D. GRAHAM, M.A., B.Sc.

Economic Botanist, Central Provinces.

I.—INTRODUCTORY.

Preliminary.—The work of classifying the rices of the Central
Provinces has been in progress during the past five years. The
present memoir is a summary of the work to date, and is published
in the hope that the results attained may prove of service to workers
in other parts. The materials on which the enquiry was started
were the rices collected from all the districts in the Central Provinces
for the Central Provinces and Berar Exhibition of 1908. In the
succeeding years this collection has been added to, largely through
the kindness of District Officers, till at the present time a collection
of 670 rices has been received. In cataloguing the specimens the
vernacular names by which the rice was known in its own district
have been made use of, and it must be clearly understood, that the
670 rices referred to above represent only vernacular names and
not necessarily distinct varieties. As a matter of fact, the ver-
nacular name is no guide to the identification of a rice, e.g., the same
vernacular is often applied to totally distinct rices in different
districts ; while, on the other hand, the same rice often bears different
names in different districts ; for example, the outstanding purple
rice is named Nagkesar in Chhattisgarh, Jalkesar in Jubbulpore and
Mahaprasad in Nagpur.

Area.—In 1911-12, the most recent year for which figures are
available, the area under rice in the Central Provinces was 4,780,560
acres out of a total of 17,961,358 under crop. In Berar 41,487

1

210 CLASSIFICATION OF RICE

acres out of 7,057,414 were under rice. Thus in 1911-12 rice
constituted 26°6 per cent. of the total area under crop in the Central
Provinces or 19°2 per cent. of the whole area under crop in the
Central Provinces and Berar.

The earliest complete figures available are for the year 1885-86,
when rice occupied 3,170,360 acres out of 13,466,675 under crop,
in the Central Provinces, which at that time included Sambalpur,
while in Berar the area was 25,832 out of 6,558,379 acres under
crop, with percentages of 23°5 per cent. and 15°9 per cent. for the
Central Provinces and Berar respectively. Rice moreover occupies
the largest area in the Central Provinces, its nearest rival being
wheat which occupies 14°8 per cent. of the area. In the Central
Provinces and Berar rice and cotton respectively occupy nearly
the same area, cotton being slightly the greater and taking up 19°5
per cent. of the whole.

Detailed figures for 1885-86 and 1911-12 for each district are
given below in a tabular form.

a tT hl Pe eee Dennen en eal nes LS

aiid Area in Area in
District. 1885-86. 1911-12.
Acres. Acres.
Saugor 12,575 17,816 |
Damoh 49,163 49,358 |
Jubbulpore 1,22,160 1,336,131 |
Mandla 47,385 1223312
Seon ae 1,06, 103 94,634
Narsinghpur .. 25,659 44.144 |
Hoshangabad 11,465 11,984 |
Nimar 12,569 12,357
Betul 12,096 12,058
Chhindwara 5,415 11,578
Wardha 4,050 4,349
Nagpur 32,763 28,948 |
Chanda 1,89,330 Deol tala
Bhandara 2,72,855 4,85,153 |
Balaghat 2,27,385 2,84,252 |
Drug a 655,099 |
defor 11,44,757 | 14,28,568 |
Bilaspur 7,16,620 | 11,19,023 |
Samba] pur 1,76,013 ee | ‘Transferred to Bengal.
Amraoti 559 6,150
Akola 331 16,171
Ellichpur 4 on Included in Amraoti.
Melghat 4,293 a Ditto
Buldana 6,180 5,282 |
Wun 2,956 33 Included in Yeotmal.
Yeotmal ois 13,884
Basim 11,509 Ly Included in Akola.

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REFERENCE
Province Boundary
District. ... Do
Feudalory Slate Do
Tahsil, De
Hoa. Q°8 of the District

Do Tahsuh

Town .,

Railway completed « ..

Do Projected...»

Rivers,

R. J. D. GRAHAM 211

The principal rice growing tracts (vide Map) of the south of the
Province are the portions of Bhandara, Balaghat and Chanda lying
in the Wainganga river basin all in the Nagpur Division, and Drug,
Raipur and Bilaspur in the Chhattisgarh Division. In the north of
the Province the south-east portion of Seoni with Burghat as a centre,
parts of Mandla and the southern portion of Damoh with the two
adjoining Tahsils of Jubbulpore, constitute the most important
areas.

Rice Soils.—The rice-producing soils of the Central Provinces
are formed almost entirely upon Pre-Cambrian rocks of the Purana
and Archean groups. The formations generally represented are
the Upper and Lower Vindhyan in Damoh, Bilaspur, Chanda, Drug
and Raipur; the Chilpi Ghat series of the Dharwar formation,
formerly known as the Transition or Sub-metamorphic, in Bhan-
dara, Balaghat, Drug and Bilaspur, and the Metamorphic and
Crystalline formation in parts of Jubbulpore, Chanda, Balaghat
and Bhandara.

The actual material derived from the weathered rock is greatly
modified in many cases by rain wash and admixture with soils of
other types. The typical rice soils of the Central Provinces have
a high proportion of coarse particles and a correspondingly low
proportion of clay, being suitable only for single cropping. The
heavier soils suitable for double cropping are much finer in
texture and therefore more retentive of moisture.

Composition of typical rice soils of the Central Provinces :—

A. Locality, Adhartal, Jubbulpore, a gneissic detritus.

B. > Katangdhara, Chanda, a gneissic detritus.

C Labhandi, Raipur, a yellow metasi soil overlying Raipur limestone.
1D} a Lanji, Balaghat, yellow sehar soil formed from Dharwar rocks.

BE. 33 Tharsa Nagpur, a black morund soil overlying crystalline rocks ; suit

able for double cropping.

A B C D E
Coarse sand 1—"2 m.m. diam. ; 22°59 43°38 5°66 12°69 16°45
Fine sand ‘2-04 m.m. diam. ae 31 bT 20°15 19°08 28°51 11°53
Silt ‘04-01 m.m. diam. eg Vicia 7°88 38°82 24'79 13°66
Fine silt ‘01-002 m.m. diam. m 7°93 10°83 19°22 14°31 15°76

Clay below ‘002 m.m. diam. AA 1110 13°05 11°67 13°90 34°79

212 CLASSIFICATION OF RICE

A B Cc D EK
Calcium Carbonate a a 0°31 0°20 0-08 0°04 118
Loss on ignition 2°42 4°47 3°58 3719 5°02
Nitrogen ay “27 “44 “45 “48

Cultivation.—Rice 1s grown throughout the Central Provinces as
a rains crop, only in the Sironcha tahsil of Chanda is a cold weather
rice crop grown. This is locally known as jimsara or grishma
kal. Briefly stated there are three methods of rice cultivation.
The first is broadcast sowing, general names for which are bootva
in Bhandara and 6ota or boar in Balaghat. This takes place either
before or immediately after the first showers of the monsoon. Dry
broadcast sowing is known as khurra in Chhattisgarh and as topa
or jhura in Damoh, while wet broadcast sowing is termed rasbeetara
in Chhattisgarh and also batar in Bilaspur and Drug. In the latter
district hogdum is the name applied to wet broadcast sowing when
the seed is covered by the plough. Antia is the term used for wet
broadcast sowing in Chanda and Bhandara. A variation of wet
broadcast sowing is common in the Chhattisgarh Division, in
Balaghat and Chanda and in Damoh and Jubbulpore districts. The
rice grains are placed in an earthen pot or a basket and thoroughly
moistened. This is then covered with cow-dung and straw. In 2-4
days the grains germinate. The artificially germinated grain is
then broadcasted in fields which have been carefully puddled. This
process is known as lehi in Chhattisgarh, kaorak or lehi in Drug and
Balaghat, mulka or kaorak in Chanda and kaorak in Bhandara. In
the Northern districts it is called machhawa.

The second method of rice cultivation is biasi which consists
of broadcasting the rice by any of the methods mentioned above,
followed by a second process of ploughing when the plants are a
foot high. ras: is confined almost entirely to the Chhattisgarh
Division.

Thirdly, transplantation called ropa in Drug, rohna in Chanda,
and rohna or parha in Balaghat and Chanda, a process by which
the seed is sown in carefully prepared and heavily manured nursery
beds at the beginning of the monsoon ; when the plants are 12—18

Rk. J. D. GRAHAM 213

inches high, they are planted out in the rice fields, which have been
especially prepared. The advantages of this method, when pro-
perly carried out, are a considerable saving in seed, a marked in-
crease in yield and an improved quality. The chief disadvantages
attaching are the necessity of a large supply of labour at one time,
the necessity of opportune rain, in the absence of irrigation, to en-
able the fields to be prepared and a decided lengthening of the time
necessary for the crop to mature. The physiological factors involved
in transplantation are somewhat obscure, but the conclusion at
present, arrived at after a number of experiments, is to the effect
that transplantation acts in a way like root pruning, the injury
to the root system stimulating the growth of the sub-aerial portion
and resulting in increased tillering. ‘The root system of the trans-
planted rice is developed from the lower nodes of the stem, the
first or seedling root system in many cases dying completely. In
fact a series of experiments shewed that amputation of the root
system of the seedlings did not interfere with the development
of the transplanted plants.

In Basim taluk of Akola rice is sown in open fields in lines by
ineans of a drill. In the Zamindaris of Chanda the sAhanor: system
of rice cultivation is practised. A piece of jungle is cleared in the
hot weather. When the ground has been covered with wood to a
depth of a few inches, the whole is set alight. Rice is then sown
in the ashes and this ends the cultivation. This method which is
also followed in other jungly parts of the Province is known else-
where as dahia.

Broadcasting is the usual method of cultivation in the Cen-
tral Provinces. Out of the total m 1911-12, 3,154,908 acres were
under broadcast rice. The practice of transplantation which until
a few years ago was unknown in Chhattisgarh is being introduced into
the Division with great success. The area has risen by 25,000
acres within the last five years. Practically throughout the Pro-
vince the crop is grown in embanked fields. These embankments
collect the rain water. Only in hilly tracts ina mixture with Kodo
and Kutki or as a catch crop incerta parts of Wardha, Jubbul-

214 CLASSIFICATION OF RICE

pore, Betul and Hoshangabad are quickly ripening rices sown in open
fields.

Methods of work.—The line of attack has been by means of pure
cultures. Every rice received has been observed in cultures as
follows:—From each variety sent in, four of the largest and at the
same time most typical heads were selected. In a number of
instances where the sample was very impure, subordinate types were
selected ; in some cases as many as six were found necessary as for
instance in Sikia Bikia (Jubbulpore). Of these heads three were
mounted as herbarium specimens for reference, while the grains from
the fourth were removed and placed in an envelope preparatory to
sowing. The contents of each envelope were sown in a line and the
produce compared with the specimen on the herbarium sheet.
Where the produce differed from the type, further pure cultures
were made, heads being selected from the original sample, and at the
same time the types found in the produce were separated out as
indicated in the case of the original selection. In this way it was
possible to check results, and at the same time to observe how far
natural crossing took place. So far no cases of natural crossing
have occurred in the line cultures. The occurrence of the grains
of a wild rice, Tari, in many samples of rice received from Raipur
together with the fact that the presence of Tari in a rice field is held
by the cultivators to lead to the deterioration of the crop suggests
however the occurrence of natural cross-pollination. This is parti-
cularly interesting on account of the occurrence of a similar pheno-
menon in Texas.(1) The presence of a small quantity of red rice
plants is sufficient to vitiate a white rice crop which can only be kept
pure by the weeding out of the red grains before sowing. Since
this preliminary note was written, Hector(2) has shown that natural
cross-pollination takes place at Dacca to an extent provisionally
estimated at 4 per cent. The absence of any cross-pollination at
Nagpur would seem to indicate that this only takes place under
certain limited conditions. A further check was maintained by the
selection of the produce of the best lines for trials on large plots.
The produce of each of the original lines was compared and the best

R. J. D. GRAHAM 215

were chosen for further trials against the best local varieties. This
second selection served a double purpose. It raised the work of
classification from the level of a mere piece of academic research to a
work of practical value and it furnished a check, were such needed,
on the observations made while the particular rice was grown in
hne culture.

Problems involved in Improvement.—There are two main openings
for the improvement of the rice crop, depending on the quality and the
quantity or yield of the rice. Generally speaking, the smaller and
more slender gramed rices are considered of a finer quality. A
second quality found in combination with the finer grain is fragrance.
This is a peculiar mouse-like smell possessed by the grain and also
noticed when the rice is in flower in the field. This fragrance, though
not appreciated by Kuropeans,—it is said that a distinguished mem-
ber of the Agricultural Service soon after arriving in the country
fell foul of his cook for serving him with rice with a mousey past—is
held in high esteem by the natives of India.

These finer rices naturally weigh lighter than the coarse ones.
The weight in 1911 of the grain from 200 heads of a coarse rice
usually lay between 650 and 750 grams though Gangawaloo (Chanda)
weighed 1,041 grams. On the other hand, a similar number of
heads of a fine rice usually weighed between 300—400 grams, the
lowest weight being 223 grams in Bagmochh (Jubbulpore). The first
problem, then, is to get a high yielding fine rice. The second pro-
blem is simple, to improve the yield of the coarse rice.

A minor problem is introduced by the practice of polishing the
rice before it goes to the market. In the polishing process the rice
is hable to be broken and the price lowered. Generally the longer
the grain the more lable it is to be broken. This is a second reason,
the first being the hghter weight of the grain, why the finer rices are
usually nearly double the price of ordinary.

Wild Rice.—The investigation of the wild rices of the Province
is still in progress. The following is a summary of the facts. Wild
rice is common on the margins of tanks, Dewdhan (Jubbulpore), Ghori

216 CLASSIFICATION OF RICE

Pashar and Bal Pashar (Raipur), m marshy places and as weeds in
the rice fields ‘ Pashi* and ‘ Sada’ (Jubbulpore), Tari, Kala, Karanga
and Karanga Pashar (Raipur) and ‘Parsad,’ of which there is an early
and a late variety, from Chanda. The most important distinction
between the wild rice and the cultivated is that the spikelets, when
the grain is mature, are very deciduous. They are usually stoutly
awned and have a dark red grain. The wild rices growing in tanks
are usually tall, having the powers of adapting themselves to the
depth of water in which they grow, and their grain takes longer to
mature. The grain of the wild rices is harvested by Gonds and
Dhimars who tie the rice plants into clumps and thus prevent
the grain falling. The grain is sold to the poorer classes and
is also used by devout Hindus on fast days (Upass) and on the
‘Harchhat ° festival.

The awned wild rices seem to agree with Oryza sativa var.
fatua, Prain.(3) A wild rice from Raipur with a dirty white
grain and an awnless wild rice from Bhandara appear to be culti-
rated rices and run wild. Watt (4) records the discovery by Duthie
of O. officinalis, Wall, syn. O. latifolia, Desv., in Chanda. This wild
rice differs from the othersin having multiveined leaves. Its
occurrence is reported from Allapilli and Sironcha. As already
mentioned the occurence of * Taz,’ whose grains do not fall down,
mixed with many varieties from Raipur, suggests the occurrence of
natural cross-pollination.

1I.—CLASSIFICATION.

Preliminary.—The most recent publication on the classification
of rice is the work of 8. Kikkawa.(5) In this work two schemes
of classification are given. The first divides the rices according
to their agricultural characters, the second by the characters of the
grain. Only the second is applied in classifying the Burmese rices.
An older classification of rice, based solely on the grain characters,
occurs in the ‘“* Handbuch des Getreides ” by I. Koernicke.(6)

The use of agricultural characters in a scheme of classification
demands a considerable amount of caution. In the rice plant we

R. J. D. GRAHAM 217

may distinguish two characters which are developed owing to the
special environment in which the plant grows. The first is the time
of ripening and the second the water requirements of the plant.
The first, however, is largely determined by the second. In a large
area with markedly different climatic conditions in the North and
South like the Central Provinces, where the crop is not entirely
dependent on seasonal rainfall, and where many different systems of
cultivation are in vogue, it is obvious that no hard and fast system
of classification based on these agricultural characters can be laid
down. This will be made clear by taking up the agricultural
characters in detail.

Time of ripening.—The rices are broadly divided in these
provinces into three main classes, v7z., early, medium and late,
which correspond with the sub-divisions mentioned by Kikkawa.(7)
Roxburgh, on the other hand, in classifying the Bengal rices recog-
nises only early and late rices.(8) These terms refer to the time of
harvesting of the rices. Early rices, many of which are ready for
harvest by September, are sown on the higher fields where the water
does not collect to a great extent, owing to the catchment area being
small and their receiving comparatively little drainage. These are
also the only rices sown as catch crops and in unembanked fields.
The earlier the variety the poorer and more exposed the field it is
sown in. Late rices are always sown on heavy soils, in embanked
fields, and in low-lying places ; as the name implies this is the last
rice to be harvested, at times being on the ground until the middle
of December, and unless the field is protected by irrigation, the suc-
cess of this crop depends on the rainfall of September and October.
Generally the late rices are grown in the fields that command the
most assured and continuous supply of water. Medium rices are
sown on land intermediate in character. They are longer in matur-
ing than the early rices, but not so long as the late rices. Taking
the average of the rices grown under observation an early rice
matures in 121 days, the range being from 106 to 145 days ; a medium
in 125 days, the range being from 110 to 169 days; and a late rice in
133 days, the range being from 119 days onwards. The period be-

218 CLASSIFICATION OF RICE

tween sowing and the appearance of the ears in the three classes is
87,90 and 91 respectively. It is evident, therefore, that the earliness
and lateness of a rice depends not so much upon the vegetative
period of growth as upon the reproductive period during which the
fruit matures and ripens. Though the rices fall broadly into these
three classes, there appears to be a considerable latitude, even in one
district, in placing any given rice in its class. The difference is
still greater, when rices of the same class from different districts
are considered. Thus early rices from Jubbulpore matured on the
average in 113 days, the range being from 109 to 132 days; medium
rices in 120 days, the range being from 117 to 129 days, and late rices
in 128 days, the range being from 119 to 155 days. The early rices
from Raipur matured on the average in 124 days, the range being
from 105 to 145 days ; the medium rices in 127 days, the range being
from 112 to 141 days; and the late rices in 138 days, the range being
from 123 to 155 days. Comparing these figures :—

Class. Jubbulpore. | Raipur.
Early 113 124
Medium 120 127
Late ie = i e oe ke 128 138

We find that the Jubbulpore rices are distinctly earlier in matur-
ing than rices of the same class from Raipur. This is probably due(1)
to irrigation in Raipur being far ahead of Jubbulpore, (2) to the
practice of biasi in Raipur which delays the maturing of the plant.

This difference is due to irrigation and methods of cultivation in
these districts, coupled with the wide range for the same class even
from one district, shew how unsatisfactory a classification must be,
based on the time of ripening of the rices.

Cold weather Rice.—Geemsal, a rice from the Sironcha tahsil of
Chanda, is generally sown in January and harvested in March.
This is an interesting variety as it is the only cold weather rice report-
ed in the Central Provinces. ‘The yield is said to be poor im spite

R. J. D. GRAHAM 219

of heavy manuring and irrigation. Its survival is probably due to
the isolated position of Sironcha and it will probably disappear as
soon as the country is opened up. The rice can be grown in the rains,
when it behaves like an early rice, and hence it may be placed with
these.

Resistance to drought and flooding.—Rices fall into three main
groups in regard to their water requirements, v7z., drought resistant,
normal and flood resistant rices. The most highly drought resisting
rices can exist for from 20 days to one month without water. In the
North of the Province these are found amongst the early rices, else-
where amongst the late rices. Flood resistant rices are found mainly
amongst late rices. The longest period that a rice can withstand
flooding is for about 15 days. If a suggestion may be made in ex-
planation of the curious phenomenon that amongst the late rices
both drought resistant and flood resistant varieties occur, may it not
be that the danger of flooding to which the late rices, growing in the
lowest fields, are exposed in their early stages, and the late date of
maturing, occurring long after the seasonal rainfall has ceased, have
evolved a type which may be both flood and drought resistant.

Height.—The average height of a rice plant is between two
and half and three and a half feet. There are, however, amongst the
early rices a large number which attain a height of only 18 inches
to two feet. These may be classed as short rices. Only one example,
Pongha, of the tall rices mentioned by Watt(9) has been found in
the Central Provinces. It occurs in tanks in Chhattisgarh, Balaghat
and Nagpur. The height varies with the depth of water, but speci-
mens over 6 feet long are not uncommon. It is not proposed to
make use of the height of the plant as a main point for classification.
The tall rice is interesting as a systematic variety. It along with the
short rices, falls into a group of systematic varieties which may be
termed ‘ specialrices.’ As pointed out by Kikkawa(10) the shorter
rices are less liable to lodge, but any advantage gained from this
is more than counterbalanced by the fact that, so far as the
Central Provinces are concerned, the short habit is associated with
poor tillering and a small panicle.

220 CLASSIFICATION OF RICE

Leaj.—The first diagnostic character noticeable in the young
rice plant is the colour of the first leaf sheath. This can be observed
five days after sowing. The normal colour is yellowish green, or a
green usually distinctly lighter than the colour of the leaf. In
certain classes, however, the colour is a shade of red or purple, in
the latter case sometimes almost bordering on black. The colour
may extend throughout the length of the leaf sheath, or may be
confined to the lower portion, just above the ground level. The
colour may further be either temporary or permanent. Generally
speaking, the deeper shades are lasting, while the lighter shades may
or may not disappear later. One interesting correlation has been
observed in connection with the coloured leaf sheath. All rices
which have a coloured leaf sheath have a dark coloured apiculus to
the glume and palea. The converse is possibly also true, but there
are still a few exceptions which are probably due toa very fleeting
colour in the leaf sheath.

Coloured Rice-—In one variety of rice the plant is a bronze
purple red and a field planted with this variety forms a_ striking
contrast to its verdant neighbours. This is due to a coloured
sap in the cells of the epidermis. In the young leaves the colour
is developed mainly in the epidermal cells lying in the furrows be-
tween the sclerenchyma ridges. As was pointed out by Mooker-
jee(11) this variety would be a convenient one to select for mtro-
duction on account of the ease with which a field could be rogued.
I agree with Kikkawa(12) in his decision that from an agricultural
point of view the difference in colour should not be made a main
point in a scheme of classification. Froma systematic stand-point
the colour of rice might well be described as a variety.

Ligule.-—In the developed leaf another diagnostic character is
found. ‘The ligule which in the rice is continued round the sheath of
the leaf as two ears or “ sickles (13) is either a light yellow colour
or may be black with red. In the latter case the stem appears to
have coloured bands or garters round it. These coloured sickles
are commonly associated with a coloured leaf sheath, but the
association is by no means invariable, as more than one rice with a

R. J. D. GRAHAM rid |

green leaf sheath have been found to have coloured sickles, e.q.,
Parewa (Raipur).

Peduncle.—The peduncle in the rice is variable in length and
may either be exserted from the last leaf (flag) or enclosed by it. In
classifying rices, a peduncle is said to be ‘ enclosed’ when the lowest
branch of the inflorescence is within the leaf sheath, and ‘exserted’
when the lowest branch, easily recognised by its complete ridge of
ciliate hairs, encircling the stem below the branch, is free from,
that is visible above, the leaf sheath. The length to which the ped-
uncle is exserted varies. Those with a peduncle more than half
the length of the inflorescence have been described as ‘ far exserted,’
those with a short peduncle simply as ‘ exserted.’

Inflorescence.—The panicle consists of a main rachis bearing
primary and secondary branches. The rachis may be in one of three
positions. It may be erect, curved or drooping. The last term
is applied to a rachis which bends in a sharp curve.

Branches.—The branches of the inflorescence are all of the
first and second order. The primary branches may either be ap-
proximate or separate. Further the primary branches, like the rachis,
may be either erect, curved or drooping. In certain rices, branches
of both kinds occur. In this case the lower branches are erect and
the higher branches drooping. Taking the three types of rachis
and the three types of branching, the following combinations occur.
The rachis may be erect with approximate, that is, erect, branches—
when the inflorescence is described as “erect.” Secondly, the rachis
may be curved more or less in a semi-circle with approximate
branches—when the term “curved” is used, or the branches may
be separated and themselves curved or drooping when the
inflorescence has a feathery appearance. Lastly, the rachis may be
“drooping” with approximate branches, when the inflorescence is
drooping, or with separate branches when the inflorescence has
_again a feathery appearance.

Gammie (14) makes use of the average number of primary
branches in order to distinguish the rices of the Bombay Presidency.
It is not possible to use this as a diagnostic character in the Central

222 CLASSIFICATION OF RICE

Provinces rices as the number of primary branches is extremely
variable. Thus in‘ Meghai’ (Raipur) the average number of branches
was 10 with a range from 6 to 14 in 83 specimens examined ; ‘Jiro’
(Narsinghpur) had an average of 16 with a range from 13 to 21 in 61
specimens examined ; ‘Jhamul! Meli’ (Raipur) had an average number
of 1] with a range from 8 to 15 in 105 specimens examined. These
three examples, taken from 64 medium rices examined, show how
unsatisfactory this character is as a basis for classification. A range
of eight is by no means uncommon, the ordinary range being six, and
a range of less than five the exception. Two further examples
will show that in certain cases the average number of primary
branches does not even represent the number of greatest frequency.

Rice. | No. of branches. No. of specimens.
= oes eee emit te ae! 3

Laxmibhog .. 1] 2
Raipur 12 10
13 15

14 1)

15 22

16 14

17 13

Is 10

19 I

Average number 14; number of greatest fre|quency 15,

Ari Motan au 4
Raipur 8 14
9 10

10 7

11 14

2 4

Average number 9; number of greatest freq/uency indefinite,

}

Spikelet.—The spikelets are articulated on their pedicels which
may be long or short. The pedicel is enlarged annularly at the top,
the two sides being distinctly oblique, that is, one side is higher
than the other. Just below the enlargement the pedicel is bent or
contorted. In many rices the facet(15) or annular enlargement
is expanded so as to appear scale-like. Hence Cook(16) in deserib-
ing O.coarctata gives the number of glumes as 5, the spikelet being
jointed above the lowest pair. The size of the facets is of a certain
amount of value in distinguishing the varieties.

R. J. D. GRAHAM 223
Three main types of facets may be distinguished, viz., ordinary
(Plate III, Fig. 1), membranous (Plate III, Fig. 2), and ciliate
(Plate III, Fig 3). In the ordinary type the facets are cup-like,
the rim sometimes being thickened: in the membranous type the
margin of the facets is flanged or might be described as having
an expanded lip which is a membranous expansion. The ciliated
facets have short erect hairs on the margins of the facets,

Each spikelet has three glumes, and a glume-like palea. The
outer glumes are usually small and sub-equal, not exceeding 1—} of
the inner glume. The inner glume and palea are sub-equal.

Clustered spikelets (17).—Although the spikelets are usually solit-
ary, in ‘ Bahia Baikoni’ (Raipur) and‘ Benie’ or * Shingofri’ (Jubbul-
pore) the spikelets are clustered on the secondary branches, from 2-7
occurring close together (Plate IV). The branches of the panicle
have consequently a slightly interrupted appearance. The double
grained rice indicates a direction in which we may look for an

increase of the yield.

Outer Glumes.—The outer glumes are usually small, coriaceous
and shiny. Glume I is inserted at a distinctly lower level than
glume II. Both glumes bear the impress of the facets of the pedicel.
The colour is usually somewhat paler than that of the inner glume
and palea, ranging from a pale yellow approaching white through
red to black. Ordinarily the outer glumes are inconspicuous, but in
‘ Bolita’ (Drug) and in ‘ Kalidhan’ (Hoshangabad) they are pale, while
the inner glumes and paleaare black. Further in ‘ Bhojraj,’ « Naku’
(Hoshangabad) and ‘ Ranikajar’ (Jubbulpore) the outer glumes are
dark, while the inner glume and palea are light. The colours of
the outer glumes are useful in differentiating the varieties.

Rachilla— Between the outer glumes and the inner glume the
rachilla is frequently expanded in an annular thickening. This
thickening tends to force the outer glume and palea apart. The
extent to which the thickening is developed can be used to
distinguish the rices.

b>
bo
—

CLASSIFICATION OF RICE

Broadly speaking, there are two types of rachilla, viz., comma-
shaped and elbow-shaped (Plate II).—The comma-shaped raehilla
has an enlarged thickened portion immediately under glume III,
and the palea with a more slender curved stalk (Plate II,
Figs. 1, 4). The elbow-shaped rachilla is uniformly thickened and
does not show the distinction into head and stalk (Plate LI,
Figs. 2, 3).

Winged Spikelets(18).—The outer glume in ‘Lal Pankha’
(Chanda), ‘ Pankha’ (Chanda), and ‘ Pakharya’ (Chanda) are large,
equalling or exceeding the spikelets. Such rices have a characteristic
winged appearance. This phenomenon, though interesting from
a systematic standpoint, is limited to only a few rices and is not
of much importance in the present scheme of classification.

Inner Glume and Palea.—The inner glume and palea are boat-
shaped, nearly equal, longer than the grain and enclose it com-
pletely. Glume II is strongly 5-nerved, spinescently hairy on the
nerves. The palea is narrower than the glume, 3-nerved, the nerves
being spinescently hairy. The commonest colour is a pale yellowish
white, but all colours from this to black, through red, occur. The
colours are constant within limits, and are useful as diagnostic
characters. Frequently the colours are confined to the furrows be-
tween the nerves, the nerves being themselves pale, while in a few
rices the upper position of the glume and palea are one colour—com-
monly red, while the lower portion is another, usually pale yellow,
the spikelet thus having a piebald appearance.

Apiculus.—The portions of glume III and _ palea are solid
and project in a larger or smaller terminal point or apiculus.
The apiculus and tip of the glume may be uniformly coloured
with the rest of the spikelet or may be differently coloured,
usually dark brown. At the time of flowering the tip of the
glume is either concolourous with the rest of the glume, red
or black. These two latter give the dark brown apiculus and
tip of the ripe spikelet. As already pointed out, this colour is
normally associated with a coloured leaf sheath. On either side
of the apiculus are two smaller projections which make the tip

R. J. D. GRAHAM 225

of glume III tridentate (Plate II, Fig. 2). These are described
by Watt(19) as glandular processes. As stated later by Watt,
they are simply excrescences of the lateral veins of the glume.

Awn.—An awn, if present, is borne on glume II]. The awn may
or may not be articulated on the glume. The length varies in dif-
ferent varieties from } inch to 3} inches. Even within the same
variety the length of the awn is by no means constant. The colour
may be white, red or black, the first bemg the most common. The
awn is scabrid and in the wild rice was undoubtedly associated with
fruit dispersal and fixing the grain in the soil. In the Central Pro-
vinces no aversion is met with to awned rices; in fact, the best and
highest quality rice Chinoor has an awn. In Damoh and
Jubbulpore awned rices are sought after as the awns protect the
crop from the attack of pigs.

Size.—The spikelets may be roughly divided into four groups.
Long spikelets in which the length is more than four times
the breadth; fine in which the length is more than three times
the breadth; coarse in which the length is more than twice the
breadth; yound in which the length is less than twice the
breadth. Any attempt to give definite measurements of Central
Provinces rices has ended in failure. True, an average measure-
ment can be made out, but the labour and time required for
this is In no way compensated by the results. For practical
purposes the above is sufficient.

Shape.—Kikkawa(20) groups the rices into six classes according
to the shape of the grain. No description is given of the classes,
and an unlettered photograph does not help to elucidate them.
Broadly speaking, there are five distinctive shapes found in the
Central Provinces. The shape naturally depends on the curvature
of the glume and the palea. The classes are as follows :—

I. Glume and Palea slightly convex. (Plate I, Fig. 1.)
II. Glume and palea convex. (Plate I, Fig. 2.)
III. Glume and palea very convex. (Plate I, Fig. 3.)

bo

226 CLASSIFICATION OF RICE.

IV. Glume slightly convex, palea convex. (Plate I, Fig. 4.)

Y. Glume slightly convex or straight, palea straight or slightly
concave. (Plate I, Fig. 5.)

Among these five classes the first is by far the commonest.
In the fifth class the shape often appears twisted or = shaped.

As a rough guide to the meanings of these terms a slightly
convex glume is four times as long as broad, convex three times,
very convex less than three times: a slightly convex palea is five
times as long as broad, convex four times, very convex less
than four times.

Stigma.—Kikkawa(21) mentions that the colour of the stigma
can be used to distinguish the rice varieties. Three colours are
found in the stigma, v7z., white, red and black. The white stigma
is found in the rices which have a green leaf sheath, the red in
association with the red leaf sheath and the black with the purple
leaf sheath. It is therefore of little use to bring in an additional
character which is associated in the Central Provinces rices with
another which can be more easily made out.

GRAIN.

Glutinous Rices.—So far as the present collection has gone,
no examples of glutinous rices have been found in the Central
Provinces. Glutinous rices have been experimentally grown on
the Raipur Farm, but with little success. They were also imported
during the shortage of the rice crop in 1907, but were not ap-
preciated even in a year of scarcity, so there seems little likelihood
of an opening for them in the Central Provinces.

An analysis of a number of Central Provinces rices by Hooper
shows that the rices are in no way inferior to those grown in other
provinces.

Colour.—According to the colour of the grain, the rices of the
Central Provinces fall into two groups. The first group contains
all the white rices, while the second contains the coloured rices.
By far the commonest in the Central Provinces are the white rices.

R. J. D. GRAHAM 227

The colour varies from pale yellowish white to a deep yellow.
Amongst the coloured rices a dark terra-cotta is the commonest,
but this may shade off into a light red or orange. The colour
is contained in the pericarp. The process of polishing rice
consists in the removal of the pericarp. This is rendered difficult
by ridges on the grain. So far as can be ascertained, no objection
is found to coloured rices, though, it is admitted that they
are more difficult to polish and consequently are more liable
to break in the process. There is no relation between the
colour of the spikelet and that of the grain. In‘ Naku’ (Hoshanga-
bad) the spikelet is coloured, but the grain is terra cotta, while in
‘ Bainssa’ (Raipur) the spikelet is almost black with a pale yellow
grain. .
Size.—The size of the grain corresponds approximately with
the size of the spikelet and the grains may accordingly be allowed
to fall into the same classes in which the spikelets were placed with
the exception of those with the long spikelet. No grain witha
length greater than four times the breadth has been found in the
Central Provinces. The length of the grain of a long spikelet and
of a fine spikelet are greater than three times the breadth, of a coarse
spikelet greater than twice the breadth and of a round spikelet
less than twice the breadth. F

Here it may be mentioned that while rices are naturally bought
and sold on the character of the grain, a general preference is shown
for rices which have a narrow grain, but opinion differs with regard
to the length. In Jubbulpore a short narrow rice is esteemed,
while in Raipur and Chanda a fine, that is, a long narrow grain is
sought after.

Shape—In shape, also, the grain usually follows the spikelet,
though this is by no means invariable. Class I has a grain in which
both sides are slightly convex and the shape is ellipsoid. Class I
has a grain in which both sides are convex and the shape is broadly
oval. Class III has a grain which is nearly round. In class IV
the side of the grain next the glume is only slightly convex, while
that next the palea is convex. In class V the two sides are nearly

228 CLASSIFICATION OF RICE

symmetrical, both sides being parallel, or if the palea is concave, the .
grain is almost crescentic.

As already mentioned, the grain has ridges upon it which
correspond with the nerves of the glume and palea.

Double Grains.—In a rice from Bilaspur and another from
Narsinghpur the glume and palea are reported to enclose two or more
grains. This variety has already been referred to by Prain(22)
under the name of Oryza sativa var Plena. Like the coloured
rice and the rices with clustered spikelets this also ranks as a
systematic variety. In the present classification the distinction
between single and double grained rices cannot be made a main
distinction.

Abdominal White—This term is used by Kikkawa(23) to
describe the presence of a chalky white portion on the ventral side
of certain rice grains. When it occurs in the centre of the grain
surrounded by a translucent portion, the term ‘Shiratama’ is used
to distinguish the grain. Inagaki, quoted by Kikkawa, believes
that the abdominal white is nothing but a portion of the rice grain in
which the spaces between the starch grains have not been filled up
with albuminous substances. Most immature rices show an
abdominal white and it appears on germination in grains which
previously showed none. »

Micro-chemical examination of the abdominal white shows
that the starch grains are less closely packed in the cells in the chalky
white portion and that the cells contain a large amount of dextrin.
It would therefore seem that the abdominal white is an indication
that a grain is immature and that the carbohydrate has not been
converted into its most concentrated form. The appearance of
abdominal white in germinating grains is then due to the conversion
of starch into dextrin. It is obvious therefore that the use of the
presence or absence of closely related carbohydrates in a portion
of the endosperm as a basis for classification has nothing to recom-
mend it.

Notched Grain.—Gammie(24) mentions the presence of a notch
on the side of the grain as a diagnostic character in the rice varieties,

Her-
barium
=

No.

Name
lcu-

5.

Br.
Baniphul

Baisur
Baisur

B
_ Benibhog :

Bhakwa f

B
Benikat z

AWN.
| Sha pe.
| Length. | Colour.
| None ae WW
| 23” Red .. IV
[See leg oT

None ¥ 2 II
Apic os. i
None 4 I]

Her.
barium
No

2,2, 11

Name. District.
|
|
Baniphul | Raipur
|
|
Baisur | Raipur
Baisur | Raipur
Benibhog | Raipur
Bhakwa Drug

Benikat Raipur

| Dura.
| tion.

| M
|

Peduncle.

En.
En.
Ex,
Ex

Ex.

VEGETATIVE CHARACTERS.

Rachis.

(1) Lone Sprkecets.

(2) Coanse| SPIKELETS.

| Erect “|

Erect :
Curved and drooping | Approx.

Curved

Drooping

Separate curved
|

. | Separate drooping

Facets.

If—RICE WITH COLOURED LEAF SHEATH

2—ReD Grain

SPIKELET

GLUMES AND Paves.

| Glume THT and

| Empty Glumes as

Ordinary Comma... | Common Common
| |
| | |
>2 | Ordinary | Comma | Brown, base dark | Black
>2 | Membranous | Comma. | Purple black —.. | Purple dark brown
>2 | Ordinary | Elbow | Common Common
Soul Membrenpues eee | Comma Common | Common
>2 | Ordinary | Elbow .| Common ) Dark common

Apicu

lus.

P. Br.

Length.

None

Apie

None

Awn.

Colour.

Red
Red

Shape

Size.

>2

Colour:

Pink yellow.

Red
Red
Pale red
Red
Pale red

| Red

Embryo.

Yellow

Red

Dark yellow

Red

Yellow |

Apiculus
ing.

Apiculus
ing

REMARKS,

black at time of flower-

red at time of flower-

Apiculus red at time of flower-

ing
ulus

Apiculus
ing.

black at time of flower-

black at time of flower-

ALEA. AWN.
Her
barium ig =———=> z ‘
No. | Shape.
£& IIT and Apicu- Length. | Colour.
Palea. lus.
rie Bl. Apic . IV
2,1, 4| Badgi E
( Ae Br. Apic Pe ]
2, 2,16 | Bansjiral e
wy mone C None a I
2, 6,13 | Bhursi mon .. C Apiec He IV
2,7, 4) Bisanbhgo, _, » None oe I
3,1, 9) Chinga non .. b Apic = I
- 2 3 | Donapralt” Ay [2 ae None a IV

Her
barium
No.

Name.

Badgi
Bansjira IL

Bhursi
Bisanbhog
Chinga

Chitar Kote I
Donaprasad I

VEGETATIVE CHARACTERS.

I—RICE WITH GREEN LEAF SHEATH

2.—ReEpD Grain
(1) Coarse Spikelets

SPIKELET.

tt Dura-

District. tion y
Peduncle. | Rachis.

Bilaspur E Ex. | Ourved

Raipur M Ex. | Drooping

Raipur E En. | Erect oa

Bilaspur L Ex. | Curved oe

Chanda E En. | Curved and drooping

Raipur M | Much Ex. | Drooping

Raipur E Ex. | Erect, curved

Branches,

Lower branches se-
parate.
Separate, drooping

Curved
Approx. —

Separate

| Size.

Facets.

Ordinary
Ordinary

Ordinray
Thickened
Membranous
Ordinary

Membranons, ciliate

Richilla.

Comma
Comma

Comma

Flat comma
Elbow
Comma
Flat comma

Empty Glumes.

Common, base dark |

Light red

| Common

Common
Common
Common
Common

GLuMEs AND PALea.

| Glume IL and)

Palea
Black
| Common
|
Common

Common |
Common
Common
- | Common

Aws.
5 a l Shape.
Apicu- | Length. | Colour.
Jus.
|

Bi. Apie IV
Br. Apic !
C \
Cc Iv
c I
C :
P. Br. ny

Size,

GRAIN.

Colour,

Dark red .. |

Pale red ..

| Red

Red

Red
Red

Embryo.

Red
Yellow
Red

. | Red

| Dark red ..

Red
Orange red
Orange red

Fra-

grance:

REMARKS.

Ridges on glume light.

Apiculus black at time of
flowering.

Glumes suffused with brown.

Her- AWN.

LO I Sha
|
Apiculus. Length. | Colour.
|

(1) Foxe SPIKELETS.

2,2,9\| D. Br. | None a I
2, 3, 13 Br. | None a I
2,7, 6\| D. Br. Ap. 26 I
a2) 01. § 2. Br. Ap. <e J

(2) CoARsSE SPIKELETS.

as aed |e DE Br. None Il
11,7 || D- Br. 1/10” Brown .. I
2,1,13|| Red Ap. a I
251, 16 Ds Br; None A I
Oy | DS Br: és a ]
a Pee br. None a |

2,3, 14|| D. Br. None - |

I.—RICE WITH COLOURED LEAF SHEATH

I.—Warre Graw

VEGET SPIKELET. GRAIN.
Her- Giumes anp P. AWN,
a y, eee Dura- LUA AN 'ALEA, Awn, |
ie | Name. District. ae, | | Remarks,
Peduncle- Rachis. Branches. Shape. | Size. Facets. Rachilla, 7 Shape. | Size. | Colour. | Embryo. Fragrance,
| ' |
Empty Glumes. | Glumelifand | spicutus. | Length. | Colour. |
| | |
gh _ ere x —— ae = |S = [apes a | | |
| | | . =
(1) Foxe Spreevers. | |
2,2,9 | Banaspatri .. | Raipur eos Ex. Drooping .. | Separate, drooping | I >3 | Membranous . | Comma .. | Pale red .. | Common ..| D.Br. { None I >2 | Pale yellow . | Pale yellow ot ! black at time of
23,13 | F | | ng. Rachis dark brown.
2,3,13 | Basmoti .. | Chanda ae Ex. Curved ee Boos I >3 | Cil Comma .. | Common .. | Darkcommon .. Br. | None I >2 Pale yellow - | Dark yellow Apiculus black at time of flower-
7 | | ing.
%,7,6) Bobita .. | Raipur telat Ex. Curved ie ae I >3 | Thickened & ciliate | Comma .. | Brown .. | Dark brown D. Br. Ap. I >2 | Pale yellow . | Yellow Apiculus black at time of flower-
3211 ; 2 7 | | ing. Ligule black.
3,211 | ChitarChunilI ., Raipur 5 M Ex. Curved -. | Drooping 2 I >3 | Ciliate .. | Comma .. | Common .. | Common .| RB Br Ap. I >2 | Yellow A Yellow Apiculus black at time of flower-
| | ing. Ligule black.
(2) Coarse Srreecers. | |
1, 1,2) Ajan . | Raipur .| M Ex. Curved .. | Drooping aL. >2 | Ordinary Comma .. | Common -. | Common . | D. Br. None ; W& IV] >2 | Pale yellow | Pale yellow Glume with light brown furrows
tan : | culus black at time of flow-
11,7) Angormoti Raipur =. |f a3 Ex. Curved .. | Drooping Iv >2 | Ordinary .. | Comma .. | Common -. | Common +l Ds Bre 1/10” Brown ..] IV >2 | White . | Pale yellow ering. Ligule black. Apiculus
Peal : | | black at time of flowering.
21,13) Bahai Jira T .. | Raipur Bait | (a En. Erect .. | Separate elt a >2 | Ordinary . | Comma .. | Common -. | Brown Red Ap. J >2 | Yellow . | Yellow Glume suffused with black, Api-
9 : | | cults black at time of flowering,
21,16 | Bainssa Raipur 6) Ex. Curved .. | Drooping I >2 | Ciliate Comma .. | Common -. | Dark brown .. | D. Br, None 0 I >2 | Pale yellow . | Yellow Glumes with light: ridges, Api
907/| : | | culus black at time of flowering.
“*7 | Banaspatri . | Bhandara ote Ex. Curved . | Drooping a >2 | Ordinary Comma .. | Pale red -. | Common .. || D: Br. 3 I >2 | Pale yellow . | Yellow Apiculus black at time of dower
21] } | ing.
22,14! Bansbhirs Drug albany Ex. Curved . | Drooping Bie >2 | Ordinary .. | Comma .. | Common -. | Common .. | P. Bre None ; I 2 | Pale yellow... | Pale yellow Ligule red. Apiculus black at
3 time of flowering.
43,14 | Batripale . | Seoni seas) En. Curved .. | Drooping mi 2 | Slightly ciliate Elbow .. | Common -. | Common | D. Br. None ul <2 | Paleyellow — .. | Pale yellow Ligule red. Apiculus black at
aeal| | time of flow ‘
25,2 | Bhakuwa Raipur ae E Much ex. | Curved Drooping F it >2 Ordinary Comma .. | Common Common . Br. None I >2 | Pale yellow - | Yellow Ligule poe Apiculus black at
25,3| | time of flowering.
25,3 | Bhakwa Drug se ake Ex. Curved | Drooping I >2 | Ciliate .. | Shortcomma .. | Common -. | Darkcommon .. | D. Br. Ap. 5 I >2 | Yellow . | Yellow Apiculus black at time of flower
oe “ ing. s
2,5,7 | Bhas Telasi . | Chanda L En. Curved .. | Drooping || ae >2 | Membranous -. | Comma -- | Dark yellow .. | Purple brown .. | D. Br. None ; I >2 | Yellow .. | Yellow Ligule black. Apiculus red at
os time of flowering.
*5,8| BhataGurmatia .. | Drug .. | E | Much ex. | Curved .. | Drooping Af at >2 | Ordinary . | Comma .. | Common -. | Common .. | Brown None I >2 | Pale yellow . | Yellow 5 Ligule black Apiculus black at
5,9 o | | time 0 lowering. Ad
25,9 Bhataphul -. | Jubbulpore .. | EB Ex. Drooping . | Separate, drooping..| I >2 | Membranous - | Elbow .. | Brown -. | Common =a | Ds Br Ap. I | >2 | Pale yellow . | Yellow Rachis brown at the origin of
| | branches. Apiculus black at
| | | time of flowering.
2,0,2| Bheda Kabor -. | Raipur meu ann Ex. Curved .. | Drooping 5 I >2 | Ciliate .. | Flat comma Common -. | Darkcommon .. | Brown None a I | >2 | Pale yellow | Yellow r Bi Apiculus black at time of flower-
q g | | | - ing.
$2.3) Chipra .. | Bhandara .,| E Ex. | Curved ; < Iv. | >2 | Mombranous . | Comma | Common .. | Darkcommon .. | P, Br. None Iv. | >2 | Paleyellow .. | Yellow . | Slight eda
*2,5) Chipra Ching .. | Bhandara ..| © | Ex, | Curved .. | Drooping -| Iv | >2 | Ciliate .. | Blatcomma —.. | Common +. | Darkeommon .. | Brown | Ap. Iv | >2 | Yellow - | Dark yellow. | «+ | Apiculus black at time of flower
. E | | | | ing.
$,2,20 | Chitar Kote I Raipur -.| M | Muchex. | Drooping an Bree I >2 | Membranous Flatcomma .. | Common -. | Dark yellow s (eo Ap.or} Common ., | I | 2 | Dark yellow } Yellow oxo |H oR D |) Mn wetermie
short. | “ i
41,4) Door Sangar .. | Raipur .. | M_ | Much ex. | Curved .. | Separato, curved .. | I >2 | Ordinary . | Flatcomma .. | Common .. | Common D. Br. Ap. or I | >2 | Pale yellow . | Yellow | Glumes with rusty patches. Api-
41,11 | p} short. | | | cobs biec ety fuinelo# Howards
32 hanii i ch ex. ing | Se i 2 | Ciliate sale a5 | hie’ soe: : . y 2 | Yelle .. | Yellow Glumes with rusty patches, \pi-
Ss = -- | Raipur re L Much ex. | Curved and drooping | Separate, drooping IV > Ciliate Comma Common Common | Brown Ap. ci IV | > } Yellow | ellow nea ee Ai SBHEGEREE
m8) Donk .. | Raipur ..| M a Curved =e a m1 >2 | Membranons, ehortly | Elbow -- | Common -. | Common | Brown None z It | <2 | Paleyellow .. | Pale yellow 5 Rigula zed Ap ie black at
4,2 ; : ciliate, | sa Ke ‘time of flowering. ;
P14) DodkiTelusi —.. | Chanda aoa Rx. | Curved -. | Drooping Bike >2 | Membranous . | Comma +. | Dark brown — -. | Dark brown D. Br. None : I | >2 | Yellow Yellow .. | Ligule red. Glumes lighter above.
| | | Apiculus red at time of flower-
; | | ng. &
43,2] Dudhee -. | Chanda we EB Ex. Curved -. | Drooping Fi] tos >2 | Ciliate -. | Comma .. | Common -» | Dark common | Brown Ap. ele Iv | >2 | Pale yellow -. | Yellow oe] - Apiculus black at time of flower-
a | | } | ing. ;
43,3) Dudhia Sela .. | Raipur ..| M Ex. Drooping | sees I >2 | Ciliate -. | Flatcomma —.. | Common -. | Darkcommon .. | D. Br. Ap. 3 mi | >2 | Yellow 2 Yellow 5 a Ligule red ieee black at
| | ime of flowering.
410 Duaki .| Mondo = .. | L Ex. | Curved | see I >2 | Thickened ciliate .. | Comma -» | Dark brown —.. | Dark brown D. Br. None . I | >2 | Darkyellow .. | Dark yellow. OR OR ee teri
| (3) Rounp Spreever. | | lighter above.
41,3) Donkhi .. | Raipur a L Ex. Curved. . | seas it <2 | Ciliate -. | Flat comma -- | Common Common -. | Brown | None 5 Wt <2 | Pale yellow Ae | Pale yellow as cs prone black at time of flower-
a = =" a | ing. “

Her-
barium.
Number.

|
-
|
|

Palea. see Length. |
(@} 1}’ |
ORY Ap. |
C iy
a Fees
C a

I.—RICE WITH GREEN LEAF SHEATH

1.—Wuite Grain

7 | ie Rams... 0 wage

VEGETATIVE CHARACTERS.

7
fain. Name. |
Number Peduncle.| — Rachis.
|
|
1, 7 | Bagmochh M Ex. | Drooping
+7 2 | Bighli .. M Ex. | Drooping
113 Chinoor L Ex. | Curved
9,14 | Chitrakot E Ex. | Curved
43, 1| Dubraj L_ | Much ex. | Drooping
1, 1 | Anterbed En. | Curved
3.1, 8 | Bagmochh En. Drooping .. |
21/14 | Bahai Jira 11 Muchex. | Curved and |
2 | drooping. |
Basant Ex. | Curved .. |
Basant Ex. | Curved |
Basmatia Ex. | Drooping .. |
Basmatia En. Curved 3
Basmatio En. | Curved and
| drooping. |
Beni Much ex. | Curved «|
Bhari Ramkel | Ex. Curved >|
Bhejri | Much ex. | Curved 21)
6,10 Bhudo Bapu -| Ex. |
6,12 | Bhuri Ramkel : a0
3,1, 3 | Chilekar | Ex. |
3,1, 6 | Chilekath =| Much ex.
3,1,15 | Chinoor Ex.
4,1, 5 | Deppi Tokaloo En.
4,1, 6 | Dhamni Dhawool . Ex. Drooping
4,1, 7, Dhamni Dhawool . Much ex. | Curved
4,1,15 | Dhongad (red) Much ex, | Curved .
4,2,13 | Dongarsar Ex. Ereet, curved
|
|
1,1, 1| Ajan .. Ex. | Curved
1,1, 5 | Alchi Kabar Muchex. | Curved |
1,1, 8) Anjan Ex. | Curved :
1,1,12 | Anterbed Ex. | Curved
1,1,13 | Anterbed 5 En. Ascending ..
1,1,16 | Ari Motin : *s oe
21, 1) Bablapuri na an Oueved
21, 3) Badeli L Ex. Curved
31, 5 | Badlai ; .. | Muchex. | Curved
ae | Bahia Baikoni E Much ex. | Curved
oi,11 | Bohuja Chingar M Ex. Drooping
21,17 | Bairbuta E Ex. Drooping
23,4 | Barbi Sela E Ex. Curved
23,12 | Batro E Ex. Curved and
; | drooping.
Hi i a | Benara zr Ex. Curved
sate Bheda L Ex. Curved .
#8, 5 | Bhesara E Ex. Curved and
P di i
31,1 | Changar Ex. ja an
3.4, 7) Chilekath iF Ex. Curved
31, 8 | Chini Sakhar Ex. Curved and
P FS drooping.
3,1,11 | Chingori Ex. inane
32,11 | Chitar Chuni I i
4,8 se Much ex. | Di
3,2, 18 | CT Gar- Much fe Gievedeend
wallo, i
41, 8) Dhana L Ex. ES
42,1) Ditbay i
9 9 8a :
49 9 Dilbeleha 0 Much ex, Drooping
4,3, 5| Dugh Ex. Drooping
Cen lsc b khowa L_ | Muchex. | Curved
v) * | Gagli Mothi L_ | Much ex. | Curved
6 :
41, 6 | Gandur th | ma. || curvea
|
|
4,2,10 | ;
Fy 4 Dongar Manki BE Ex. Curved
42 ongar Manki M Ex. Curved :
12,12 Dongar Manki B Ex. | Curved and
4,3, = drooping.
» 4) Dudhiyar E Ex. Curved and

Branches.

Approx.

Drooping
Spreading
Drooping
Drooping

Drooping
Approx.
Drooping

Spreading
Drooping

Spreading
Approx.
Drooping

Drooping
Drooping

Drooping
Spreading
Drooping
Drooping

Drooping -

ing.

Drooping
Drooping
Drooping
Drooping

Drooping
Drooping

Drooping

Drooping
Drooping
Lower branches

drooping.
Curved

Drooping

Lower, drooping
separate,

Lower, drooping
separate.

Drooping

Drooping
Approx.
Approx.

drooping.

SHR

Drooping a
Approx, lower spread.

Bee

Slightly drooping . .

Separate, drooping 4

Drooping, separate .

Separate, drooping .

Separate, drooping .

Drooping, separate .

Facets, Rachilla.
> 4 | Ordinary Comma
> 4 | Thickened Short comma
> 4 | Membranous Elbow
> 4 | Membranous Comma
> 4 | Membranous Comma
> 3 | Ordinary Comma
> 3 | Ordinary Elbow
> 3 | Ordinary Comma
> 3 | Ordinary - | Comma
> 3 | Ordinary .. | Elbow
> 3 | Ordinary - | Elbow
>3 Membranous Comma
> 3 | Ordinary Comma
>3 Membranous Elbow
> 3 Ordinary | Comma
> 3 | Ordy, or sltly. ciliate | Comma
> 3 | Ordinary Comma :
> 3 | Ordinary Elbow -|
> 3 | Ordinary Short comma 5
> 3 | Membranous Comma - |
> 3 | Membranous - | Elbow
> 3 | Membranous - | Plat comma
> 3 | Ciliate - | Plat comma
> 3 | Thickened Comma
> 3 | Ordinary Comma
> 3 | Ordinary Flat comma
> 2 | Ordinary Comma |
> 2 | Ordinary Comma .
> 2 | Ordinary Comma
>93 Ordinary Comma
> 2 | Ordinary Elbow
> 2 | Ordinary Comma
> 2 | Ordinary Comma
> 2 | Ordinary Comma
> 2 | Ciliate Comma
> 2 | Ordinary Comma
> 2 | Ordinary Comma
> 2 | Ordinary Elbow
> 2 | Ordinary Flat comma
> 2 | Ciliate Elbow
> 2 | Ordinary Short comma
> 2 | Ordinary Comma
> 2 | Ordinary Comma
> 2 | Ciliate Short comma
> 2 | Ordinary Plat comma
> 2 | Thickened Elbow
> 2 | Ordinary Flat comma
> 2 | Ordinary Flat comma
> 2 | Thickened Plat comma
> 2 | Membranous Elbow
> 2 | Membranous Plat comma
> 2 | Membranous Plat comma
> 2 | Membranous Elbow
> 2 | Ciliate Plat comma
> 2 | Membranous Flat comma
<2 | Ordinary Elbow
< 2 | Ordinary Flat comma
< 2 | Thickened Elbow
<2 | Ordinary Comma

GuuMEs AND PaLea.

Empty Glumes.
|

(1) Lone

Common
Common

| Common

Common

| Common

(2) Fixe Sprkevers.

Common

.. | Common
.-| Black brown

Common
Common

Common

Common
Common

Common
Common
Common
Common
Common
Common
Common
Common
Common
Common
Common
Common
Common

(3) Coarse

Common
Common
Light brown
Common
Common
Common
Common
Common
Common
Common
Scarlet
Common
Common
Common
Common
Common
Common
Common
Common
Common

Common
Common

Brown

Common
Brown

Common
Common

Red brown
(4) Rounp.
Common
Common
Common

Common

SPIKELETS.

Glume ITT and Palea.

Common
Orang» yellow
Common
Common
Common

Dark yellow common
Common o
Dark brown

Common
Common
Common

Pale common
Common

Common
Common = ||
Common
Common 5
Dark common... |
Common :
Common
Common
Common
Light brown
Dark common
Brown

Dark common

SPrKeLets

Common
Common
Common

Dark yellow
Dark yellow
Common
Common
Common dark
Common
Dark common
Common
Common
Common
Common
Dark common
Dark brown
Common
Common
Dark common
Common, slightly
dark.
Common
Common
Dark common
Common
Brown
Dark common
Orange brown
Light brown
SPIKELETS.
Common

Orange yellow
Common

Common and darker

Colour,

| Common

Common
Commen
Common

Common

Common
Common

Dark common
Common

Light brown

Common
Dark common

Scarlet

Red

GRAIN.

wot

wen

me
ton

=

VVVVVVVVVVVVVY VV VV¥V VV
NO 8 b9 bo toes tone to ROR to

Hee ae eee

torr

AAKVVV VVV

torrets

bo pornos

w

NV AAVY

wren

t wo wo

AN V AV A VVV
toro to

“A
to

AN ANN V VAVY
to wr to porter

Pale yellow
Pale yellow
Pale yellow
Yellow
Yellow

Pale yellow
Pale yellow
Yellow

White
Pale yellow
Pale yellow

Pale yellow
Pale yellow

Pale yellow
Dark yellow
Pale yellow
Yellow
Pale yellow
Pale yellow
Yellow
Yellow
Yellow
Yellow
Pale yellow
Dark

Pale yellow
Pale yellow
Pale yellow

Pale yellow
Pale yellow
Pale yellow
Pale yellow
Pale yellow
Pale yellow
Pale yellow
Pale yellow
Pale yellow
Pale yellow
Yellow
Yellow
Yellow
Yellow
Yellow
Pale yellow
Yellow

Pale yellow

Pale yellow
Yellow

Pale yellow
Yellow
Yellow
Yellow

Yellow

Dark yellow

White
Yellow
Yellow

Pale yellow

Colour.

ellow
Dark yellow

REMARKS,

Tip of glume red at time of
flowering.

Dwarf plants, furrows on glumes
slightly dark.

Furrows on glumes slightly dark.

Spikelets clustered.
Furrows on glume dark.

Furrows on glume dark.

Furrows on glume slichtly dark.

Glumes with red patch.
Apiculus black at time of flower-
ing.

Ligule black.

Apiculus black at time of flower
ing.

Glume with dark furrows.

Clustered spikelets.

Furrows on glume patched with
orange. 4

Furrows on glume with orange

patches,

Apiculus black at time of flower-
ing.
Short rice. Apiculus black at
time of flowering.
Glumes with yellow brown fur-

rows.

Glumes with brown furrows.

Top of glume III dark brown,
apiculus black at time of
flowering.

Apiculus black at time of fower-
ing.

R. J. D. GRAHAM 929

This character is difficult to make out and presumably refers to the
depression left by the base of the style. In the rices of the Central
Provinces this notch is found to be present or absent in the grains
of the same head. It may be that the true notch has been overlooked
in the present examination of the Central Provinces rices, but at
present it is not possible to use this as a diagnostic character.

EMBRYO.

The embryo occupies an oblique position on the side of the grain
covered by the glume. The embryo is normally the same colour
as the grain, but the portion occupied by it differs from the rest of
the grain in not being translucent. In certain cases the embryo,
instead of being of the same colour as the grain, is distinctly dark,
in the white rices a dark yellow and in red rices orange.

Outline Classification.—The distinctions brought out in this
paper enable a comparatively simple classification to be made.

All rices fall into one of two groups, viz., rices with a green leaf
sheath and those with a coloured leaf sheath. The second class may
be sub-divided into those with a red leaf sheath and those with a
purple leaf sheath.

These classes further sub-divide on their vegetative characters,
those of the spikelet and those of the grain. In addition to these
morphological characters, the time of ripening, though not definite
enough to form a main point in the classification of rices from a large
area, is of considerable local importance. As already mentioned,
the special rices are fitted into the general scheme of classification.
They are so distinct that they may well be considered systematic
varieties.

DESCRIPTION OF PLATES.

Pruate I.

Principal shapes of the spikelet in Oryza sativa, Linn, x C7.

Fig. 1. Chitar Chuni (Raipur) ‘+ Type 1. Palea slightly convex, glume
slightly convex.
» 2. Mendha(Chanda) °° ‘+ Type 2. Palea convex, glume convex.
» & Donkhi (Raipur) ay ‘+ Type 3. Palea very convex, glume
very convex.
» 4. Selo (Raipur) ue ‘+ Type 4. Palea slightly convex, glume
convex.
»5 d+» Dhamni Dhawool (Chanda) ‘* Typed. Palea straight, glume slightly
convex,
Puarte II.
Principal shapes of rachilla in Oryza sativa, Linn, x C7.
», 1. Bainssa (Raipur) ie >> Comma.
» 2 Dhana (Jubbulpore) °° ‘+ Elbow. Outer glumes red, glume IT]

and palea common.
» 9 Chini Sakhar (Jubbulpore) ‘+ Hilbow.
The projection of the lateral nerves
of glume [II is distinct.

» 4. Deor Sagar (Raipur) ‘+ Flat comma.
Puate III.
Types of facets in Oryza sativa, Linn, x C10.
,, 1. Deor Sagar (Raipur) ‘+ Ordinary.
,, 2. Dhana (Jubbulpore) °- ‘+ Membranous.
., 3» Dudhia sela (Chanda) *: Cihate.
Plate IV,

Clustered spikelets in Bahia Baikoni (Raipur).

I,

PLATE

PRTC RPE revi:

.
d - = F + ee e eg
~~ * Pa = " % ten = =
¥ . - ot, - a a "a :
pt. ™ ~ sa * “ - a - . ‘ a - ~
. r 4 D Soe 4” * 4d <n 6 Te eae
aes > - - n _ pt
. ear . . ‘ , A * = re |
‘ * 4 = - a ls gm eA. ”
* = F x ot a .
J 7 - : — = e+ » «ft a4 a
viet : : =
‘ = ee > — ‘ ,
vis ¥ j = ae »
ic | “ + - - > +i : rd
a 2 . A oo és - r Sw
‘ . ¢ 5
a * c ~,
a ~ 7 - ~ .
= : a
. ~ ~
»' -
eu < - ‘
~ - = .
‘ - - . 7
ve = . 7 < ; P ‘ =e =e
< 4 mates ‘ sy
‘ "s “ .
, m) . =tp ve wae
. - = 7 x
. E a
y ‘ oF
' ~ ‘ ars fs
oa He » ‘
‘ < :
+* - ° .
; : Fie . dodetyt cite a?
ie of ach? ak ms Pens oan ia 5 aer-h
he ~ - x 3 ; iL ‘ She : }
. Be hy rn ued Auk flee dene) aes“ no Ot * t. ms uf had
’ 2. a x, re oe ag, Carey,"
y ve | Pe whe = - 4 : + O76 . .
> ‘3 . . + , fs - -
ya ‘ * : : ~ s 7 : “5
' _ - 2 7 “
+ We ce Pe ane - ee a . : ee
« ro ; : t d , Fee
! a Ed a . 4 :
r * cs i ~ Nd *
i .
‘
. |
‘
4:
La .
* sd 4
~ 1
.

Tl ALVW1d

PEATE SIM:

0

“|
<e.
nt +
'
‘ ‘
A
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. Koernicke
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S.
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S.

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LIST OF REFERENCES,

D. O. No. 400.

Mem. Dept.. Agri., India, Bot. Series, Vol. V1, No. 1.

Bengal Plants, p. 1184.
Dictionary of Economic Products, Vol. V, Part II
p- 501.

Jour. of Agri., Imp. Univ., Tokyo, Vol. III, No. 2.

Handbuch des Getreides, Band I, S. 232-242.
Ibid, p. 15.

Flora Indica, Vol. II, p. 203.

Ibid; Vol. V, Part Ul, p< 504,

Ibid, p. 18.

Address to Indian Congress, 1906.

Ibid, p. 21.

Organography of Plants, Vol. II, p. 377.
Bull. Dept., Agri., Bombay, No. 30, p. 89.
Ibid, p. 503.

Flora of Bombay Presidency, Vol. II, p. 1042.
Ibid, p. 89.

Ibid, p. 21.

Ibid, Vol. V, Part II, p. 503.

Ibid, p. 28.

Ibid, p. 22.

Ibid, p. 1184.

Ibid, p. 28.

Ibid, p. 90.

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LENO: een.

II, No. VI.
Ill, No. « I.
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III, No. IIT.
ELT No: DV.
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I, No. I
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I, No. Ill
To No; oY
fe SNossel¥
Hee koe MAG
II, No. i
11, No: “ait:

IJ, No. III.

Tee NGss vv
II, No. Ve

RiyeNo; | VT
Rio Vk.
DigiNo. WELT.

My No. xX:

PeNo, | 3s
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EY. No.) LU,
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IV; No: ‘IV.
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December 1914, BoTaNIcAL Srnaigs, Vor, VI, No. 8,

MEMOIRS OF THE
DEPARTMENT OF AGRICULTURE
IN INDIA

THE INFLUENCE OF THE ENVIRONMENT ON
THE MILLING AND BAKING QUALITIES
OF WHEAT IN INDIA

No. 3. THE EXPERIMENTS OF 1911-12

BY
ALBERT HOWARD, C.L.E., H. M. LEAKE, M.A
M.A Economic Botanist to the Government
Imperial Economic Botanist of the United Provinces

AND
GABRIELLE L. C. HOWARD, M.A

Personal Assistant to the Imperial Economie Botanist
Associate and former Fellow of Newnham College, Cambridge

TANICAL
GARDER.

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Students and

PREFACE.

THE experiments described in the present paper are a continua-
tion of those dealt with in two previous communications (Mem.
Dept. Agr. in India, Botanical Series, Vol. III, No. 4, 1910, and
WolkvV.. No. 2; 1913).

We desire to take this opportunity of expressing our indebted-
ness to several officers of the Indian Agricultural Department for
their valuable co-operation in the conduct of this work. For
facilities at Dumraon and Bankipore we are indebted to Mr. G.
Sherrard, Deputy Director of Agriculture, Bihar. Dr. Parr and
Mr. B. C. Burt, the Deputy Directors of Agriculture of the United
Provinces, have assisted us at Aligarh and Orai, respectively. At
Meerut, Babu Jagannath Pershad gave us all facilities on his farm.
Mr. Sharma has kindly placed the resources of the Partabgarh Ex-
periment Station at our disposal. In the Punjab, Mr. Roberts,
Professor of Agriculture at Lyallpur, and Mr. Southern, Deputy
Director at Gurdaspur, have been good enough to assist in the
work, while in Sind, Mr. Henderson very kindly assisted us at Mir-
purkhas. For facilities at Hoshangabad in the Narbada Valley
we are indebted to Mr. G. Evans. At Raipur and at Tharsa in the
Hastern Circle of the Central Provinces, Mr. D. Clouston, Deputy
Director of Agriculture, has given us very valuable help.

For the large number of nitrogen determinations involved
in the work we are indebted to Dr. J. W. Leather, Imperial Agri-
cultural Chemist.

In the milling and baking aspect of the subject we have been
fortunate enough to secure the invaluable assistance of Mr. A. E.
Humphries, formerly President of the National Association of
British and Irish Millers and a well-known authority on these ques-
tions.

ALBERT Howarb.

July 28th, 1914. H. Martin LEAKE.
GABRIELLE L. C. Howarp.

If
iil fe
Tt.

CONTENTS.
Introduction os “

The influence of environment on grain quality

Summary of conclusions .. ee

THE INFLUENCE OF THE ENVIRONMENT ON THE
MILLING AND BAKING QUALITIES OF
WHEAT IN INDIA.

No. 3. THE EXPERIMENTS OF 1911-12.

BY
ALBERT HOWARD, C.LE., H. M. LEAKE, M.A.,
M.A., Economic Botanist to the Government
Imperial Economic Botanist, of the United Provinces,
AND

GABRIELLE L. C. HOWARD, M.A,
Personal Assistant to the Imperial Economic Botanist,
Associate and former Fellow of Newnham College, Cambridge.

1. InrrRopvucrtron.

In the improvement of the wheat crop in India, the question
of the influence of the environment on the quality of the grain is a
matter of great importance. It affects not only the breeding of
improved wheats but also the question of seed distribution. It
has long been a vexed question as to what extent quality is deter-
mined by environment and how far it must be considered as charac-
teristic of the race. To the plant-breeder, who wishes to combine
in one strain the largest number of valuable qualities, a knowledge
of the respective parts played by breed and by environment in
producing and maintaining such qualities becomes essential. In
the general aspect of seed distribution in India, it is necessary to
know in what tracts wheats with high milling and baking qualities
are possible. For instance, at the present time, the black cotton
soils of the Peninsula and the canal irrigated tracts of Northern
India produce for the most part soft, weak wheats often with poor
milling qualities. One of the objects of this investigation is to
determine whether or not wheats of better quality can be grown
in these important areas,

LIBR 440 ¥
NEW YORE
BOTAN lGal

GARUGR,

234 ENVIRONMENT AND BAKING QUALITIES.

The characters of the wheat grain which may be affected by
change of environment are the following :—

1. Colour. While the general colour, red or white, of any
wheat remains the same no matter under what conditions it is
grown, nevertheless the depth or tone of colour in red or white
wheats is not constant. In India, white wheats, when well grown
under dry farming conditions, are frequently much darker in tint
than the same wheat grown carelessly under a superabundant
supply of canal irrigation. Similar differences are to be seen in
red wheat.

2. Size and weight of grain. The size and absolute weight of
the grain vary very considerably both in different localities in the
same year and also in the same locality in different seasons.

3. Composition. Much of the work on the effect of environ-
ment on the characters of the wheat grain has been concerned with
the effect of change of environment on the nitrogen content of the
grain—the nitrogen content being taken not only as a measure of
the percentage of gluten, but also as a rough indication of the strength
of a wheat. There are, however, exceptions to the general rule
that the higher the nitrogen the greater the strength so that, in
the present state of knowledge, the only safe method of estimating
strength is by milling and baking tests. Quality as well as quantity
of gluten is important in this respect. For a flour to be really
strong there must be sufficient gluten of the right quality present.
So far, while no accurate relation has hitherto been found between
chemical composition and the bread-making value of wheat, never-
theless the trend of recent investigations on this subject affords
hope that the strength of wheat may be explained from a chemical
standpoint. Thus Wood’ has found in the case of Fife and other
strong wheats that the water soluble phosphates in these flours is
high—over 0°1 per cent. and the chlorides and sulphates very low.
They also contain more magnesia than lime. Weak wheats, on the
other hand, yield flours with a low proportion of soluble phosphates,

1 See Jago—The Technology of Bread-making, London, 1911, p. 323,

—_ s+ —

HOWARD, LEAKE AND HOWARD. 235

a high percentage of chlorides and sulphates with more lime than
magnesia. Harvey and Wood’ have published a preliminary note
on a method of determining the baking strength of single ears of
wheat by taking advantage of the differences in opalescence which
exist between water extracts of various wheats.

4. Consistency. The effect of environment on the consistency
of the wheat grain, 2.¢., its translucent or starchy appearance, has
been perhaps more thoroughly investigated than any other aspect
of the question. There is no doubt that consistency depends very

largely on the soil, on the available moisture and on the nutrition

of the crop, and that, in many cases, great changes take place in this
character in any variety according to the climatic conditions under
which itis grown. From the miller’s point of view, the consistency
of the sample is of the highest importance and is one of the
factors in determining the value of wheat. Consistency is com-
mercially important in two ways. Firstly, it affects the process
of conditioning or the adjustment of water previous to grinding
—as arule translucent wheats take up more water than soft samples.
Secondly, translucent grains usually behave better than soft wheats
in the mill and are more free grinding, thus enabling separation of
bran from flour to be made with ease. As strong wheats are fre-
quently translucent, translucency is sometimes considered to be
an indication of strength, but this is not always the case as both
translucent weak wheats and mellow strong wheats occur. In
spite of these exceptions, however, the consistency of the grain
remains a very important factor in the commercial valuation of a
wheat.

Quality. Good quality in wheat flour has been defined by
Humphries’ as “‘the sum of excellence on several points ”’ and these
are five in number : (1) flavour; (2) colour of the flour ; (3) strength,
i.e., size and shape of loaf; (4) stability of dough; (5) yield of

1 Harvey & Wood, A Method of determining the Baking strength of single ears of Wheat,
British Association Reports, 1911, p. 597.

2 Humphries, Quality in Wheaten Flour—a paper read before the Joint Session of the
Chemistry, Botany, and Agriculture Sections of the British Association at Winnipeg, 1909,

236 ENVIRONMENT AND BAKING QUALITIES.

bread per sack of flour. It is obvious that information on such
points cannot be determined in any other way than by milling and
baking tests. In seeking information on the effect of environment
on the quality of the grain it is therefore essential to submit the
sample to a complete test in the mill and bakehouse.

The experimental investigation of this subject in India was
commenced in 1907 and the results obtained up to the harvest of
1911 have already been published.'. In the present paper an
account is given of the progress made during the wheat growing
season of 1911-12. 7

The results published in the last paper were summed up as
follows :—

“ Usually in India the consistency of a wheat varies greatly
according to the conditions under whichit is grown. Some trans-
lucent wheats however are affected to a much less extent than
others while a few soft wheats have always remained soft.

“Weak wheats like Muzaffarnagar can be improved to some
extent in millmg and baking qualities by cultivation but they have
not been made to behave like strong wheats.

“Strong wheats with good milling qualities have been found to
retain strength and milling qualities both under canal irrigation on
the alluvium and also on the black soils of Peninsular India. In
the future improvement of the wheats of these tracts, the question
of grain quality should receive particular attention.

“ Adverse factors, such as waterlogging and late cultivation
affect both the yield and quality of wheat in the plains of India.
In any particular wheat, the conditions which produce the highest
yield are those which also produce the best sample. In the same
wheat, high yield and high quality can be combined. To obtain the
greatest financial return for his labour the cultivator should grow

' Howard, Leake & Howard. Memoirs of the Dept. of Agr. in India (Bot. Series), Vol.
ILI, No. 4, 1910, p. 191, and Vol. V, No. 2, 1913, p. 49.

HOWARD, LEAKE AND HOWARD. 237

to perfection a wheat which combines high yielding power with
high quality.”

The earlier investigations carried out in Europe and North
America, which bear on this subject, have been restricted to a great
extent to the influence of external conditions on the composition
of wheat. This literature is referred toin the previous paper. Since
it was published one contribution to the subject has appeared which
is dealt with below.

In the United States, Le Clerc' has contimued his investiga-
tions on the influence of the environment on the composition of
wheat. The object of these experiments was to determine the
part played by soil on the one hand, and by climatic conditions on
the other hand, in bringing about the well known differences in the
appearance and composition of wheat caused by changes in the
environment. For this purpose, samples of soil were interchanged
among three localities—Maryland, Kansas, and California, which
differ widely in climatic conditions. From each locality, sections
of a normally fertile wheat-producing soil, five feet square and
three feet deep, were dug up in three-inch layers, sacked and re-
placed in the same original position. The various samples of
wheat grown were analysed and the results are set out in tabular
form. They indicate that climatic conditions, far more than the
soil, influence the composition and appearance of the wheat grain.
As in previous years, milling and baking tests have not been in-
cluded in the scheme and the conclusions are drawn from the analy-
tical data only. It is unfortunate that the various samples were not
milled and made into bread as these experiments would then have
been most useful in throwing light on the influence of climatic
conditions on milling and baking qualities. In order to obtain
accurate information on the effect of environment on the behaviour
of the same sample in the mill and subsequently in the bakehouse
it is, in the present state of knowledge, unsafe to rely on
chemical data only and on the appearance of the samples. Such
important matters as strength of flour and the free-milling nature

1 Le Clere and Yoder, Jour. of Agr. Research, Vol. 1, 1914, p. 276.

238 ENVIRONMENT AND BAKING QUALITIES.

of the wheats might easily be masked by changes in consistency
and nitrogen content.

The agricultural conditions under which wheat is grown in
India have been referred to in detail in the previous papers. There
are two great wheat tracts in India which differ widely from each
other, both as regards soil and as regards the source of moisture.
The more important of these regions is the alluvium of the Indo-
Gangetic plain, stretching from Bihar on the east through the
United Provinces and the Punjab to Sind on the western coast.
In parts of Bihar, wheat is grown on high moisture retaining loams
without irrigation. in Oudh, wells supplement the rainfall, while
in the western districts of the United Provinces, canal water is
commonly employed in wheat growing. In the Punjab, the crop
is largely watered from perennial and inundation canals while, in
Sind, inundation from the Indus takes the place of the monsoon.
The predominant features of the wheat tracts of the plains are the
alluvial character of the soil and the occurrence of some form of
irrigation. ‘The second great wheat growing tract in India is found
in the Peninsula on the black cotton soils of the Central Provinces
and Bombay. Here irrigation is the exception and most of the
wheat is grown on the moisture left in the soil after the previous
monsoon.

Besides moisture and soil, another factorin Indian wheat pro-
duction is of importance. Thisis the limited growth period. Wheat
can only be sown with safety as soon as the temperature falls suffi-
ciently for germination to take place and for the seedlings to deve-
lop. Any attempt to lengthen the growth period by early sowing
leads to the partial or entire destruction of the seedlings by heat.
The duration of the growth peried is equally limited by temperature
at harvest time. The Indian wheat crop ripens under a rapidly
ascending temperature, often accompanied by hot, dry winds. Any
late crops dry up rather than ripen and the rapid advance of the
hot season prevents the cultivation of late maturing wheats. The

HOWARD, LEAKE AND HOWARD. 239

growth period is shortest in Bombay and Central India and longest
in the Punjab and North-West Frontier Province.

The main directions in which Indian wheat can be improved
are two—yield and quality. The limited growth period, the fre-
quent shortness of the water-supply, and the fact that in many
tracts the moisture retaining capacity of the wheat soils is not
great, all indicate that moderate yielding wheats are likely to be
the most profitable to the grower over an average of seasons. Higher
yielding wheats can be grown with safety to a very limited extent in
places where the retentive power of the soilis considerable and where
the growth period is longer than the average. Compared with the
wheats of Western Europe, where the growth period is long, the
rainfall well distributed and the standard of agriculture far higher,
the wheats of India are only moderate yielders. Generally speak-
ing, the season is too short in India for the growth of such high
cropping wheats as those of France and England. As yield is
determined by the length of the growth period and the average water-
supply, the plant-breeder in increasing the amount of wheat grown
soon reaches the limit imposed by these conditions. In the im-
provement of the quality of Indian wheat, however, there is much
greater scope forthe breeder. In general, the wheats of the country
have poor grain qualities, both from the milling aspect and also
from the point of view of bread-making. Some Indian wheats do
not mill well while all those exported have a reputation for pro-

ducing weak flour.

The method adopted in this investigation has been to compare,
as regards consistency, absolute weight, nitrogen content, and mill-
ing and baking qualities, several pure lines, of widely different
quality, grown at various centres. The stations have been selected
so as to include as many as possible of the most important wheat
tracts of the Indo-Gangetic plain as well as a few places represen-
tative of the black cotton soils of Peninsular India. In the
present paper, the behaviour of Pusa 12 at a large number of sta-
tions is the chief subject dealt with.

II. Tue Inruvence or Environment on GRAIN
(QUALITY.

Six Pusa wheats were employed in the environment experiments
of 1911-12. Three of these (Nos. 4, 12, and 22) are selections and
three (Nos. 107, 108, and 110) are hybrids. All are white wheats
of good quality and would be described as strong, free milling sorts.

The fourteen stations at which these wheats were grown are
representative of the general agricultural conditions of the Indo-
Gangetic plain and of the soils of Central India. These stations,
which are indicated on the map at the end of this paper, were as
follows :—

STATIONS ON THE GANGETIC ALLUVIUM.

1. Pusa. This station is situated on the older alluvium in
North Bihar, a tract in which wheat is grown without irrigation
on high moisture retaining loams containing about 30°% of calcium
carbonate.

2. Partabgarh. This station represents the alluvial tract of
Oudh where wheat is chiefly grown on well-irrigation.

3. Cawwpore. This centre is typical of the wheat tract known
as the Middle Doab, where wheat is grown on strong alluvial loams
with canal irrigation.

4. Meerut and Aligarh, These are duplicate stations, typical
of the large alluvial canal irrigated tract known as the Upper Doab,
where wheat is an important cold weather crop. The agricultural
conditions at these twe centres closely resemble those of the neigh-
bouring Muzaffarnagar District.

HOWARD, LEAKE AND HOWARD. 241

STATIONS IN THE INDUS VALLEY.

6. Gurdaspur. This centre is situated in the submontane
tract of the Eastern Punjab, where wheat is largely grown as a
dry crop without irrigation.

7. Lyallpur. This station is typical of the canal irrigated
tract known as the Chenab Colony, which produces a large proportion
of the wheat exported from Karachi. Overwatering the wheat
crop is a common spectacle in the Chenab Colony where the general
standard of cultivation is below that practised by the best culti-
vators in the Kastern Punjab and in the Upper Doab.

8. Mirpurkhas. This station is situated in Sind where
wheat is largely grown on the moisture left after inundation from
the Indus. The wheat production of this area is likely to increase
very considerably when it is commanded by perennial canals.

BLAcK Soin STATIONS.

9. Raipur and Tharsa. These stations are situated in the
Kastern portion of the Central Provinces where wheat is grown
on the moisture left in the black soils after the monsoon.

11. Hoshangabad. At this centre, which is situated in the
Narbada Valley in the west of the Central Provinces, the black
soils of Central India are seen to perfection. Wheat is perhaps the
most important crop in this tract and irrigation is exceptional.

12. Ora. The agricultural conditions of this Bundelkhand
station resemble those of the Central Provinces mentioned above.
The soil is, however, not true black cotton soil and the wheat crop
round Orai is watered to some extent by canals.

SoutH BIHAR STATIONS.

13. Dumraon. This station is situated in South Bihar to the
south of the Ganges and outside the Gangetic alluvium. The soil
is sandy and possesses little moisture-retaming capacity. In
consequence, even with irrigation, the yields of grain and straw
are small.

242 ENVIRONMENT AND BAKING QUALITIES.

14. Bankipore. This centre is also to the south of the Ganges
where the soil is not true Gangetic alluvium. It is a dark, heavy,
moisture-retaining clay, not unlike some of the soils of Peninsular

India.

The soil and moisture conditions at these fourteen stations
vary greatly as well as the general standard of agriculture. It is
no exaggeration to say that these stations represent the entire
range as regards soils and agricultural practice in India at the
present time and every gradation between dry cultivation, in the
monsoon-fed areas, and canal irrigation, in tracts which would
otherwise be desert. The selection of these stations was purposely
made in order to decermine how a wheat with good grain qualities
would behave under such widely differing conditions. In the pre-
vious paper, tables were published giving the agricultural details
relating to the preparation for wheat, the seed rate in general use,
and the amount of water given after sowing. General information
relating to the production of the wheat crop can also be found in
Wheat in India.’

After the harvest of 1912, twenty-eight samples of wheat,
grown at the fourteen stations mentioned above, were selected for
complete milling and baking tests and Mr. Humphries’ report is
given in full below.

Report By Mr. A. E. Humpuries (Past PrRestpENT oF THE In-
CORPORATED NATIONAL ASSOCIATION OF BritisH AND IRISH
MILLERS) ON TWENTY-EIGHT SAMPLES OF WHEAT FROM THE
INDIAN CROP OF 1912 SENT TO ENGLAND IN 1913.

‘In continuation of similar work performed by me in several
previous seasons, I have examined, cleaned, conditioned and milled
separately twenty-eight sample lots of wheat sent me by Mr. Albert
Howard, Imperial Economic Botanist and Mr. H. Martin Leake,
Economic Botanist to the Government of the United Provinces,

' Howard & Howard, Wheat in India, 1909, pp. 1—46,

HOWARD, LEAKE AND HOWARD. 243

and I have baked the flour produced from each lot by at least three
different methods, so that I may be able to form a confident opinion
as to their respective commercial merits on the markets of the

United Kingdom.

In previous reports, I have discussed various technical details,
and, therefore, need not herein do more by way of preamble than
reiterate a few definitions. Conditioning is a term applied to
the adjustment of the physical condition of wheat, whereby an
optimum separation of branny husk from kernel can be made in
milling. Where Indian wheats are concerned, it includes an ad-
dition of water varying widely in degree according to the nature of
the variety. Some kinds are described as free-milling, because
the separation of such wheat in milling into its various commercial
constituent products can be effected easily, other kinds are
described by the expressive term woolly because separations in
milling are effected with great technical difficulty. The colour oi
the resulting flour depends to a considerable extent upon the facility
with which the necessary separations in milling can be effected.
The term strength, applied to flour, means its capacity for yield-
ing large shapely loaves. Stability indicates the facility with
which the baker can handle large masses of dough. Yield of bread,
which must not be confounded with strength, is the measure of
the quantity of bread which can be produced trom a given quantity
of flour. Good flavour implies a moistness and sweetness inthe
taste of bread at least one day old. The term good colour,
applied to flour, indicates its whiteness or brightness. A wheat
which is translucent may be strong, but translucency is not a true
index of strength. Nor is it correct to use hard and strong
as correlated terms, for some hard wheats are weak; some soit
wheats are strong. A wheat is said to be red or white ac-
cording to the colour of its skin. The term red really implies various
shades of brown; the term white various shades of yellow.
A red skin may cover an endosperm of good colour; a white
skin may cover an endosperm of poor colour. Many red
wheats are strong; most white wheats are weak, but

244 ENVIRONMENT AND BAKING QUALITIES.

colour of skinis one Mendelian unit, quality of endosperm 1s another ;
and a white wheat may be of maximum strength ; a red wheat may
be quite weak. The relative darkness of the medium and lower
grades of flour is due principally to the presence of comminuted
bran, and it is obvious that red bran particles will discolour flour
much more than those from a white bran. As a rule, white
wheats are more inclined to woolliness than red ones; but the
former can behave perfectly in milling, and the ideal wheat will

probably be a white one.

Wheats Received.
The wheat growing districts of India may be grouped as
follows :—
1. The Ganges Valley.
The Indus Valley.
3. The Black Soils of Central India.

I have received twenty-eight sample lots, and these can be

nh

grouped as follows :—

Ganges Valley. Indus Valley. Black Soits.

° Pusa 12 from Pusa. Gurdaspur. Raipur,
Bankipore. Lyallpur. Tharsa.
Dumraon. Mirpurkhas. Orai.

Partabgarh.,
Cawnpore.
Aligarh.
Meerut.

Pusa 22 from Pusa. Lyallpur.
Partabgarh.
Cawnpore.
Orai.
Aligarh.
Meerut.

Pusa 4 from Pusa. Tharsa,
Hoshangabad.

Pusa 107 from Pusa Tharsa.
», 108 Pusa. Tharsa.
oe LLU Pusa.

I have, in previous seasons, tested many varieties grown in
India and from among them have selected for commendation those

HOWARD, LEAKE AND HOWARD. 245

known as Pusa 4, Pusa 12, and Pusa 22. I am very pleased to see
that these varieties have now been grown at several places repre-
senting widely differing sets of natural conditions, so that Iam now
in a position to ascertain the effect of environment upon their
qualities.

Methods of Testing.

When a new kind of wheat is offered to a buyer, he forms an
opinion as to its merits upon its appearance, and he probably buys
the first lot with no better guide to its intrinsic worth than its good
or bad looks. Of course, an experienced buyer is better able to
appraise the real value of good looks than a beginner. To the latter,
a fine development and cleanliness may be all important: the
former has learned to know that a fine exterior may cover many
faults, and dirt, which can easily be removed, may nevertheless
obscure real beauty. Even so, the best judges know quite well
that their judgment in such a case may be faulty. Various methods
of rapid and accurate testing have been suggested. For instance
chewing has been recommended, and in certain cases for certain
points of quality that rough and ready test is valuable : but it is
useless or worse than useless in other cases, and obviously it would
not be used in the case I have pre-supposed, if the wheats were coated
with dirt of unknown origin. I need not labour the point that clean-
liness and a good appearance must in any and all cases materially
affect the price which a wheat will realize in our markets. But
there are other points which in most cases will militate against a new
kind of wheat at the outset of its commercial existence. A miller
has to learn how to treat it in conditioning and milling, so as to
secure optimum results, and he has to ascertain definitely not
only how it will behave in the bakehouse when used by itself
but how it will behave when blended with many other wheats
in various proportions. All this takes much time and trouble and
the inevitable mistakes cost money, so that a buyer is not likely
to pay a full price for the first few lots of a new kind of
wheat. But if he finds by many and various tests and by long

246 ENVIRONMENT AND BAKING QUALITIES.

experience that it has great intrinsic merits and can be relied
upon to vary in its qualities within small limits only, the reputa-
tion of the wheat will grow and its relative commercial value will
increase. So it may easily happen that a new wheat of great
intrinsic worth, arriving in unattractive guise from a district
which has hitherto produced wheat of poor quality, may realize
at the outset relatively poor prices on our markets and have to
overcome slowly the well or ill-founded prejudices due to its ap-
pearance, or even to its geographical source of origin, yet in time
the same wheat, even in the same unfortunate or deplorable guise,
may win its way to the real esteem of buyers, and command relatively
high prices in our markets. On the other hand, a wheat of
beautiful appearance, coming from likely quarters, may, in time,
because it is nondescript in quality, possessing no outstanding
merit of real importance, recede in relative commercial value
and leave the beginner or outsider wondering why merchants and
millers have such’ apparently curious ideas as to the value of
wheats.

In recent years, a good deal of work has been done by chemists
to ascertain the ultimate cause or causes of quality in wheat, but
it is true even now to say that they are unable to state with precision,
in terms of their own science, the characteristics of wheat which are
the ultimate causes of, or at any rate are correlated with, good
baking qualities. For present purposes, | have confined my en-
quiries, as tothe quality of the samples sent me, to their appearance,
and to their behaviour in the mill and bakehouse under commercial
conditions.

Cleaning. Almost all of the sample lots arrived in first rate
condition. The wheats from Lyallpur and Gurdaspur, however,
were dirty and Pusa 12 from Bankipore was very weevilly, so these
wheats had to be specially cleaned.

Conditioning. From previous experience, | was aware that to
obtain flour and bread of optimum quulity, some Indian wheats

HOWARD, LEAKE AND HOWARD. 247

had to contain after ‘ conditioning ’ a relatively low, others a high
or even a very high percentage of water. Although I bad in pre-
vious years tested these varieties, [ have made a few tests this year
to settle in my own mind how the samples should be condi-
tioned.

Baking. (have used the baking method with which I regularly
and continuously test flours for commercial purposes. The flour
from each of these Indian sample lots has been tested in at least
three ways :—

A. In this set, the flour obtained from each wheat indicated
was used without admixture of any other flour, and in making the
dough, flour, water, salt and yeast were the only ingredients.

B. This set was made in the same way as A except that in each
case a highly diastatic malt extract was added in making the dough,
in a proportion equal to 0°2 per cent. of the flour used.

C. Because British millers very rarely use Indian wheats alone,
I have in this set made a mixture of flours, using in each case one-
third of a straight run flour made from No. 2 Northern Manitoba
wheat of the 1912 crop, and two-thirds of the flour from each Indian
wheat indicated. 1 have fcr this set used the B baking method
except that the proportion of malt extract was reduced to 0°14 per
cent. of the total flour used in each case.

Pusa 12.

This may be described as a long berried, white wheat inclined
to be opaque. ‘The berries are not really large, but in most cases
are well developed and may be described as of good size.

The three groups of this wheat, judged by appearance, differ
from each other, principally as regards hue. Those from the Ganges
Valley and from the Black Soil Districts are of yellow hue, those from
the Indus Valley are different. It is difficult to describe the differ-
ence in words, but a miller wiil understand the phrase if I say they

248 ENVIRONMENT AND BAKING QUALITIES.

are of a white or yellow colour with a grey hue. When I was
examining the samples for the first time, I did not know the
geographical position of Gurdaspur or Mirpurkhas, and I was very
ereatly interested when I discovered that these placesare in the
Indus Valley. The hue to which I am referring appears, therefore, to
be characteristic of the district so far as this variety 1s concerned,
and it is a good point, for to me it is indicative of good quality. It
is very unfortunate that the Lyallpur sample, and, in lesser degree,
the Gurdaspur lot are so dirty. That blemish detracts very seriously
from their appearance, and it would militate against their commercial
value, especially when the wheat is first introduced to our markets
in commercial quantities. In a previous season, I received a sample
lot of wheat grown at Raipur, which possessed a most attractive
appearance and was in fact of very superior quality. This Pusa
12 sample grown at Raipur in 1912 has a very peculiar appearance.
It contains a few grains of a pink hue, and other grains which have
a black discoloration at the germ end of the berry. If that wheat
had been offered to me in commerce, | should have bought only
a small quantity at a low price, because its unusual appearance would
have left me in great doubt concerning its real worth. The Tharsa
wheat is smaller in berry than the average Pusa 12, but British
millers would not object to it for that reason. The remaining Black
Soil lot, Orai, is very pale and therefore unattractive in appearance

to millers who have to make strong flour.

In recent years, a great deal has been heard of strength in
wheat, and insome connections that point is of great importance,
but it is easy to exaggerate its importance in other connections. So
although anincrease inthe strength of Indian wheat is desirable,
it is far from being the only point of quality which buyers will take
into consideration. For instance, Pusa 12, grown at Orai and
Bankipore, has the appearance of weakness, but in other respects
are good, well-grown wheats. The Dumraon lot, although it contains
a considerable proportion of translucent grains, does not appear to
be strong, nevertheless it is a beautiful sample of wheat. 1 need
not attempt to describe the appearance of each lot in detail, but

HOWARD, LEAKE AND HOWARD. 249

taking into consideration all points of quality, I should place

them, according to appearance, in the following order :—
1. Aligarh.

. Meerut.

. Cawnpore.

. Dumraon.

. Partabgarh.

. Mirpurkhas.

. Pusa.

. Gurdaspur.

Oral.

. Tharsa.

11. Bankipore.

12. Lyallpur.

CAND oO — Ww WD

—
=)

From the foregoing list I have omitted Raipur, for the reason
already mentioned, but if I were compelled to place it, I would
bracket it equal to its sister lot Tharsa. Furthermore, when I
was examining the samples for the second time, I appraised them
differently. It then seemed to me desirable to put the grey hued
ones, Mirpurkhas, Gurdaspur, and Lyallpur into a separate group,
for I began to suspect that they were in fact of better quality than
their appearance at first sight would lead one toexpect. However.
as I am at this moment setting out a sequence of relative merit
according to appearance only, I have left the list as I first made it.
Accordingly, Lyallpur is certainly at the bottom. It is a dingy,
dirty looking lot, and the germ end of the berry appears to be swollen
in a way which one would expect to find if the earliest stages of ger-
mination had been passed. This small point of appearance is
characteristic of all Pusa 12 samples, but it is specially noticeable
in the Lyallpur sample.

In conditioning and milling, all these wheats behaved satis-
factorily except the Bankipore lot, and as that was badly weevilled
on its arrival in England, its inferior behaviour in the mill
is explained. The miller’s note on the Tharsa lot is that it “‘ milled

about the same as dry English.”” The lots from the Indus districts
2

250 ENVIRONMENT AND BAKING QUALITIES.

(Mirpurkhas, Lyallpur and Gurdaspur) behaved quite well at these
stages. After conditioning, they were more uniformly mellow than
the Ganges Valley lots, but they required the same percentage of
water as those to yield optimum separations.

The baking trials provided an interesting surprise ; for by each
of the methods described hereinbefore as A, B and C, the Indus
Valley lots showed a substantial superiority in strength and stability
over the others, and the Black Soil lots a slight but still appreciable
superiority in strength over those from the Ganges Valley. Having
regard to my previous experience, it was no surprise to find that
even the last named were distinctly superior in these respects to
the ordinary Indian wheats of commerce, Karachi, Calcutta, or
Bombay. Of course, it must be remarked that I have been handling
in these trials small quantities of wheat on an experimental scale,
but as the results set out in the following tables are in conformity
throughout with this verdict, my opinion as to stability and strength
is a confident one, but because it is impossible to obtain optimum
results as regards colour of flour and bread in testing such small
lots, 1 do not want to say much concerning that point of quality,
but it was obvious that, in this respect also, the real worth of the
Lyallpur lot is at variance with its appearance, for the colour of the
crumb was very good, fully equal to any other of the series and
superior to most of them.

The following tables will best put on record the results obtained

in the bakehouse as to ‘ stability’ and ‘ strength’ of the thirteen
lots of Pusa 12.

It has for many years seemed to me easier to record one’s
opinion in marks than in words, and a long and varied experience
has enabled us to express ourselves in this way with facility, but it
must be understood that the figures do nothing more than express
the baker’s and my own opinion of the relative merits of the samples
concerned on the points of quality specified. We may sometimes
aid our judgment by measuring some of the loaves, but essentially
the markings are based on the impression conveyed to our mind

HOWARD, LEAKE AND HOWARD. 251

by the senses of touch and sight : the method by which bakers in
commerce almost invariably judge the flours they use. Indeed,
I would like to emphasize the point, that I have deliberately chosen
to test, according to commercial methods, all the Indian wheats
I have handled on an experimental scale, for it seems to me of great
importance to apply, even to these small lots, the methods which
will be used in commerce in handling shiploads ultimately. Cer-
tain forms of laboratory work may assist the miller and baker to
obtain optimum results in the mill and bakehouse, and I am very
keen on seeking to apply such knowledge to commerce, but in the
present state of knowledge such work may sometimes mislead, and
for the purposes now in view, I have carefully applied well-known
commercial methods and based my judgment upon them exclusively.

Ganges Valley.

STABILITY. STRENGTH.

Baking method. Baking method.

A. B. (Of Average. A. B. | GC: Average.

Cawnpore 80 84 85 83 84
Pusa 82 88 88 86 83
Aligarh 80 84 85 83 83
Meerut 80 84. 85 83 83
Bankipore 78 84 86 83 82
Partabgarh 80 84 |° 82 82 81
Dumraon 78 82 84. 81 78

AVERAGE 80 84 85 83 82

Black Soils.

Eel

STABILITY. STRENGTH.

Baking method. Baking method.

| i
|

A. B. C. Average. | A. B. C. | Average.

| | | |

Tharsa 80 88 85 84 84 86 89 86
Raipur 78 84 88 83 75 82 92 83
Orai 78 84 88 83 76 80 84 80

AVERAGE

252 ENVIRONMENT AND BAKING QUALITIES.

Indus Valley.

STABILITY. STRENGTH.
~ Baking method. Baking method.
A. B. C. Average. : B. C. Average.
Mirpurkhas 82 86 86 85 é 90 92 89
Lyallpur | 82 88 88 86 86 92 88
Gurdaspur 78 86 88 84 85 88 86
AVERAGE 81 87 87 85 91 91 88

The flour from the ordinary Indian wheat of commerce, Choice
White Karachi, baked by itself would, according to the scale
used, obtain, say, 70 marks for stability and 70 for strength ; No. 2
Northern Manitoba, 1912 crop, would get, say, 92 and 95 respectively,
so my opinion as to the relative merits of these wheats as regards
stability and strength may be stated in the following summary :—

Stability. Strength.
No. 2 Northern Manitoba, 1912 crop... ae 92 95
Pusa 12, Indus Valley Ee St oe 87 87
Pusa 12, Black Soils AC = ae 85 83
Pusa 12, Ganges Valley oe x bed 84 82
Choice White Karachi ue we ma 70 70

It will be seen that in this summary, I have used the figures
concerning the Pusa 12 wheats, from the B set of trials.

Before | make any comments on these baking results, I should
like to put side by side the two sequences of merit relating to these
Pusa 12 wheats, one based on appearance, in which all points of
quality are taken into consideration, the other based on baking
trials in which the only points of quality considered are stability and

HOWARD, LEAKE AND HOWARD. 253

strength. For the latter sequence, the marks based on the three
sets of baking trials A, B and C are used.

Appearance. Strength & Stability.
Aligarh. Lyallpur.
Meerut. Mirpurkhas.
Cawnpore. Gurdaspur.
Dumraon. Tharsa.
Partabgarh. Cawnpore.
Mirpurkhas. Pusa.

Pusa. Aligarh.
Gurdaspur. Meerut.
Orai. Raipur.
Tharsa. Bankipore.
Raipur. Partabgarh.
Bankipore. Orai.
Lyallpur. Dumraon.

Although, of course, I did not expect, having regard to what
they purport to be, to get the same order of merit in both cases, the
differences between the two sequences are striking, and I have made
some enquiries with a view to ascertaining the causes or explana-
tion of the discrepancies.

The most extraordinary difference concerns the Lyallpur lot,
certainly at the bottom of the list when judged by appearance,
at the top so far as strength and stability are concerned. I have
already said that this lot as regards colour of “ crumb” was very
good, fully equal to any other of the series and superior to most of
them, but on this point of quality, I have this further important
remark to make. Bread produced from Manitoba wheat has at its
best a beautiful grey white colour, that produced from Choice White
Karachi is intensely yellow, and relatively bad. The Pusa 12
from Lyallpur yielded bread similar to the Manitoba in this respect.
Here, therefore, is a case to which my remarks under the heading
“* Methods of Testing” apply. No miller would buy the first lot,
or few first lots of it, except at a low price ; he might well hesitate
to spend time and trouble in making trials to ascertain its intrinsic
merits. Surely something can be done to improve its looks.

Another striking result is the position of the Indus Valley
wheats in the second sequence. I shall be very interested in seeing
in later seasons how other varieties behave when tested in a similar

254 ENVIRONMENT AND BAKING QUALITIES.

way. for present purposes, it is sufficient to say, that so far as
quality is concerned, Pusa 12 seems to suit the Indus Valley. If this
wheat be extensively growninthat district, it will probably command
sooner or later higher prices in our markets than the Choice White
and Red Karachi now so largely used in this country. If its
yield per acre of grain and straw be no higher than that obtained
from the varieties now commonly grown in the district, the relative
increase in value per quarter should commend Pusa 12 to growers ;
if it will also yield increased crops of grain and straw on an average
of seasons, it should replace existing varieties within a few seasons ;
but it is only right to add that no likely increase in the price per
quarter will compensate for a substantially diminished yield. I
shall therefore be much interested in seeing the returns as to yield
of grain and straw.

The Dumraon lot was particularly well developed, and might
be described as a really beautiful sample. Nevertheless, it was
relatively ‘weak.’ I find on enquiry that it was grown on over-
irrigated sandy loam, which might account for the baking result.
It is necessary, however, to point out that this lot, though it is at
the bottom of the list as regards bakehouse results, is nevertheless
much better than the ordinary Indian wheat of commerce (Karachi,
Calcutta or Bombay) as regards stability and strength. In this con-
nection | should like to say that the Bankipore lot is stronger than
its looks and record would lead one to expect. I understand that
it was grown after rice, on land which had been over-irrigated for
that crop, and it seems that so far as strength is concerned, it was
made to appear worse than it is by that treatment.

It will be seen from the tables that the ‘ strongest ’ earned
89 and the ‘weakest’ 78 marks for strength. I am_ surprised
that the difference is not greater. Some years ago, we grew wheat
from the same seed on several typical soils in England, and found the
differences greater than those with which we are now dealing.
However, I have made enquiries to ascertain whether any of the
districts represented in these trials are unlikely to grow wheat for

‘SLVJHM NVIGNI GNV NVIGYNVO WOdd SHAVO'T

‘IHOVUVS ALLY AA AOLOH,) ‘(aMadsvVauns)) ZL vsodg ((NUBHLUON % ‘ON) VHOLINVIK

HOWARD, LEAKE AND HOWARD. 255

export, and I find that Bankipore and Dumraon are in that category
and that Tharsa, Raipur and Orai, if they furnish any at all for that
purpose, are likely to provide only relatively small quantities. It,
therefore, appears from my information that the likely exporters
of substantial quantities are as follows. Alongside each name, I
have placed its marks for ‘strength’ (average of A, B, C methods

of baking).

By Karachi. By Calcutta,
Lyallpur is So, the) Pusa os BA) ee
Mirpurkhas .. sp GY) Partabgarh .. o- 81
Gurdaspur... sq tele
Cawnpore 3¢ .. 84
Aligarh as oS
Meerut 36 22) oo

If the growers and merchants of the Indus Valley will send their
wheats to our markets in a cleaner state than the samples I have
received from Lyallpur and Gurdaspur, I see no reason why, for
commercial purposes, the produce from the districts named in the
Karachi list should be sold here separately, but if the wheat
from the Punjab, when shipped on a commercial scale, will be as
unsightly as the two samples named, the produce from the other
districts should be marketed separately. The former will require
much time to establish the reputation they deserve in this country,
but for the latter with their fine appearance, enhanced prices may
be expected on their first arrival.

It is obvious that there is no reason for marketing the Pusa
and Partabgarh wheats separately.

In Bulletin 22, issued in 1911 by the Agricultural Research
Institute, Pusa, a coloured photograph of three loaves baked by me
was included, and I was asked to provide a similar photograph
showing a loaf made from the Lyallpur lot of Pusa 12, alongside a
loaf produced from No. 2 Northern Manitoba and another from
Choice White Karachi. I had made several trials with this Lyall-
pur wheat, and, in doing so, had used up the whole of the lot sent
me, so as the Gurdaspur wheat, of which I had some left, was similar
though slightly inferior to the Lyallpur, 1 made a further test and
had the photograph taken which is reproduced herewith. It was

256 ENVIRONMENT AND BAKING QUALITIES.

coloured by hand to show the characteristic differences in the colour
of the crusts. I need hardly say that the better coloured crusts are
not only thinner and better in all respects than the grey one, but
they give the loaves a much more appetising appearance.

Pusa 22.

Of this variety, I received sample lots from seven places. It isa
round berried wheat of attractive appearance, similar to Canadian
Fife in grain shape and in its behaviour in the mill. This in its
category is high praise, for Fife whether it be grown in Canada,
England or Germany, or whether it is represented by its Australian
progeny Comeback, or its English child Burgoyne’s Fife, is ideal in
its behaviour in the mill. To get optimum results, I found it desir-
able to raise the water content of these samples to a high figure.
In this connection, I would like to remark that the use of water in
wheat conditioning, has been reduced to a science. To obtain
successful results, close attention must be paid to details, more
particularly as to quantity and time, and amateur efforts in apply-
ing water to wheat may be as harmful to it, as the unskilled use of
water at spas may be injurious to the human system.

| append, in this case also, the two sequences, one constructed
on appearance only for all points of quality, the other on baking
results for stability and strength only. In these cases, I found that
the use of diastatic malt extract did very little good, so my opinion
as to baking results is based on trials made with flour, water, salt
and yeast only, in other words, the method known as ‘ A’ in the
Pusa 12 trials is the only one referred to in this connection.

Appearance. Stability & Strength.
Pusa. Lyallpur ae .. 84
Aligarh. Pusa oF .. 84
Cawnpore. Meerut aE .. 84
Meerut. Cawnpore 5c sin Da
Partabgarh. Partabgarh se «ot 4O
Orai. Orai aie aa
Lyallpur. Aligarh ei ee

Here we have the same striking difference concerning Lyallpur
and the remarks I made on that point in the case of Pusa 12 apply

HOWARD, LEAKE AND HOWARD. 257

here also. The only other striking difference in these two series
concerns the Aligarh lot. It certainly was weevilled on its arrival
here, and that may be the indirect cause of a falling off in baking
value, but apart from that, our work on English wheat has shown
that a given soil may grow a relatively strong sample of one variety
and under identical conditions a relatively weak sample of another
variety, which simply means in effect, that we have to provide for
each environment a variety or varieties inthe highest degree suitable
forit. Lunderstand, however, that mainly for agricultural reasons,
this wheat is not likely to be extensively grown, and I may, there-
fore, dismiss it with these few remarks.

The Pusa lot is of particularly attractive appearance and the
Orai wheat though very pale is nevertheless hard.

Pusa 4.

-I am at present particularly interested in this variety because
it has got beyond the experimental stage and I have received a
small sample of a large quantity grown in 1913 on a commercial
scale in Bihar.' I have shown this sample to several millers and
merchants, who all, with one consent, say it is very fine looking
wheat. One miller, who tested it by a comparatively new but
secret laboratory method, praised it very highly, and said such
wheat “ will never go begging.” Some of a cautious turn of mind,
well acquainted with ordinary Indian wheats, refrain from
expressing an opinion concerning its real merits until they have
tested it under commercial conditions.

I have received for milling and baking tests three sample lots
grown in 1912, respectively, at Pusa, Tharsa and Hoshangabad.
The last named is the best of the three, judged by appearance and
is very fine wheat beautifully grown. The Pusa sample has the
blemish of a black spot on the germ end of the berry which it had in
previous years, and in addition this year, a similar black spot in the
crease at the opposite end of the berry. Thisis not a very detri-

1 This sample was grown on the Hathowrie Estate in the Darbhangha District.

258 ENVIRONMENT AND BAKING QUALITIES.

mental fault, but if it can be removed it will be an improvement.
Apart from that, this lot is not so well-grown as the same wheat
raised in previous seasons at Pusa. The Tharsa lot contains some
unsound grains. It lacks brightness and appears to have been
substantially damaged by unfavourable climatic conditions. How-
ever, all three lots behaved very wellin the bakehouse, and in the
markings for stability and strength were quite close together. The
flours from the Hoshangabad and Tharsa lots behaved like typical
London milled flours, the Pusa lot was tougher inthe dough than the
others, and to some extent resembled the flours made from American

or Canadian spring wheat.
Pusa Nursery Lots.

In addition to the foregoing, I received the following sample
lots ; two lots of Pusa 107, one grown at Pusa, the other at Tharsa ;
two lots of Pusa 108, one from each of these places, and one lot of
Pusa 110 grown at Pusa. Pusa 108 is a round berried wheat.
The grains of the Pusa lot are small in size and those of the Tharsa
lot still smaller, but that is not a serious fault. The formeris wholly
hard and wholly translucent, a very fine looking lot. The latter
is not so bright or so translucent, and appears to have suffered from
bad weather. Both behaved very well inthe bakehouse and yielded
bread of beautiful appearance.

Pusa 107 is a much larger berried wheat, quite as large as Pusa
12. This variety as grown at Pusa may also be described as wholly
hard and wholly translucent, is particularly well-grown, and is in
fact a lovely sample of wheat. The Tharsa lot also appears to have
suffered from bad weather. In this case the relative inferiority in
appearance of the Tharsa lot corresponds with the baking result,
but the bread from it was good, that from the Pusa lot very good.

Pusa 110 is a long berried grey white wheat, is almost wholly
translucent and quite hard. This lot (from Pusa) is a very nice
sample of wheat, although the blemish of the dark spot found in
Pusa 4 occurs in this variety also. It behaves quite well in the

bakehouse.

HOWARD, LEAKE AND HOWARD. 259

Wheats grown at Pusa.

I have this year received in all sample lots of six varieties
grown at Pusa, and having regard to their appearance and to the
bakehouse results, I should place them in the following order of
merit :—

Pusa 108)
” +
” 170 -very close together.
pr alu
» 1

5)
99 level.

They are all fine wheats, and there is no great difference
between the best and the worst of them.

Wheats from Tharsa.

I placethe four lots from Tharsa in the following order of merit :—

Pusa 12
LOT
ste
», 108

but there is very little difference between the best and the worst of

them.
Summary.

It has again been demonstrated that wheats of the highest
class can be grown in India on several kinds of soil, and on land
which has beenirrigated. It has been shown that the high excellence
of quality possessed by wheats grown at Pusa in previous seasons
was not due to environment or agricultural practice, for the same
varieties of wheat have yielded still better results elsewhere. It is
interesting to note that this high excellence of quality was found
existing in wheats indigenous to India, and that inthe Pusa Nursery
varieties, the progeny appear to possess intact the great strength
of the strong parents. I have no doubt that any or all of the wheats
tested will realize, some at once, some later, relatively higher prices
on our markets thanthe existing Indian wheats of commerce. If
these new varieties yield no more grain and straw per acre than those
ordinarily grown, their extended distribution as seed is desirable ;

260 ENVIRONMENT AND BAKING QUALITIES.

if, in addition, the new varieties will yield greater quantities of grain
and straw than those ordinarily grown, the position of the Indian
grower will be greatly improved, and the propagation of the new
kinds should be pressed forward.”

WEYBRIDGE, ENGLAND,
A. K. HUMPHRIES.
3rd October, 1913.

Some exceedingly interesting results are to be found in the above
report which it is now proposed to deal with briefly. The first
relates to the importance of milling and baking tests in environment
experiments with wheat. In the case of the Pusa 12 sample grown
at Lyallpur, the report shows that a miller of great experience, who
had tested this variety in previous years, was entirely misled by the
external appearance of the badly grown and harvested sample.
Great flour strength and good milling qualities are therefore lable
to be masked by the effects of poor cultivation and overwatering.
This experience with the Lyallpur sample is of the greatest
value as it shows that if the Indian cultivator 1s to obtain immedi-
ately the greatest financial return for his labour he must not only
grow a wheat with good quality but this wheat must be well-grown
so that it at once takes the eye of the buyer. The appearance
of the sample is therefore a most important matter in the work of
introducing to advantage a new grade of wheat on the Home markets,

The general results obtained with Pusa 12 at the thirteen sta-
tions confirm and extend those obtained in 1910-11. These are
summed up in Table 1, which also gives the consistency, absolute
weight and nitrogen percentage of the various samples. It will
be seen that the nitrogen percentage of Pusa 12 is, on the whole,
not very high. The actual tests bring out the point that no matter
what the agricultural conditions were, the milling and baking

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262 ENVIRONMENT AND BAKING QUALITIES.

qualities were little influenced by environment. Flour strength and
milling qualities were not in the least destroyed by transferring a dry
crop wheat raised at Pusa in the extreme east of the wheat-growing
area of India to North-Western India, or to the black soils of Central
India. Indeed, the wheat was improved by this treatment, as the
best results were obtained at Lyallpur, Mirpurkhas, and Gurdaspur,
in the Indus Valley and at Tharsa in Central India.

A single grade of white wheat of improved quality can therefore
be grown over a very large part of the wheat producing area of India.
If the yield of grain and of bhusa (straw) of this wheat under culti-
vators’ conditions is satisfactory over these areas, then a great step
in advance will have been made. That this step has been accom-
plished will be evident when it is remembered that this wheat is now
being distributed to cultivators in the Punjub, United Provinces,
Bihar, and Central Provinces. Over avery wide range of. condi-
tions and of soils, both under dry cultivation and with canal and well
irrigation, this wheat has done uniformly better than the local sorts
when grown by the cultivators themselves. It is now only a matter
of time and of organization for this variety to replace the existing
crop over large tracts and for its produce to influence the Indian
wheat trade. Besides its good yielding power, Pusa 12 stands
out inthe field from the country wheats asared chaffed, beardless
wheat with strong straw among bearded white chafted kinds mostly
characterized by very weak straw. This fact is of considerable
advantage inthe work of replacing the country wheats by the new
variety.

The milling and baking results obtained with the samples of
Pusa 12 grown in the Indus Valley and also on the black soils prove
that there is no longer any reason why grain quality should be ignored
in the work of improving the wheats grown in these two areas.
In the first place, the performance of this wheat when grown by
cultivators shows that yield and quality can be combined in the same
variety. In the second place, these areas, which now produce only
poor quality wheats, have been shown to be capable of growing
samples of the same class as the best grades of Manitoba wheat.

HOWARD, LEAKE AND HOWARD. 263

It has been demonstrated in previous papers that all the Punjab
wheats tested up to the present are of poor quality and of the same
class as the grade known as Choice White Karachi. In like manner,
the white pissi wheats of the black cotton soils which have been
tested are far inferior to Pusa 12. From the results obtained by the
cultivators with this wheat in the Eastern Circle of the Central
Provinces, it 1s exceedingly likely that it will replace the wheats now
grown over large areas of the Central Provinces. To be conclusive,
the trials will have to be made under actual cultivators’ conditions
and on a sufficiently large scale.

In the case of Pusa 22, the results of the last two years are
summed upin Table 2. This wheat was selected from the local Bihar
mixtures and in good years yields very fine looking samples. Its
weak straw and liability to rust will prevent its ever being taken
up by cultivators.

The results of the tests of the other Pusa wheats are also of
interest. The details relating to the consistency, absolute weight
and nitrogen percentage are given in Table 3. Pusa 4 is a white
wheat with good straw which is exceedingly early and which can
ripen an attractive looking sample on the minimum of soil moisture.
For this reason it is being widely distributed to cultivatorsin Bundel-
khand where the soil often contains too little moisture to ripen the
local wheats. In Bihar, this variety, on account of its rapid growth,
strong straw and limited foliage has proved a suitable cover crop
for Java indigo, and it is hoped to establish this wheat in North
Bihar in sufficient quantities for export as an improved grade. The
results of the tests of Nos. 107, 108 and 110 are of interest as they
show that in the work of breeding improved varieties it is possible
to transmit unimpaired the high quality of one parent. These
three types were obtained by crossing Pusa 6 and Muzaffarnagar.
In the case of No. 110, an improved Muzaffarnagar has resulted
with stronger straw, red chaff and a high quality grain. Where an
improved bearded wheat is required on the Gangetic alluvium
this variety is well worth a trial.

QUALITIES.

AND BAKING

EN VIRONMENT

264

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HOWARD, LEAKE AND HOWARD.

TABLE 3.

265

Comparison between Pusa 4, 107, 108, 110, grown at Pusa and in the
Central Provinces, 1910-11 and 1911-12.

nn ssssssssssssssssssssssssee

Pusa

”

9

Weight of
1,000 grains
in grammes.

CONSISTENCY.
Name of wheat and 1910-11. 1911-12.
locality.
ee Ae
| : E : : E 1910-11,

4, Pusa Oo 491i 3 53 441] 45°67

4, Raipur 0 50 50|;— —~— —] 45°30

4, Hoshangabad — — —|]1 40 59 —

4, Tharsa —_—- — — | 12 69 19 —
108, Pusa —- — — 0 ee 98h
108, Tharsa —- — — A 2 24 —
107, Pusa —- —- — 0 4 96 _—
107, Tharsa ———) — jAlG 51 33)
110, Pusa _—_- —- — 0 15 85 —

1911-12.

Nitrogen
percentage.

!

1910-11.) 1911-12.

III. Summary or Concuvusions.

The conclusions arrived at as a result of the investigations
described in this paper may be summed up as follows :—

1. Pusa 12, a large grained, white wheat, grown at thirteen
stations on the Indo-Gangetic alluvium and on the black soils of
Peninsular India under widely differing conditions as regards soil,
available moisture, and agricultural practice has maintained its
high milling and baking qualities in all cases even under unfavour-
able conditions. It behaved in the mill as a free-milling wheat and
yielded strong flour and high grade loaves.

2. The best results with Pusa 12 were given by the samples
from the Indus Valley, the second best by those from the black
cotton soil stations followed closely by those grown at Pusa and
other stations on the Gangetic alluvium.

3. Inthe case of Pusa 4, another white wheat with good grain
qualities, the samples grown at Pusa, Tharsa, and Hoshangabad
behaved very wellin the bakehouse and in the markings for stability
and strength were quite close together.

4. The milling and baking tests of the new Pusa hybrids,
Nos. 107, 108, and 110, show that, inthe process of hybridization, the
milling and baking qualities of the strong parent have been trans-
mitted unimpaired to the offspring. As far as the tests with these
wheats have gone, the grain qualities have not been affected to any
extent by change of environment from Pusa tothe Central Provinces.

5. The results obtained generally confirm and amplify the
conclusions reached in the previous paper, namely, that “ strong
wheats with good milling qualities have been found to retain strength
and milling qualities both under canal irrigation on the alluvium
and also on the black soils of Peninsular India. In the future im-
provement of the wheats of these tracts, the question of grain quality
should receive particular attention.”

erred aN

fuzaffarnagar
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such as Chemistry, Botany, Entomology and the like.

I, No. I,

I Nove il:
1 sta LN
NOs Ve
rs sNo. ihe
Ie avigg. VAG
II, No. 1G
II; No: » UII.
I, No. Ii:
RIANo:;= Eve
De Now ve

II, No. VI.
II, No. VII.

II, No. VIII.

II, No. IX.
III, No. I.

Hi Nos iI:

III, No. III.

Tit, No; DV.

RET, eNO; uv.

Ci, Now viL:

EVGNG sl.
IV, No. Il.

IV, No. III.

IV, No. IV.

nV. ‘No. cy.
IV, No. VI.
Vig. INOy ha

BOTANICAL SERIES.

Studies in Root Pavrasitism. The Haustorium of Santalum album.—
Part I.—Early Stages, up to Penetration by C. A. BARBER, M.A., F.L.S.
Price, Re. 1. (Out of print.)

Part I1.—The Structure of the Mature Haustorium and the Inter-relations
between Host and Parasite, by C. A. BARBER, M.A., F.L.S. Price, Rs. 3.
(Out of print.)

Indian Wheat Rusts, by E. J. BUTLER, M.B., F.L.S., and J. M. HAYMAN,
D.V.Ss. Price, Rs. 3. (Out of print.)

Fungus Diseases of Sugarcane in Bengal, by E. J. BUTLER, M.B., F.L.S.
Price, Rs. 3. (Out of print.) ;

Gossypium obtusifolium, Roxburgh, by I. H. BURKILL, M.A. Price, Re. 1.

An Account of the Genus Pythiwm and some Chytridiacee, by E. J. BUTLER,
M.B., F.L.S. Price, Rs. 4-8.

Cephaleuros virescens, Kunze ; The Red Rust of Tea, by HAROLD H. MANN,
p.se.; and C. M. HUTCHINSON, B.A. Price, Rs. 4. (Out of print.)

Some Diseases of Cereals caused by Sclerospora graminicola, by K. J.
BUTLER, M.B., F.L.S. Price, Re. 1-8. ,
The Indian Cottons, by G. A. GAMMIKE, F.L.S. Price, Rs. 7-8. (Out of print.)
Note on a Toxic Substance excreted by the Roots of Plants, by F.

FLETCHER, M.A., B.Sc. Price, Re. 1-8.

Studies in Root Parasitism. III.—The Haustorium of Olaa scandens, by
C. A. BARBER, M.A., F.L.S.° Price, Rs. 2-8.

Studies in Root Parasitism. IV.—The Haustorium of Cansjera Rheedii, by
C. A. BARBER, M.A., F.LS. Price, Rs. 2-8. (Out of print.)

Some Experimentsin the Hybridising of Indian Cottons, by P. F. Fyson,
B.A., F.L.S. Price, Re. 1-8. (Out of print.)

The Varietal Characters of Indian Wheats, by ALBERT HOWARD, M.A.,
A.R.C.8., F.L.S.; and GABRIELLE L. C. HOWARD, M.A. Price, Re. 1.
(Out of print.)

The Mulberry Disease caused by Coryneum mori, Nom., in Kashmir, with
Notes on other Mulberry Diseases, by E. J. BUTLER, M.B., F.L.S. Price,
Re. 1-8. (Out of print.)

The Wilt Disease of Pigeon-Pea and the Parasitism of Neocosmospora
vasinfecta, Smith, by E. J. BUTLER, M.B., F.L.S. Price, Rs. 3.

Studies in Indian Tobaccos. No. I.—The Types of Nicotiana rustica,
L., Yellow Flowered Tobacco, by ALBERT HOWARD, M.A., A.R.C.S., F.LS. 5
and GABRIELLE L. C. HOWARD, M.A. Price, Rs. 4.

Studies in Indian Tobaccos. No. 11.—The Types of Nicotiana tabacum,
L., by ALBERT HOWARD, M.A., A.B.C.S., F.L.8.; and GABRIELLE L. C,
HOWARD, M.A. Price, Rs. 9.

Studies in Indian Fibre Plants. No. I.—On two varieties of Sann,
Crotalaria juncea, L., by ALBERT HOWARD, M.A., A.R.C.S., F.L.S.5 and
GABRIELLE L, C, HOWARD, M.A. Price, Re. 1. o

The Influence of the Environment on the Milling and Baking Qualities of
Wheat in India. No. L—The Experiments of 1907-08 and 1908-09, by
ALBERT HOWARD, M.A, A.R.C.S., F.L.S.; H. M. LEAKE, M.A., F.L.S.; and
GABRIELLE L. C. HOWARD, M.A. Price, Re. 1-8.

The Bud-Rot of Palms in India by E. J. BUTLER, M.B., F.LS,
Price, Rs. 2. ; ;

The Economic Significance of Natural Cross-fertilization in India, by ALBERT
HOWARD, M.A., A.R.C.S., F.L.S.; GABRIELLE L. C. HOWARD, M.A.; and
ABDUR RAHMAN KHAN. Price, Rs. 4-8. :

The Millets of the Genus Setaria in the Bombay Presidency and Sind, by
G. A. GAMMIE, F.L.S._ Price, Re. 1. 2 ae Pe)
Studies in Indian Fibre Plants. No. 2.—On Some New Varieties of Hibiscus
cannabinus, L., and Hibiscus Sabdariffa, L., by ALBERT HOWARD,
M.A., A.R.C.S., F.L.S.; and GABRIELLE L. C. HOWARD, M.A. Price, Rs. 3.

Notes on the Incidence and Effect of Sterility and Cross-fertilization in
the Indian Cottons, by H. M. LEAKE, M.A., F.1.S.; and RAM PRASAD.
Price, Re. 1. : Re

The Inheritance of Red Colour and the regularity of self-fertiliz-
ation in Corchorus capsularis, the common Jute Plant, by I. ,
BURKILL, M.A.; and R. 8. FINLOW, B.8c., F.C.S. Price, Re. 1.

Observations on Certain Extra-Indian Asiatic Cottons, by H. M. LEAKE,
M.A., F.L.S.; and RAM PRASAD. Price, Re. 1-8.

The Morphology and Parasitism of Rhizoctonia, by F. J. F. SHAw, B.Sc.,
A.R.C.S., F.L.8. Price, Rs. 2

The Inheritance of some Characters in Wheat, I, by A. HOWARD, M.A,,
A.R.C-S., F.L.S. ; and GABRIELLE L., C, HOWARD, M.A. Price, Re. 1.

Viol, -V; Nos IL.

Vol. V, No. III.

Vol: VV. Nos EV.
Nol; V; No: V
Wools Vil; Nos
Vol) VI; No: I.
Vol. VI, No. III.
Wole VI; No. IV.
Wols Vil; No. Vi
Vol. VI, No. VI.
Vol. VI, No. VII.
Vol. VI, No. VIII.

Vol. I, No. I,
Vol. Ig No;) IL.
Vol. I, No. III.
Viole Iy-No. LV.
Vol. I, No. Vz.

Wolsds Now» Vi.
Vol. I, No. VII.

Viol, 1, No: VEIT.
Vol. I, No. IX.
Vol. I, No. dG,
Vol. II, No. I,

Viol. TIeNo. / IF.
Vol. II, No. III.

Vol. II; No. IV.
Vol. TiS No: Vi.

Vol: Dice Now aVii.

Vol. IIT, No. I.

BOTANICAL SERIES—contd.

The Influence of the Environment on the Milling and Baking Qualities
of Wheat in India. No. 2.—The Experiments of 1909-10 and 1910-11, by
A. HOWARD, M.A., A.R.C.S., F.L.S.; H. M. LEAKE, M.A., F.L.S.; and
GABRIELLE L. C. HOWARD, M.A. Price, Re. 1,

The Varieties of Soy Beans found in Bengal, Bihar and Orissa and their
Commercial possibilities, by E. J. WOODHOUSE, M.A.; and C. S. TAYLOR,
B.A. Price, Rs. 2.

On Phytophthora parasitica nov. spec. A new Disease of the Castor Oil
Plant, by J. F. DASTUR, B.Sc. Price, Rs. 2.

Studies in Peronosporacee, by E. J. BUTLER, M.B., F.LS.; and G. §.
KULKARNI, L.Ag. Price, Rs. 2.

Notes on Pollination and Cross-Fertilization in the Common Rice
Plant, Oryza sativa, Linn., by G. P. HECTOR, M.A-, B.Sc. Price, Ke. 1.

A Sclerotial Disease of Rice, by F. J. F. SHAW, B.Sc., A.R.C.S., F.L.S.
Price, Re. 1.

Studies in Indian Tobaccos. No. 3—The Inheritance of Characters in
Nicotiana tabacum, L., by GABRIELLE L. C. HOWARD, M.A. Price, Rs. 3.

Studies in Indian Cottons, Part I—The Vegetative Characters, by H. M
LEAKE, M.A., F.L-S.;and RAM PRASAD. Price, Rs. 3-8. (Out of print.)

The Red Rot of Sugarcane, by HE. J. BUTLER, M.B., F-L.S.; and A. HAFIZ
KHAN. Price, Re. 1.

Some New Sugareane Diseases, by E. J. BUTLER, M.B., F.L.S.; and
A. HaFiz KHAN. Price, Rs. 2.

Preliminary Note on the Classification of Rice in the Central Provinces, by
R. J. D. GRAHAM, M.A., B.Sc. Price, Re. 1-8.

The Influence of the Environment on the Milling and Baking Qualities
of Wheat in India, No.3. The experiments of 1911-12, by A. Iiowarp,
U.1.E., M.A. ; H. M. LEAKE, M.A.; and G L. ©. HowaRD, M.A. Price,
Re. 1 or 1s. 6d.

CHEMICAL SERIES.

The Composition of Indian Rain and Dew, by J. WALTER LEATHER,
Ph-D., F:1.¢. Price, Re: 1.

The Composition of the Oil Seeds of India, by J. W. Learuer, Ph.v., F.L.¢,
Price, Re. 1. (Out of print.)

The Pot-Cuiture House at the Agricultural Research Institute, Pusa, by
J. W. LEATHER, Ph.D., F.1.c. Price, Rs. 3.

Experiments on the Availability of Phosphates and Potash in Soils, by
J. W. LEATHER, Ph.D., F.1.C. Price, Re. 1-8.

The Construction of Drain Gauges at Pusa, by M. H. ARNOTT, M.INST.C.E.,
with a Preface by J. W. LEATHER, Ph.D., F.I.C. Price, Rs. 3. (Out of
print.)

The Loss of Water from Soil during Dry Weather, by J. WALTER
LEATHER, Ph.D., F.I.C. Price, Rs. 2. (Out of print.)

The System Water, Calcium Carbonate, Carbonic Acid, by J. WALTER
LEATHER, Ph.D., F.I.C.; and JATINDRA NATH SEN, M.A., F.CS.
Price, Re. 1.

Water Requirements of Crops in India, by J. WALTER LEATHER, Ph.D.,
F.1.c. Price, Rs. 3.

The Nature of the Colour of Black Cotton Soil, by H. E. ANNETT, B.Sc.
(Lond.), F.C.S., M.S.E.A.C. Price, Re. 1.

Water Requirements of Crops in India—II, by J. WALTER LEATHER,
Ph.D., F.1.C. Price, Rs. 2-8.

The Composition of the Milk of some Breeds of Indian Cows and Buffaloes
andits Variations, Part I, The milk of some breeds of Indian cows, by A.
A. MEGGITT, B.Sc. (Lond.), F.c.s.; and H. H. MANN, D.Sc. Price, Re. 1-8.

Records of Drainage in India, by J. WALTER LEATHER, Ph.D., F.1.C,
Price, Re. 1.

The Rab System of Rice Cultivation in Western India, by H. H. Mann,
p.sc.; N. V. JOSHI, B.A., B.Sc., L.Ag.; aud N, V. KANITKAR, B.Ag.
Price, Re. 1. ;

The Composition of the Milk of some Breeds of Indian Cows and Buffaloes
and its Variations, Part II, The milk of some breeds of Andian buffaloes,
by A. A. MEGGITT, B.Sc.; and H. H. MANN, D.Sc. Price, Re. 1-8.

A contribution to the knowledge of the Black Cotton Soils of India,
by W. H. Harrison, M.Sc., and M. R. RAMSWAMY SIVAN, B.A.
Price, Re. 1. neh ed } ,

The Date Sugar Industry in Bengal. An investigation into its Chemistry
and Agriculture, by H. E. ANNETT, B.S8c., F.C.S., assisted by G. K
LELE, L.Ag., and BHAILAL M. AMIN, B.A. Price, Rs. 3

Evaporation from a Plain Water Surface, by J. WALTER LEATHER, Ph.D.,
F.1.C. Price, Re. 1.

Vol. III, No. II.

Vol

IE Ne; TET,

Vol. III, No. IV.

Volt, Ne.-sV.

Vol. III, No. VI.

Vol. III, No VII.

Vol. 111, No. VIII.

Vol. III, No. IX.
Vol. I, No. I
Vol. I, No. II
Vol. I, No. ILI
Vol. I, No. IV.
Vol. I, No. Mic
Vol. I, Mo. ” Wii.
Vol. 11, No. I,
Vol. II, No. II.
Vol. II, No. III.
Wolotl; No» ‘LV.
Vol. II, No. V.
Vol. II, No. VI.
Vol. If, No. VII.
Vol. II, No. VIII.
Vol. II, No. IX.
Vol. II, No. X.
Vol. III.

Walks a0. Vas
Vol. 1V, No. Il.
Vol. IV, No. III.
Vol. 1V, No. IV.
Vo).IV, No. V-
Vol. IV, No. VI.
Vol. “WV, anor: ae

CHEMICAL SERIES—conid.

Studies in the Chemistry and Physiology of the Leaves of the Betel-vi
(Piper Betle) and of the Commercial Bleaching of Betavine Eaeaee
by H. H. Mann, p.sc.; D. L. SAHASRABUDDHE, B.Sc., L-Ag. ; and
V. G. PATWARDHAN, B.Ag. Price, Re. 1-8.

The Gases} of Swamp Rice Soils, by W. H. HARkISON, M.Sc., and P. A.
SUBRAMANIA AIYER, B.A. Price, Re. 1-8.

Experimental Error in Sampling Sugarcane, by J. W. LEATHER, Ph.D.

ee Price, es i ,

e Fractional Liquefaction of Rice Starch, by F. J. WARTH, M.Sc. ;

es - eee B.Sc. Price, Re. 1. MES a
‘he Yield an omposition of the Milk of the Montgomery herd J
and Errors in Milk Tests, by J. W. ieee nae pA =
A. ©. Doses, Price, Re. 1 or I's. 6d. ;

The System Potassium Nitrate, Sodium Chloride, Water, by J. W.
LEATHER, Ph.D., F.1.C., and JOTINDRA NATH MUKERJEE, B.A., B.SC.
Price, Re. 1 or 1s. 6d.

The Systems—(A) Water, Magnesium Carbonate, and Carbonic Acid, (B)
Water, Calcium Carbonate, Magnesium Carbonate and Carbonic
Acid, by J. W. LEATHER, eh.D., F.1.C., and JATINDRA NATH SEN, M.A,
F.c.S. Price, Re. 1 or 1s. 6d.

Studies of an Acid Soil in Assam, by A. A. MEGGITT, B.Sc., F.C.S. Price
Re. 1-8 or 2s. 6d. ;

ENTOMOLOGICAL SERIES.

The Bombay Locust—A Report on the investigations of 1903-04, by H.
M. LEFROY, N.A., ¥.4.8., F.Z.S. Price, Rs. 2-8.

The more Important Insects injurious to Indian Agriculture, by H.
M. LEFROY, M.A., F.E.S., F.Z.S. Price, Rs. 3. (Out of print.)

The Indian Surface Caterpillars of the Genus Agvrotis, by H. M. LKFROY,
M.A., F.E.S., F.Z.S.; and C, C. GHOSH, B.A. Price, Re. 1-8. (Out of print.)

Individual and Seasonal Variations in Helopeltis theivora, Waterhouse,
with description of anew species of Helopeltis, by HaroLp) H. Mann,

p.sc. Price, Re. 1-8.

The Coccide attacking the T’ea Plant in India and Ceylon, by I. E. GREEN,
¥.K.S. ; and HAROLD H. MANN, D.Sc. Price, Re. 1. (Out of print.)

The Mustard Sawfly, by H. M. LEFROY, M.A., F.E.S., F.Z8.; and C. C,
GHOSH, B.A. Price, Re. 1. (Out of print.)

The Rice Bug, by H. M. LEFROY, M.A., F.E.S., F.Z.S.__ Price, Re. 1.

Remarks on Indian Scale Insects (Coccide@), Part III, by E. K. GREEN,
F.E.8., £-Z.8. Price, Re. 1-8.

The Red Cotton Bug, by H. M. LEFROY, M.A., F.E.S., F.Z.S. Price, Re. 1.
(Out of print.)

The Castor Semi-Looper, by H. M. LEFROY, M.A., F.E.S., F.Z.8. Price, Rs. 2.
(Out of print.)

The Tobacco Caterpillar, by H. M. LEFROY,M.A., F.E.S., F.Z.S. Price, Re. 1-8.
(Out of print.)

The Cotton Leaf-Roller, by H. M. LEFROY, M.A,, F.E.S., F.Z.S. Price, Re. 1-8.
(Out of print.)

Notes on Indian Scale Insects (Coccide), by H. MAXWELL-LEFROY, M.A.,
F.E.S., F.Z.8. Price, Re. 1-8. (Qut of print.)

Life-Histories of Indian Insects—I (Coleoptera), by H. MAXWELL-LEFROY,
M.A., F.E.8., F.Z.8. Price, Rs. 2.

Life-Histories of Indian Insects—II. Some Aquatic Rhynchota and
Coleoptera, by D. NOWROJEE, B.A. Price, Re. 1-8,

Life-Histories of Indian Insects—III. The Rhinoceros Beetle (Oryctes
rhinoceros) and the Red or Palm Weevil (Rhynchophorus ferrugineus),
by C. C. GHOSH, B.A, _ Price, Rs. 2.

The Food of Birds in India, by C. W. MASON, M.S.E.a.C., edited by H,
MAXWELL-LEFROY, M.A., F.E.S., F.Z.S. Price, Rs. 7-8.

Eri Silk, by H. MAXWELL-LEFROY, M.A., F.E.S., F.Z,S., and C, C. GHosH,
B.A. Price, Rs. 3.

Tetrigine (Acridiine) in the Agri. Research Institute, Pusa, Bihar, with

descriptions of new species, by J. L. HANCOCK, F.E.S. Price, Re. 1.

The Big Brown Cricket (Brachytrypes achatinus, Stoll), by C. C. GHOSH,

B.A. Price, Re. 1.

Life-Histories of Indian Insects—IV (Hymenoptera), by G. R. Dorr,

B.A. Price, Rs. 2.

Inquiry into the Insecticidal Action of some Mineral and other Compounds
on Caterpillars, by H. MAXWELL-LEFROY, M.A., F.E.S., F.Z.8: 5 and RLS
FINLOW, B.Sc., F.C.8. Price, Re-1-8. (Out of print.)

The ‘Psylla” disease of Indigo, by A. J. GROVE, M.Sc., and C. C,

GuHOSH, B.A. Price, Re. 1-8.

Life-Histories of Indian Insects—V (Lepidoptera), by C. C. GHOSH, B.A.

Price, Rs. 2-8.

BACTERIOLOGICAL SERIES.

" Studies in the Bacteriological Analysis of Indian Soils, No. 1, 1910-11 by
C..M. HUTCHINSON, B.a. | Price, Rs. 2-8. C
Rangpur Tobacco Wilt, by C. M. HurcHINsON, B.A. Price, Rs, 2.

VETERINARY SERIES.

[, Anaphylaxis in the larger Animals, by Major J. D. E. HOLMEs, M.A., D.Sc
Parc etgls ae Aba Rs. 2. PES hog
alvarsan in the Treatment of Surra in Horses, Dogs and Rabbi

Seige : Major J..D, E. HOLMES, M.A., D.Sc., M.R.€.V.S, Price, Re, 1-4, poe

-». Vol, I, No. III, Some more Successful Experiments on the Treatment of Surra in the

LT aga Camel with Recommendations for Systematic Treatment, by A. S. Leese,

SAC SWol,

r)

fame M.R.C.V.S.._ Price, Re. 1.

*~ Vol, I, No, IV. On the Immune Bodies occurring in Anti-Rinderpest Serum and on the
) _ Variations occurring in the Serum Proteins of Animals during Rinderpest

: and during Immunisation and Hyper-immunisation, by. P. HARTLEY,

Ph Basse f D.Se. Price, Rs. 2,

DS Syvoi,: IL, No, I. Some cases of Surra treated in the Field and in the Laboratory during the

These ; autumn of 1911, by Major J, D, E. HouMgs, M.A., D.Sc., M.R-C.V.S.

ee as Price, Re. l. -

af Vol. II, No- II, Rinderpest: Further Investigations on questions connected with the

“i ; Economical Production of Anti-serum, by Major J. D. E. HOLMES, M.A.,

42 ps D,Se.,-1.C.¥.D. Price, Re. 1.
,- Vol. Il, No. 111, The Curative Treatment of Hemorrhagic Septicemia in Cattle by the
: ; ». administration of Iodine and other notes on Chemiotherapy in Rinder-
, pest and Hxmorrhagic Septicemia, by Major J. D. E. HOLMES, 0.1.£.,
fi Rica ee M.-A., D-Sc,, M.R.C.V.S.. Price, Re. 1 or 1s. 6d.
> — Vol. TI, No. IV. The Vitality of the Hemorrhagic Septicemia Organism outside the body,
tite by Engl J. D, E. HOLMES, C€.1.E., M-A,, D-Sc., IC.V.D. Price, Re, 1
or 1s, 6d.
II, No.. V.. Bursati, by Major J. D. E, HOLMES, C.I.E., M.A., D-SC., M-R.C.V.8._ Price,
45 eee ; Re. 1-8 or 2s, 3d. ;
Vol, If, No: VI, Experimentson the treatment of Surra in Camels, by H. E. Cross, M.R.-0.v.s.,
i D-V.H., A.Sc.” Price, Re. 1 or 1s. 6d.
Vol. II, No. VII. Anthrax—some experiments on the Immunising Effect of the simultaneous
: injection of an Anthrax Attenuated Virus and an Anthrax Anti-Serum,
by Pa #4 J. ). B. HOLMES, €.1.E.,M.A-, D.Sc., M.R.C.Y.S,. Price, Re, 1
or ls. 6d.

BULLETINS ISSUED BY THE AGRICULTURAL RESEARCH
x - INSTITUTE, PUSA,

“Bieg ee a pe on Cotton in Bihar in 1904, by H. M. LEFROY, M.A, F.£.S,, F:Z.8, Price, As. 4
or * 7 {

An Outbreak of Cotton Pests in the Punjab, 1905, by H, M. Lerxoy, M.A,, F.E,8.,
B28. Price, As. ‘6 or 7a. (

The Extension of Jute Cultivation in India, by R, 8S. FINLOW, B.Sc.,'F.C.8, Price,
As, 12 or 1s, 2d. (Out of print.) :

First Report on the Fruit Experiments at Pisa, by A, HOWARD, M.A., A.R.C.Si,
F.L.S.. Price, As. 6 or 6d. 4

Report on Trials of the South African. Locust Fungus in India, by E. J. Bouter,
M.B., F.L.S. ; and H, M, LEFROY, M.A., F.E.S., F,Z.8. Price, As. 2 or 3d.

The Ticks Infesting Domesticated Animals in India, by C. WARBURTON, M.A,
Price, As. 4 or 6a. (Out of print.) .

A Preliminary Account of the Biting Flies of India, by H. M. Lerroy, M,a., F.B.S.,
¥.z.8, Price, Re. Lor 1s. 6d. (Out of print.)

Official and Recommended Methods for use in Chemical Laboratories of the Depart-
ments of Agriculture in India, by J. WALTER LEATHER, Ph.D., F.C. Price, As. 4
or 6d, . Blt. x}

Report on Coconut Palm Disease in Travancore, by E, J. BUTLER, M.B.,. F.LS.
Price, As. 6 or 6d. (Out of print.)

Treatment and Observation of Crop Pests on the Pusa Farm; by H. M. Lerroy, M.A,,

: F.E.S,, F.Z.8.; and ©, S. Misra, B.A. Price, As. 6 or 7d. (Out of print.)

. On Flax Dodder, by A, HowakbD, M.A., A.B.C.S., Fd.8.~ Price, As, 4 or 6d. (Out

0, nt.
Sie. re Malkin and Care of Lawns in India, by A. HOWARD, M.Ai, A.R.C,.8., F.L8.
Ey hS Price, As. 40r 6d. (Out of print.) ;
No. 13... Su ne at the Partabgarh Experimental Station, by G. OLARKE, F.1.C. ;. and Khan
ae ahadur S. M. HADI, M.R.A.C., M.R.A.S, Price, As. 6 or 6d.
--- No. 14, The Milling and Baking Qualities of Indian Wlieats, No, 1, by A. HOWARD, M.A.,
et A.B.C.8., F.L.8, 3 and GABRIELLE L. C. HowARD, M.A. Price, As. 4 or 6d.
Note on the Extension of Cultivation of Fibre Plants in India. Price, As. 6 or 8d.
Second Report on the Fruit Experiments at Pisa, by A. HOWARD, M.A, A.B.C.8.,
F.1,8, Price, As. 6 or 8d, - Sina

Fe BYE BEING ISSUED BY THE Laweuuiceal RESEARCH -
INSTITUTE, PUSA-—coneld,

No. 17... The Milling and Baking Qualities of Indian Wheats, ‘No. 2. Some new Pilea sel t a ot
tested in 1909, by A. HOWARD, M.A. A.R.C.S., F,L.8: 5 and cad ve

paar BOv SEO. MA, cae ice, As. 6 or 8d,
Arn + No. 18. Report on t utbreak of Blister- Blight on Tea in the Dai jeelin District i h )
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