NationalLibraryOfScotland/medical-history-of-british-india huggingface dataset:
29
pseudo-spicate system of female flowers—in the axils of the leafy bracts, and a
weak development—a solitary involving bract with its contained female flower—in
the axils of the concomitant stipules (PLATE 1, figs. 1, 3 ; and PLATE 2,
figs. 1-3).
Alternation between male inflorescence and female flower. —The contents of
the stipular axils, which are the primary objects of contrast, are thus, in the males
a cymose system of male flowers, and in the females a single involving bract with
a solitary female flower. This is brought out even more strongly in the case of
abnormally monœcious plants. We have seen (PLATE 1, fig. 2) how manifest the
alternation between cymose male panicle and solitary female flower is, in the case
of the contents of the axils of the basal stipules in the lowest floral branches of
otherwise normal males ; and have encountered the converse opposition in the
case of the contents of the axils of the basal stipules in otherwise normal female
inflorescences. The contrast is even more marked in the case of Sheória hemp
(PLATE 5, fig. 7). In Cannabis therefore, whether in normal or in abnormal
conditions, that this, whether in the same or in different individuals, the
change of modus from male to female does not result in an alternation of male
flower and female flower, but in an alternation of male inflorescence and female
flower, The absence of a female whorl in the male flower, so long at least as the
stamens are situated in the same plane, and the absence of a male whorl in the
female flower, are in keeping with this observation.
Alternation between male flower and female inflorescence. —If, however,
the change of modus attending the substitution of male for female be interpreted
aright as involving an alternation between female flower and male inflores-
cence, the converse change of modus, attending the substitution of female for
male, should involve a corresponding alternation between male flower and
female inflorescence. The evidence that this is the case seems very definite.
In the abnormal male flower recorded by Autenrieth, within which was produced
an axis bearing a spiral of female flowers with their subtending involving bracts ;
in those recorded by Gasparrini where a normal whorl of sepals subtends a spiral
of stamens on an axis tipped by a female flower ; and in all the intermediate condi-
tions exhibited by the abnormal female spikelets of Mória hemp (PLATE 5, figs.
2-6), we have abundant evidence that the male flower and the female
axillary spikelet are homologous throughout. But if the male flower and the
female inflorescence be equivalent their component parts must likewise be equi-
valent, and the contrast cannot in this case be between flower and flower, but
between the male stamen with its co-ordinate sepal on the one hand, and the
female ovary with its co-ordinate involving bract on the other. These Mória
spikelets show clearly that this is the only contrast possible. The bearing of
this on the structure of the male flower is therefore clear, its bearing on the
3
0
structure of the female flower has now to be developed ; the only point to be
noted before we proceed to do so is that, since the sepals of the male flower
correspond to the involving bracts of the female spikelet, no contrast is permissible
between male perianth and female perianth, the latter being an organ quite
unrepresented in the male flowers.
Channels of Diclinism. —In plants of Mória hemp it is usual to find, closely
applied to the often open monocarpellary ovary which constitutes the terminal
female flower of the spikelet, a single stamen, sometimes sessile, at other times
stalked (PLATE 5, figs. 2, 3 ; 16, 17). The two look as if they were in the same hori-
zontal plane and belonged to the same whorl (we know that at any rate they
belong to the same spiral), but the absence of a perianth, which is a characteristic
of the terminal female flowers of these spikelets, whether complete or in-
complete, renders the condition equivocal. The case is, however, different when
a study is made of the abnormal flowers so common in Sheóriahemp (PLATE
5, figs. 9-13). From these we learn that within the involving bract there are
two points of growth, one external, nearest to the involving bract, and another
internal, nearest to the floral axis. Of these two channels of growth only the
posterior internal is essentially that in which the female influence shows itself ;
the anterior external is essentially the channel in which the male influence is
manifested. In plants where the female modus is the dominant one, growth
takes place along both lines though most markedly in the posterior channel, and
the two are subsequently surrounded by a special covering, the perianth. In
the axil of the phyllome which marks the posterior channel of growth appears
an ovule that becomes intimately associated with that phyllome, and the opposed
anterior phyllome, by a process of growth corresponding to pistillody of the
andrœcium, instead of becoming a stamen, has the filament developed as a
sterile carpel which completes the ovarian cavity, and has the anther developed as
a style. The presence of the ovule thus not only serves to stimulate its own, the
posterior, phyllome to grow into a carpel but similarly stimulates the anterior
phyllome. When, on the other hand, the male modus is the dominant one and
there is no ovule present to stimulate its growth, the posterior phyllome does not
develop at all, the anterior only develops ; it grows now as a stamen and the
production of a perianth is not induced. Occasionally, even in this channel of
growth, there is a manifestation of female influence, to the extent of inducing the
formation of anthers whose connectives are prolonged as styles ; and occasionally
in the posterior channel there is a manifestation of male influence, since the fertile
carpel may be replaced by a deformed anther. But these exceptions may be
taken as proving the rule, which the majority of those teratological phenomena
appears to establish, that the diclinism of Cannabis asserts itself in two channels
of growth which are collateral and not successive.
31
Nature of the Diclinism. —The nature of this diclinism, like almost every
other point in connection with Cannabis, has been disputed, some contending that
it is inherent or primitive, others that it is abortive or derivative. The teratological
facts discussed above—and, since deviations from diclinism must in a normally
diœcious plant belong to the category of teratological phenomena, it is only
from the teratological side that the problem is to be satisfactorily discussed—leave
us in no doubt as to which view is correct. The diclinism here is in the highest
degree inherent, and is the result of a dichogamy that is essential, not accidental ;
the alternation in the sexual modus in the first place manifests itself in what are
morphologically the same organs, and in the second place precludes any compari-
son between the collection of organs that goes to form a male flower and the
collection of organs that constitutes a female flower.
Delpino,1 to whom we owe the distinction between primitive and secondary dicli-
nism, places Cannabis among diclinous plants of the former category, in which the
unisexuality is inherent, primitive and necessary. Macchiati, 2 on the other hand, believes
that Cannabis is a diclinous plant of the second category ; that it is constructed on a bisex-
ual type, becoming unisexual female by atrophy of the andrœcium, unisexual male by
atrophy of the gynœcium.
As the male and female flowers cannot be contrasted, either as regards structure or as
regards sexual modus, Macchiati's statement, which embodies the general belief, is obviously
without foundation and Delpino's view is amply substantiated.
SECTION VI. Summary of results.
The Problems.—Of the many unsolved problems relating to Cannabis only
those have been here attacked on which a study of abnormalities in the flower and
in the inflorescence is calculated to throw light. The chief of these relate to the
nature of the ovary, whether composed of one carpellary leaf or of two, or indeed
whether composed of an organ or organs that may be referred to the leaf-type
at all ; and to the origin of the ovule, whether axial or foliar. As a correlated
question the character of the diclinism exhibited by Cannabis has had to be
considered.
The monocarpellary theories hitherto held have postulated the existence of
only an anterior carpel in the female ovary ; the bicarpellary theories have
postulated the existence of both an anterior and a posterior carpel, of which only
the anterior is developed, the posterior remaining abortive. The latest view
suggests that the pistil may be an organ of intermediate character, a quid sui
generis neither foliar nor axial. The theories as to the foliar origin of the ovule
have involved either an interpretation of the placenta that is opposed to histological
characters, or the hypothetical existence of an intracarpellary organ of foliar
nature, intervening between axis and ovule, that does not actually occur.
1 La Distribusione dei sessi nelle piante : Milan, 1867.
2 Sulla sessualita, etc., della Canapa, p. 1.
32
The Results. —The instances of phyllody of the gynœcium, detailed in section
3, show that the pistil is composed of two carpels, both of leaf-type, and both
capable, under abnormal influences, of becoming leaves. But, of these two, it is
the posterior, not the anterior carpel that is fertile ; moreover, in an abnormal, but
still a closed ovary neither of the two becomes abortive ; the anterior, though
sterile and smaller than the posterior, nevertheless forms an organic part of the
pistil and, by union with the posterior, completes the ovarian chamber. This is
also precisely what occurs normally, and this interpretation, as is shown in section
4, fulfils all the requirements of organography, which no theory hitherto proposed
does ; accords with all known organogenic facts ; and finally, interprets satisfac-
torily all known histological characters.
The instances of phyllody of the ovule, detailed in section 3, show that the
ovule is truly axial, not, however, terminal but axillary to the posterior carpellary
phyllome with which it becomes from the first organically united and by which it
is carried by intercalary growth to the apex of the loculus. This interpretation,
as is shown in section 4, again satisfies all the demands of organography,
organogeny and histology.
The character of the diclinism is shown, from an examination of the facts
detailed in section 3, to be primitive and essential, not, as has been generally
contended, accidental and derivative ; the condition is further shown to be
manifested in channels of growth that are collateral and not successive ; moreover,
its phases do not, as is usual, involve a change of modus within comparable flowers
of male sex and of female sex. The flowers are, alike as regards structure and
development, incapable of comparison either as a whole or in detail ; the change
of sexual modus involves a contrast and an alternation, which are habitual and
normal, between the female flower and the male inflorescence ; and a corresponding
contrast and alternation, which are casual and abnormal, between the female
inflorescence and the male flower.
G I. C. P. O.—No. 951 S. C.—21-9-04.—495.
Explanation of the Plates.
PLATE I.
CANNABIS SATIVA Linn. ♂ .
Fig. 1.—Floral branch of male plant of Cannabis, from upper portion of stern—from
nature.
,, 2.—Floral branch from near base of stem showingbranch upraised(extra-
axillary) and the first pairs of paniculate cymes of flowers replaced by
solitary female flowers—from nature.
„ 3.—Scheme of male inflorescence: I -VI primary continuations of axis ; α smaller
proximal, β larger and successively dividing distal branchlets of the cyme-
system.
„ 4.—Male flower, young— enlarged.
„ 5.—Male flower, expanded— enlarged.
„ 6.—Diagram of a male flower.
„ 7.—Stamen—enlarged.
„ 8.—Longitudinal section of an anther— enlarged.
„ 9.—Transverse section of an anther, before dehiscence—enlarged.
„
10.—Transverse section of an anther, after dehiscence— enlarged.
„
11.—Pollen-grain— much magnified.
Prain: Cannabis.
PLATE I.
D. Prain delt.
C A N N A B I S S A T I V A L i n n .
♂
K. D. Chandra lith.
PLATE II.
CANNABIS SATIVA Linn. ♀ .
Fig. 1.— Floral branch of female plant of Cannabis—from nature.
„
2 & 3.—Schemes of female inflorescence : ax. fol. leafy axis; ax. fl. flowering
shoot; st. stipules.
„
4 & 5.—Basal pairs of female flowers of European Hemp— after Schnitzlein.
„ 6.—Diagram of basal pair of female flowers: ax. fol. leafy axis; ax.fl. flowering
shoot; st. stipules.
„
7.—Female flower of Indian Hemp— from nature.
„
8.—Involving bract of the same—from nature.
„ 9.—Female flower, half of involving bract removed— from nature.
„
10.—Female flower, perianth laid open—from nature.
„ 11.—Female flower, half of perianth and half of ovary removed, showing side
view of ovule—from nature.
„ 12.—Female flower, half of perianth and half of ovary removed, showing ovule
from in front—from nature.
„ 13.—Tip of style with stigmatic papillæ—enlarged.
„ 14.—Nutlet and husk (withered involving bract)—from nature.
„ 15.—Nutlet, husk removed, seen from below— from nature.
„ 16.—Interior view of apex of nutlet—after B. Clarke, emended.
„ 17.—Ripe kernel removed from nutlet—from nature.
„ 18.—Vertical section of ripe kernel—from nature.
„ 19.—Transverse section of ripe kernel, above the middle—from nature.
„ 20.—Transverse section of ripe kernel, below the middle—from nature.
„ 21.—Embryo, removed from albumen—from nature.
PLATE II.
D . Prain. delt.
K. P. Dass lith.
CANNABIS SATIVA Linn. ♀
PLATE III.
ORGANOGENY OF CANNABIS ♀.
Fig. 1-6.—Successive transverse sections through pedicel and lower part of ovary of
female flower of Cannabis to show position and destiny of the fibro-vascular
traces. (st., stipule ; br. inv., involving bract ; α trace innerving the
involving bract ; β, trace innerving the anterior, external, sterile carpel ;
γ, trace innerving the posterior, internal fertile carpel ; δ, trace innerv-
ing funiculus of ovule ; х , secondary trace derived from γ—after Briosi
and Tognini.
„ 7.—Vertical section through pe licel and base of ovary, showing the foregoing
traces, β, γ, δ and х —after Briosi and Tognini.
„ 8.—Apex of female inflorescence, showing earliest stages of development of
female flowers ; ov., ovules ; st. bracts, corresponding to stipules ; br.
inv., involving bract ; αх . fl., floral axis—after Briosi and Tognini.
„
9.—Slightly more advanced stage of development of a female flower ; ov. ovule ;
st., stipular bract ; br. inv., involving bract— after Briosi and Tognini.
„ 10.—Young female flower ; ov., ovule ; car., ovary ; per., perianth ; br. inv. involv-
ing bract— after Briosi and Tognini.
„ 11.—Vertical section (diagrammatic) of young female flower, at an early stage
before the axillary position of the axial ovule is marked— after Briosi
and Tognini.
„ 12.—Vertical section (diagrammatic) of young female flower, to show the position
of the ovule as axillary to the posterior carpellary phyllome — after
Briosi and Tognini.
„ 13.—Young female flower, ovarian parietes hiding the ovule but the cavity net
yet closed—after Briosi and Tognini.
„ 14.—More advanced stage of young female flower, the opening contracted but still
unclosed— after Briosi and Tognini.
„ 15.—Further stage of female flower, the styles forming but not yet united at their
bases,— after Briosi and Tognini.
„ 16.—Still further stage of female flower, the cavity closed by union of the bases
of the styles, but the line of union still visible as a faint cicatrix— after
Briosi and Tognini.
„ 17.—Longitudinal section (diagrammatic) of recently closed ovary, showing
obcampylotropous ovule still destitute of ovular integuments— after
Briosi and Tognini.
„ 18-20.—Vertical section (diagrammatic) of young flower of female Cannabis,
to show development of ovule integuments—after Briosi and Tognini.
„ 21.—Young unfertilized ovary— after Schleiden, and from nature.
„ 22.—Vertical section of young unfertilized ovary— after Schleiden, and from
nature.
„ 23.—Vertical section of young ovary, to show commencement of development of
embryo— after Schleiden, and from nature.
Prain: Cannabis.
PL ATE III.
E Briosi et Tognini, et Schleiden.
O R G A N O G E N Y O F C A N N A B I S
♀
.
K.P.Dass lith.
PLATE IV.
PHYLLODY OF GYNŒCIUM IN CANNABIS.
Fig. 1-3.—Pathological conditions due to Insect Blight— from nature.
„ 4.—Monstrous gynœcium with closed carpels, posterior view— from nature.
„ 5.—Monstrous gynœcium with closed carpels, lateral view— from nature.
„ 6.—Monstrous gynœcium with closed carpels, anterior view— from nature.
„ 7.—Monstrous gynœcium with closed carpels, laid open to show rudimentary
ovule—from nature.
„ 8.—Perianth of monstrous gynœcium, laid open—from nature.
„ 9-12.—A series of monstrous gynœcia with closed carpels, laid open to show
variations in position and nature of rudimentary ovules— from nature.
„ 13.—Monstrous gynœcium with gaping carpels, posterior view—from nature.
„ 14.—Monstrous gynœcium with gaping carpels, lateral view— from nature.
„ 15.—Monstrous gynœcium with gaping carpels, laid open— from nature.
„ 16.—Monstrous gynœcium with carpels replaced by free leaflets— from nature.
D. Prain delt.
K. D. Chandra lith.
Prain: Cannabis.
PLATE IV.
PHYLLODY OF GYNOECIUM IN CANNABIS.
PLATE V.
KHÁSIÁ, MÓRIA AND SHEÓRIA CANNABIS.
Fig. 1.—Flower from plant of khásiá Hemp, abnormal female apparently organically
perfect but functionally sterile—from nature.
„ 2.—Inflorescence from plant of Mória Hemp, i.e., Cannabis with male flowers
on a female plant—from nature.
„ 3.—Terminal flowers from Mória inflorescence—from nature.
„ 4.—A second inflorescence from plant of Mória Hemp—from nature.
„ 5.—A third inflorescence from plant of Mória Hemp— from nature.
„ 6.—Terminal flowers from a fourth plant of Mória Hemp—from nature.
„ 7.—Portion of inflorescence from plant of Sheória Hemp, i.e., Cannabis with
female flowers on a male plant—from nature.
„ 8.—Flower of Sheória Hemp showing only an involving bract and stalked
stigma—from nature.
„ 9.—Flower of Sheória Hemp with anterior style replaced by an anther—from
nature.
„ 10.—Vertical section of Sheória Hemp with anterior style replaced by an anther
—from nature.
„ 11.—Pair of flowers of Sheória Hemp, one perfect, the other deformed, the anterior
carpel and style replaced by a perfect stamen, the posterior carpel and
style replaced by a monstrous 4-celled non-polliniferous sessile anther—
from nature.
„ 12.—The same abnormal flower, half the perianth removed—from nature.
„ 13.—The abnormal monstrous sessile anther of the same flower, seen from behind—
from nature.
„ 14.—Male flower 'with sepals whorled' but with the 5 stamens spirally disposed
on an elongated receptacle tipped by a slightly modified carpel— after
Gasparrini.
„ 15.—'Hermaphrodite' flower, occurring spontaneously, the anthers removed, so as
to show scattered 'female organs'—after Autenrieth.
„ 16.—'Hermaphrodite' flower with single style, the ovary adnate to an anther—
after Autenrieth.
„ 17.—Carpel in which the ovary, adherent to the base of an anther, is almost
entirely abortive except that the styles are developed— after Gasparrini.
„ 18.—Anther with connective prolonged as a short style— after Autenrieth.
„ 19-20.—Two instances of carpels united, by means of the abortive ovary, which
is reduced to a connective, to the face of an anther of which one loculus
is almost completely abortive, the other little developed— after Gasparrini.
Prain; Cannabis.
PL ATE V.
D. Prain delt.
K H Á S I Á , M O R I A
A N D
S H E O R I A C A N N A B I S .
K. P. Dass lith.
No. 12
( NEW SERIES )
SCIENTIFIC MEMOIRS
BY
OFFICERS OF THE MEDICAL AND SANITARY DEPARTMENTS
OF THE
GOVERNMENT OF INDIA.
ON THE MORPHOLOGY, TERATOLOGY AND DICLINISM OF THE FLOWERS
OF CANNABIS.
BY
MAJOR D. PRAIN, M.B., I.M.S.
(Indian Medical Service. )
ISSUED UNDER THE AUTHORITY OF THE GOVERNMENT OF INDIA
BY THE SANITARY COMMISSIONER WITH THE GOVERNMENT
OF INDIA, SIMLA.
CALCUTTA:
OFFICE OF THE SUPERINTENDENT OF GOVERNMENT PRINTING, INDIA.
1904.
Price Annas 0-14-0 or 1s. 4d.
REPORT
ON
SANITATION,DISPENSARIES, AND JAILS IN
RAJPUTANA FOR 1919
AND ON
VACCINATION FOR THE YEAR 1919-20
Published by Authority
CALCUTTA
SUPERINTENDENT GOVERNMENT PRINTING, INDIA
1921
Price Rs. 2 As. 12.
REPORT
ON
SANITATION, DISPENSARIES, AND JAILS IN
RAJPUTANA FOR 1919
AND ON
VACCINATION FOR THE YEAR 1919-20
Published by Authority
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