exhibited in all that were in a healthy condition. I examined several,
and watched them for a long time. The phenomenon described depends
much upon the healthy condition of the frustule at the time; as the
movements of the diatoms became sluggish, the circulation gradually
slackens and then ceases altogether. I also saw a somewhat similar
action in the more active specimens of _P. hippocampus_ and _Navicula
cuspidata_, but the coarser markings and thickness of the wall of these
diatoms seemed to place greater difficulties in the way of observation
than the finer valves of the _P. angulatum_. One thing I believe is
certain, that the circulation described is precisely similar to that
seen in the Closteria, or, on a much larger scale, in Chara and the
leaf of the Anacharis, bearing in mind also that in the Closterium
the cell is divided by a transverse suture, and in _P. angulatum_
by a longitudinal one (Plate II., Nos. 38-40). About the same time
some very lively specimens of the Pinnulariæ were sent to me, and the
movements of these frustules were more closely observed. One or two
of the more active would attack a body relatively larger than itself,
it would also force its way into a mass of granular matter, and then
recede from it with a jerky motion. In more than one instance a cell
of Palmoglæa was seized and carried away by the Pinnularia, the former
at the time being actively engaged in the process of cell division.
Other diatoms present among my specimens were also in an active
condition, and the circulation of granular matter in all was distinctly
visible. In the Pinnulariæ two large colourless vesicles were seen on
either side of the median nodule, each having a central nucleus, as
represented in the accompanying sketch, made while under observation
in two positions. The central portion of each frustule was closely
packed with a rich yellowish-brown coloured endochrome, interspersed
with a few fat globules. The phenomenon of cyclosis was not seen in
any of these diatoms, but I have satisfied myself, by staining, of the
presence of a delicately fine external protoplasmic covering in many
diatoms. That their movements resemble the gliding movements exhibited
by the Amœba can scarcely be doubted. Numerous forms of Diatomaceæ are
found growing on or attached to water-plants or pieces of detached
stalks, which, although generally simple, are sometimes compound,
dividing and subdividing in a beautiful ramous manner. Pinnulariæ,
Nitzschia, &c., are seen adherent by one extremity, about which they
turn or bend themselves as on a hinge. By the process of cell-division,
groups of Synedræ become attached by a point, in a fan-like form.
The fan-like collection of frustules is said to be flabellate, or
radiate. In Licmophora, Achnanthes and other species (Plate II., Nos.
29-33) the double condition of union of frustules and of attachment by
a pedicle are illustrated. When a stipe branches it does so normally
in a dichotomous manner, each new individual being produced by a
secondary pedicle. This regular dichotomy is seen in several genera:
Cocconema and Gomphonema, the latter more perfectly in Fig. 296, from a
microphotograph, in which a branching, or rather longitudinal, rupture
of the pedicle takes place at intervals, and the entire organism
presents a more or less complete flabella, or fan-like cluster, on the
summit of the branches, and imperfect or single frustules irregularly
scattered throughout the whole length of the pedicle.
_Isthmia enervis_ (Fig. 297).--The unicellular frustule of this species
is extremely difficult to define, owing to the large areolations of the
valves; it has a remarkable internal structure. The olive-brown cell
contents are found collected, for the most part, into a central mass,
from which radiating, branched, granular threads extend to and unite
with the periphery. When viewed by a magnifying power of 600 or 700
diameters, these prolongations are seen to be composed of aggregations
of ovate or spindle-shaped corpuscles, held together by protoplasmic
matter. These bodies are sometimes quiescent, but more often travel
slowly to and fro from the central mass. The general aspect under these
conditions so nearly corresponds to the characteristic circulation in
the frustules of unicellular plants and of certain rhizopoda, that it
is difficult to realise that the object when under examination is an
elegant marine diatom.
[Illustration: Fig. 297.--_Isthmia enervis._ Microphotograph.]
There is a large section of diatoms in which the frustules are
diffused throughout a muco-gelatinous envelope in a definite manner.
Histologically this is homologous with the pedicles and connecting
nodules thrown out during the act of self-division, and in some species
(Cocconeis, Fragillaria, &c.) it often persists after that act is
complete.
Diatomaceæ, Recent and Fossil.
[Illustration: Fig. 298.--Fossil Diatoms from Springfield (Barbadoes).
1, Achnanthidium; 2, _Diatoma vulgare_, side view and front view; 3,
Biddulphia; 4, 5, 6, 7, _Amphitetias antediluviana_, front view, with
globular and oval forms; _Gomphonema elongatum_ and _capitatum_.]
_Fossilised Diatomaceæ._--Dr. Gregory was of opinion that a large
number of diatoms separated into species are only transition forms,
and more extended observations have proved that form and outline are
not always to be trusted in this matter. Species-making is a modern
invention, and can hardly apply to the indestructible fossilised forms
of the frustules of Diatomaceæ, with their beautiful sculpturings and
geometrical constructions, which have not been materially changed since
they were first deposited. Startling and almost incredible as the
assertion may appear to some, it is none the less a fact established
beyond all question, that some of the most gigantic mountain-ranges,
as the mighty Andes, towering into space 25,250 feet above the level
of the sea, their base occupying vast areas of land; as also massive
limestone rocks; the sand that covers boundless deserts; and the soil
of many wide-extended plains, are each and all principally composed of
Diatomaceæ. And, as Dr. Buckland once observed: “The remains of such
minute animals have added much more to the mass of materials which
compose the exterior crust of the globe than the bones of elephants,
hippopotami, and whales.”
In 1841 the late Mr. Sollitt, of Hull, discovered the beautiful
longitudinal and transverse _striæ_ (markings) on the _Pleurosigma
hippocampus_. A curved graceful line runs down the shell, in the centre
of which is an expanded oval opening. Near to the central opening the
dots elongate crossways, presenting the appearance of small short bands.
In the vicinity of this town many interesting varieties of Diatomaceæ
have been found, the beauty of the varied forms of which are constantly
under investigation; at the same time some of them are highly useful,
as forming that class of _test objects_ which are better calculated
than many others for determining the excellence and powers of certain
objectives. Mr. Sollitt carefully measured the markings on some of the
frustules and found they ranged between the 1/30000th and 1/130000th of
an inch; the _Pleurosigma strigilis_ having the strongest markings, and
the _Pleurosigma acus_ the finest.
Mr. J. D. Sollitt not only first proposed their use, but he also
furnished the measurements of the lines of the several members of this
family, as follows:--
Amphipleura pellucida, or Acus, 130,000 in the inch, cross lines.
" sigmoidea, 70,000 in the inch.
Navicula rhomboides, 111,000 in the inch, cross lines.
Pleurosigma fasciola, fine shell, 86,000 in the inch, cross lines.
" " strong shell, 64,000 in the inch, cross lines.
" strigosum, 72,000 in the inch, diagonal lines.
" angulatum, 51,000 in the inch, diagonal lines.
" quadratum, 50,000 in the inch, diagonal lines.
" Spencerii, 50,000 in the inch, cross lines.
" attenuatum, 42,000 in the inch, cross lines.
" Balticum, 40,000 in the inch, cross lines.
" formosum, 32,000 in the inch, diagonal lines.
" strigilis, 30,000 in the inch, cross lines.
[Illustration: PLATE XII.
MICRO-PHOTOGRAPH OF TEST DIATOMS.]
Lichenaceæ.
The lichens are a family of autonomous plants, an intermediary group of
algals or cellular cryptogams, drawing their nourishment from the air
through their whole surface medium, and propagating by spores usually
enclosed in asci, and always having green gonidia in their thallus.
Their gonidia, bright coloured globular cells, form layers under the
cortical covering of the thallus, and generally develop in the form
of incrustations, which cover stones, wood, and the bark of trees,
or penetrate into the lamellæ of the epidermis of woody plants. The
gonidia of lichens partake of both the character of vegetative and
reproductive cells.
The thallus in the fructicose group attaches itself by a narrow base,
growing in the form of a miniature shrub. Another group is met with
in a slimy condition--the gelatinous lichens. These species, for the
most part, furnished dyes before the discovery of the coal-tar dyes.
In many of the _Palmella cruenta_, commonly found growing on the walls
and roofs of houses, a colourless acid liquid is found, which, on being
treated with alkali, produces a bright yellow colour; and another,
_Avernia vulpina_, furnishes a brown dye; the _Rocella fuciformis_
and _R. tinctoria_ yield the purple dye substance known as orchil, or
archil, from which the useful blue paper of the chemist for testing
acidity is manufactured. Usnic acid, combined with green and yellow
resins, seems to be more or less a constituent of many lichens.
A vertical section of _Palmella stellata_ is given in Plate I., No.
26, in which the emission of the ripe spores of the lichens is seen
to be not unlike that which takes place in some of the fungi, Pezizæ,
Sphæriæ, &c. If a portion of the thallus be moistened and placed in a
common phial, with the apotheca turned toward one side, in a few hours
the opposite surface of the glass will be found covered with patches of
spores, easily perceptible by their colour; or if placed on a moistened
surface, and one of the usual glass slips laid over it, the latter
will be covered in a short time. As to the powers of dissemination of
these lowly organised plants, an observation led to the conclusion
that the gonidia of lichens have greater powers in this direction
than was formerly supposed. It is found that by placing a clean sheet
of glass in the open air during a fall of snow, and receiving the
melting water in a tube or bottle, quantities of what has been looked
upon as a “unicellular plant” can be taken, the cells of which may be
kept in a dormant condition for a long time during cold weather, but
upon the return of spring warmth and moisture they begin to increase,
by a process of subdivision, into two, four or eight portions; these
soon assume a rounded form, and burst the parent cell-wall open; these
secondary cells then begin to divide and subdivide again, and the
process may go on without much variation for a long time. The phenomena
described may be watched by taking a portion of the bark of a tree on
which Chlorococcus has been deposited, and placing it under a glass to
keep it in a moderately moist atmosphere; the only difference being a
change in colour, caused by the growth of the fibres, as may be seen
on microscopical examination. “And this,” says Dr. Hicks, who first
observed this phenomenon in plant life, “is an instructive point,
because it will be found that the colour varies notably according to
the lichen prevalent in its neighbourhood.”[59] He believes there can
be no doubt that what has been called Chlorococcus is nothing more
than the gonidia of a lichen; and that under suitable conditions,
chiefly drought and warmth, the gonidium often throws out from its
external envelope a small fibre, which, adhering and branching, forms
a “soridium.” “The soridia remain dormant for a very long time, and
do not develop into thalli unless in a favourable situation, in some
cases it may be for years. It will be perceived that the soridium
contains all the elements of a thallus in miniature; in fact, a thallus
does frequently arise from one alone, and the fibres of neighbouring
soridia interlace; thus a thallus is matured very rapidly. This is
one of the causes of the variation of appearance so common in many
species of lichens, more readily seen towards the centre of the parent
thallus. When the gonidia remain attached to the parent thallus, the
circumstances are, of course, more favourable, and they develop into
secondary thalli, attached more or less to the older one, which, in
many instances, decays beneath them. This process being continued
year after year gives an apparent thickness and spongy appearance to
the lichen, and is the principal cause of the various modifications in
the external aspect of the lichens which caused them formerly to be
misunderstood and wrongly classified.”[60]
The erratic lichens are found among the genus Palmella, some of which
grow among boulders of the primary and metamorphic formations, curled
up into a ball, and only fixed to their matrix by a slender thread. The
globular _Lecanora esculenta_ will at times suddenly cover large tracts
of country in Persia and Tartary, where it is eaten by the cattle.
During a scarcity of food a shower of these lichens, Mr. Berkeley tells
us, fell at Erzeroum, and saved the cattle from starvation.[61]
Another group of the Palmella, or Peltigeri, so named from the
target-like discs on their surface, spread their foliaceous fronds
over the ground, and as the fruit is marginal, it gives the thallus
a digitate appearance. These are often spotted over by a little red
fungus. The Lecidinei contains numerous species of the most varied
habits, and always crustaceous, and so closely adherent to the hard
rocks and stones on which they grow, that at length they disintegrate
them. From this low species a higher form arises, with erect branching
stems, and clothed with foliaceous, brightly-coloured scales.
The Coccocarpei is mainly distinguished by having orbicular discs
entirely deprived of the cortical envelope called an excipulum. The
discs spring at once from the medullary stratum, and contain asci and
sporidia similar to those of the minute fungi Sphæriæ. Some of the
lichens are themselves parasitic, and begin existence under the thick
skin of the leaves of tropical plants, and spread encrusting thallus
over their surface, the excipulum and perithecia being black; but
in most cases these are beautifully sculptured, and are interesting
objects for the microscope. Indeed, the sphere-bearing lichens,
with upright stems bearing globular fruit at the extremity of their
branches, are at first indicated by a swelling, but in time the outer
layer bursts and exposes sporidia, which are beautiful objects under
the microscope on account of their spherical form and more or less deep
blue tint. Humble and lowly as lichens may appear to be, they have been
divided into fifty-eight or more genera and 2,500 species. The brothers
Tulasne, De Bary, the Rev. Mr. Berkeley, and others, devoted great
attention to the peculiarities of their structure and natural history.
_Hepaticæ._--An intermediary group of much interest to the microscopist
are the Hepaticæ (liverworts). These are found growing on damp rocks in
the neighbourhood of springs and dripping banks. The scale-moss, the
_Marchantia polymorphia_ (Fig. 299), may be taken as typical of this
little group, with its gemmiparous conceptacles and lobed receptacles,
bearing archegones on transparent glass-like fruit stalks, carrying on
their summits either round shield-like discs or radiating bodies with a
striking resemblance to a wheel without its tyre.
[Illustration: Fig. 299.--_Marchantia polymorphia._]
The liverworts are closely allied to the mosses, and as much difficulty
was experienced in dividing the two, Hooker placed the whole under one
genus, the Jungermannia. More recently, however, they have been divided
into those with a stem and leaves confluent in a frond, Marchantia;
those with stem and leaves distinct, Jungermannia; and those with a
solitary capsule, filiform, bivalved, stalked, with a free central
placentation, Anthocotaceæ. Some botanists have further divided them,
but they are all extensively propagated by gemmæ.
The fronds carry the male organs, or _antherids_, and the disc, in the
first instance, bears the female organs, or _archegones_, and after
a time gives place to the _sporanges_, or spore cases. It is these
bodies which are of so much interest to microscopists; if the plant is
brought into a warm room, they suddenly burst open with some violence
the moment a drop of water is applied to them, and the sporanges are
dispersed in a small cloud of brownish dust. If this dust is examined
under a medium power, it is seen to consist of a number of chain-like
bodies, somewhat like the spring of a small watch; and if the process
of bursting be closely watched, these minute springs will be found
twisting and curling about in every direction. The structure of the
frond itself will be seen to be interesting when cut in the vertical
direction and placed under the microscope.
[Illustration: Fig. 300.--Gemmiparous conceptacle of _Marchantia
polymorphia_, expanding and rising from the surface of a frond. In the
interior are seen gopidial gemmæ already detached by the splitting of
the epiderm.]
The gemmæ of _Marchantia polymorphia_ are produced in elegant
membranous cups, with a toothed margin growing on the upper surface
of the frond, especially in very damp courtyards between the stones,
or near running water, where its lobed fronds are found covering
extensive tracts of moist soil. At the period of fructification the
fronds send up stalks, which carry at their summit round shield-like
radiating discs, which bear upon their surface a number of little
open basket-shaped “conceptacles.” These again expand into singularly
graceful cups (as in Fig. 300), and are found in all stages of
development. When mature, the basket contains a number of little green
round or oblong discs, each composed of two or more layers of cells;
the wall itself being surmounted by a glistening fringe of teeth, whose
edges are themselves regularly fringed with minute outgrowths. The cup
seems to be formed by a development of the superior epidermis, which is
raised up, and finally bursts and spreads out, laying bare the seeds.
The archegones of Marchantia are very curious bodies, while the elater
and spores are even still more so. These are elongated cells, each
containing a double spiral fibre coiled up in the interior. It is the
elasticity of this which tears apart the cell-membrane, and sends
forth the spores with a jerk, and thus assists in their dispersion.
Marchantia is the type of the malloid Hepaticæ.
Musci, Bryophyta.
Mosses are a beautiful class of non-vascular cryptogams. Linnæus
called them _servi_, servants or workmen, as they seem to labour to
produce vegetation in places where soil is not already formed. The
Bryophyta form three natural divisions: the Bryinæ, or true mosses;
the Sphagnaceæ, or peat-mosses; and the Hepaticæ, or liverworts. The
two first are commonly united. In these the sexual organs consist of
antheridia and archegonia, but they are of simpler structure than will
be found in ferns; and the first generation from the spore is asexual.
[Illustration: Fig. 301.--Screw-moss.]
The common or wall screw-moss (Fig. 301) grows almost everywhere, and
if examined closely, is seen to have springing from its base numerous
very slender stems, each terminating in a dark brown case, which
encloses antheroids. If a patch of the moss is gathered when in this
state, and the green part of the base is put into water, the threads of
the fringe will uncoil and disentangle themselves in a most curious and
beautiful manner; from this circumstance the plant takes its popular
name of screw-moss. The leaf usually consists of either a single or a
double layer of cells, having flattened sides, by which they adhere
one to another. The leaf-cells (Fig. 302) of the Sphagnum or bog-moss
exhibit a curious departure from the ordinary type; they are large,
polygonal, and elongated, and contain spiral fibres loosely coiled in
their interior. The young leaf does not differ from the older; both are
evolved by a gradual process of differentiation.
[Illustration: Fig. 302.--Section of leaf of Sphagnum moss, showing
large cells of spiral fibres and connecting apertures.]
Mosses, like liverworts, possess both antheridia and pistillida,
which are engaged in the process of fructification. The fertilized
cell becomes gradually developed into a conical body elevated upon a
footstalk, the walls of the flask-shaped body carrying the higher part
upwards as a _calyptra_ or hood upon its summit, while the lower part
remains to form a kind of collar round the base. These spore-capsules
are closed on their summit by _opercula_ or lids, and their mouths
when laid open are surrounded by a beautiful toothed fringe, termed
the _peristome_. This fringe is shown in Fig. 303, in the centre of a
capsule of Funaria, with its peristome _in situ_. The fringes of teeth
are variously constructed, and are of great service in discriminating
the genera. In _Neckera antipyretica_ the peristome is double, the
inner being composed of teeth united by cross bars, forming a very
pretty trellis. The seed spores are contained in the upper part of the
capsule, where they are clustered round the central pillar, termed the
_columella_; and at the time of maturity, the interior of the capsule
is almost entirely occupied by spores.
[Illustration: Fig. 303.--Mouth of Capsule of Funaria, showing
Peristome.]
[Illustration: Fig. 304.--Hair-moss in Fruit.]
The undulating hair-moss, _Polytrichum undulatum_ (Fig. 304), is found
on moist, shady banks of pools and rivulets. The seed-vessel has a
curious shaggy cap; but in its construction it is very similar to
that of the screw-moss, except that the fringe around its opening is
not twisted. The reproductive organs of mosses are of two kinds; the
capsule containing minute spores, _archegonia_, and the _antheridia_,
or male efflorescence. The capsule, _theca_, or sporangium, is
lateral or terminal, sessile, or on a fruit stalk (_seta_) of various
shapes, indehiscent, or bursting by four valves at the sides, or more
commonly by a deciduous cup, _operculum_. When this falls the mouth
of the capsule becomes exposed. The rim is crowned with tooth-like
or cilia-like appendages in sets of four or multiples of that
number--_peristome_. These are often brightly coloured and hydroscopic.
By simply breathing upon them they suddenly fly open, and are endowed
with motion, that is, if they contain spores. The spores on germination
produce a green confervoid-like mass of threads, from which the young
plant arises.
The Sphagnaceæ, or “bog mosses,” have been separated from true mosses
from the marked differences they present. The stem is more widely
differentiated, and throughout its structure a rapid passage of
fluid takes place. It has the power of absorbing moisture from the
atmosphere, so that if a plant be placed dry in a glass of water with
its rosette of leaves hanging over the edge, it acts like a syphon,
and the water will drop from it until the glass is emptied. As may be
supposed, the leaf is composed of large open cells, and it absorbs
more water than the root. The antherids or male organs of Sphagnaceæ
resemble those of liverworts, rather than those of mosses, both in
form and arrangement; they are grouped in “catkins” at the tips of the
lateral branches, each of the imbricated perigonal leaves enclosing
a single globose antherid on a slender foot-stalk, and surrounded by
long branched paraphyses of cobweb-like tenuity. The female organs,
or archegones, do not differ materially in structure from those of
mosses; they are grouped together in a sheath of deep green leaves at
the end of the shorter lateral branchlets at the side of the rosette
or terminal crown of leaves. The sporange is very uniform in all the
species, and the spores are in groups of fours, as in mosses, around a
hemispherical columella. These plants grow so rapidly that they soon
cover a pool with thin matted bundles of branches, and as they decay
they fall to the bottom, and become the foundation of the future bog or
peat moss.
=Felices.=--Of all the spore-bearing families the ferns are the more
universally known. They constitute an exceedingly numerous genera and
species, and vary from low herbaceous plants of an inch in height
to that of tree ferns, which attain a height of fifty or more feet,
terminating in a graceful coronet of fronds or leaves. Of whatever size
a fern may be, its spores are, for the most part, microscopic, produced
within the sporangium by cell division, and are therefore free and
variously shaped.
The true mode of development of ferns from their spores was that
furnished by Nägeli, who announced the existence of antheridia. On the
spore starting into life it sends out from the cell-wall of its outer
coat a white tubular projection, or root fibre (Fig. 305, A, B, and C),
which passes through the cell-wall of its outer coat. This attracts
sufficient moisture to burst open the outer, and then it begins to
increase by the subdivision of its cells, until the primary green
prothallus D is formed. This falls to the ground, and, being furnished
on its under side with thread-like fibres, fixes itself to the earth,
and thus is developed the rhizome, or root of the future plant. In each
of the antheridia, which are numerous, a cell is formed, chiefly filled
with albuminous matter and free spores, each having attached a flat
ribbon-like filament, or stermatoid, curled in a spiral manner. These
are ultimately set free by the rupture of the cell-wall, and commence
revolving rapidly by the agency of the whip-like appendage at the
larger end.
[Illustration: Fig. 305.--Development of the Globular Antheridium and
Spermatoids of _Pteris serrulata_.
A. Spores; B, C. Early stages of development; D. Prothallus with radial
fibres; _a_, _a_ and _a_, _b_ are stermatoids; and _h_, _h_. Enclosed
antheridia.]
The sporangia, or spore-cases, are, for the most part, globular in
form, and are nearly or quite surrounded by a strong elastic ring,
which in some cases is continued to form a stalk. When the spores
are ripe, this ring, by its elastic force, tears open the sporangia
and gives exit to a quantity of microscopic filaments, curled in
corkscrew-like fashion (Figs. 305 and 307). The ring assumes various
forms; in one group it passes vertically up the back of the sporangium,
and is continued to a point termed the stomata, where the horizontal
bursting takes place. This form is seen in Fig. 306, _a_, _b_. In other
groups it is vertical, as in _c_, _c_; in others transverse, as in _d_;
or apical, as at _e_; and in a few instances it is obsolete, as in _f_.
These are the true ferns, and their systematic arrangement is chiefly
founded on the peculiarity of the sori and sporangia, characters which
become quite intelligible by the aid of the microscope.
[Illustration: Fig. 306.--Sporangia of Polypodiaceous Ferns.
_a_, _b_. Polypodiaceæ; _c._ Cyantheineæ; _d._ Gleichenineæ; _e._
Schizeineæ; _f._ Osmundineæ.]
[Illustration: Fig. 307.--Spores of _Deparia prolifera_.]
The beautiful ringed sporangium of the fern (Fig. 307) when ruptured
gives exit to the dust-like spores; these, examined under a moderate
power, are seen to be sub-globose and pyramidal, the outer coat or
exospore being a coloured hyaline cell with nuclei similar to the
spores of mosses, but in which chlorophyll soon begins to form, and
from this little green embryonic growth the organs of reproduction are
formed.
In all ferns the pistillidia or archegonia are analogous to the ovules
or nascent seeds of flowering plants, and contain, like them, a
germinal vesicle, which becomes fertilized through the agency of the
spiral filaments, and then gradually develops into an embryo plant
possessing a terminal bud. This bud begins at once to unfold and push
out leaves with a circinate vernation, of a very simple form at first,
and growing up beneath the prothallium, coming out at the notch;
single fibrous roots are at the same time sent down into the earth,
the delicate expanded prothallium withers away, and the foundation of
the perfect fern plant is laid. When a fern acquires a considerable
stem, as in a tree fern, it consists of cellular tissue and an external
cortical portion forming fibro-vascular bundles, scalariform ducts, and
woody fibre. Fig. 308, _b_, shows an oblique section of the footstalk
of a fern leaf with its bundle of scalariform ducts.
These observations on ferns have acquired increased interest from
subsequent investigations made on the allied Cryptogams, and on the
processes occurring in the impregnation of the Conifers. Not only
have later researches furnished a satisfactory interpretation of the
archegonia and antheridia of the mosses and liverworts, but they have
made known and co-ordinated the existence of analogous phenomena in
the Equisetaceæ, Lycopodiaceæ, and Rhizocarpeæ, and prove, moreover,
that the bodies described by Dr. Brown in the Conifers under the name
of “corpuscles” are analogous to the _archegonia_ of the Cryptogams;
so that a link is hereby formed between these groups and the higher
flowering plants.
[Illustration: Fig. 308.--_a._ Vertical section of Fern-root, showing
spiral tissue and cells filled with granular bodies; _b._ Section of
Footstalk.]
_Equisetaceæ._--The development of _Horse-tails_ (Fig. 309), the
name by which they are commonly known, corresponds in some respects
with that of ferns. They comprise a little group, and the whole of
their structure is composed in an extraordinary degree by silex, so
that even when the organic portion has been destroyed by prolonged
maceration in strong acid, a consistent skeleton still remains. It
is this flinty material that constitutes their chief interest for
microscopists. A portion of their silicious particles is distributed
in two lines, arranged parallel to the axis of the plant, others are
grouped into oval forms, and connected by a chain as in a necklace. The
form and arrangement of the crystals are better seen under polarised
light. Plate VIII., No. 170, a portion of the epidermis, forms an
extremely beautiful object. Sir David Brewster pointed out that each
silicious particle has a regular axis of double refraction. What is
usually said to be the fructification of the Equisetaceæ forms a
cone or spike-like extremity to the top of the stem (Fig. 309), the
whole resembling a series of spike-like branches (the real stem being
a horizontal rhizome), and a cluster of shield-like discs, each of
which carries a circle of sporanges that open by longitudinal slits to
set free the spores which are attached to it in two pairs of elastic
filaments (shown in Fig. 291, F, G), _elaters_; these are at first
coiled up around the spore in the manner represented at G, but on their
liberation they extend themselves as shown at F. The slightest moisture
will close them up again, and their purpose having been served in the
distribution of the spores, they are no longer required. If a number of
spores be spread out on a glass-slip under the microscope and, while
watching, a bystander breathes upon them, they immediately respond,
are set in motion, presenting a curious appearance, but as soon as
the hydroscopic effect has passed off they return to their previous
condition. These spores can be mounted in a cell with a movable cover,
and made to exhibit the same effect over and over again.
[Illustration: Fig. 309.--Equisetum giganticum.
_a._ Fragment of stem showing mode of branching out; _b._ Cone or
spike of fructification; _c._ Scale detached from cone; _d._ Spore
with elastic filaments; _e._ Vertical section of stem; _f._ Transverse
section showing hexagonal cells.]
The vascular tissue of the Equisetaceæ (Fig. 309, _e_, _f_) shows
them to be of a higher grade than the ferns. More recently discovered
Horse-tails, of Brazil, grow to a gigantic size, but even these are
comparatively small when compared with the Calamites, and other fossil
Equisetaceæ of the coal measures and new red sandstone. They all
require a calcareous flinty soil for growth. A spring water-course
making its way to the sea, as in the Chines of the Isle of Wight, is
very favourable, the author having gathered them more than once in
Bramble Chine.
Nearly allied to ferns is a little group of small aquatic plants, the
Rhizocarpeæ (pepperworts), which either float on the water or creep
along shallow bottoms. These are chiefly curious from having two
kinds of spores produced from separate sporanges; smaller and larger
“microspores” undergoing progressive sub-division without the formation
of a distinct prothallium; each cell giving origin to an antherozoid,
a generative process said to belong exclusively to flowering plants,
corresponding indeed to the pollen grains of higher plants.
Structure of Phanerogamiæ or Flowering Plants.
The two great divisions of the vegetable kingdom are known as
Cryptogamia and Phanerogamia. It does not follow, however, that there
is any abrupt break between the two, as will appear from the context.
Although it is customary to speak of the flowering plants as a higher
grade of life, yet there is an intermediary class of Phanerogamiæ in
which the conspicuous parts of the generative system partake of a
condition closely resembling those of the higher Cryptogamiæ, observed
in Gymnosperms, Coniferæ, and Cycadæ. So it may be said the distinctive
character of the former is that of reproduction by seeds rather than
flowers. The progress of botanical science during the latter half of
the Victorian reign has been quite as remarkable as that of histology;
while the comparative physiology and morphology of plants have
perhaps advanced even more rapidly because the ground was newer. The
consequence is that the specialisation of botanical science has been
brought about con-currently with a more comprehensive nomenclature.
The chief cause in this instance of modern specialisation is utility.
“If we look at the great groups of plants from a broad point of view,
it will be seen that the fungi and the phanerogams occupy public
attention on other grounds than do the algæ, mosses and ferns. Algæ are
especially a physiologist’s group, employed in questions on nutrition,
reproduction, and cell division and growth. The Bryophyta and
Pteridophyta, are, on the other hand, the domain of the morphologist
concerned with such questions as the alternations of generations and
the evolution of the higher plants.
“Fungi and phanerogams, while equally or even more employed by
specialists in morphology and physiology, appeal widely to general
interest, and evidently so on the ground of utility. Without saying
that this enhances the importance of either group, it certainly
attracts scientific attention to them. However, the histology of the
minute cell, in addition to its importance from an academical point of
view, has a special interest for the microscopist.”
It would be impossible to find anything more remarkable in histology
than the detailed agreement in the structure and behaviour of the
nucleus in the higher plants and the higher animals, an agreement which
is conspicuously manifest in those special divisions which take place
during the maturation of the sexual cells.
So with regard to the question of “alternation of generations.” We
have known since the important discoveries of Hofmeister that the
development of a large part of the vegetable kingdom involves a regular
alternation of two distinct generations, the one which is sexual being
constantly succeeded, so far as the normal cycle is concerned, by the
other which is asexual. This alternation is most marked in the mosses
and ferns. In the Bryophyta the ordinary moss or liverwort plant is the
sexual generation of the ovum, which, when fertilised, gives rise to
the moss-fruit, and represents the asexual stage. The latter is once
more seen to form spores from which the sexual plant is again developed.
In the Pteridophyta the alternation is equally regular, but the
relative development of the two generations is totally different,
the sexual form being the insignificant prothallus, while the whole
fern-plant, as we ordinarily know it, is the asexual generation.
The thallus of some of the lower Bryophyta is quite comparable with the
prothallus of a fern, so as regards the sexual generation there is no
difficulty in seeing the relation of the two classes; but when we come
to the asexual generation or sporophyte the case is totally different.
There is no appreciable resemblance between the fruit of any of the
Bryophyta and the plant of any vascular Cryptogam.
“It is now known that in the higher plants a remarkable numerical
change takes place in the constituents of the nucleus of the cell
shortly before fertilisation. In angiospermous plants a reduction of
the chromosomes occurs shortly before differentiation of the sexual
cells. Thus, in the case of the lily, fertilisation is not the simple
fusion of nuclear bodies. These spheres are seen to fuse in pairs, and
then by position to determine the plane of first cleavage of the ovum;
agreeing, in fact, closely with what is observed to take place in the
animal kingdom.”
In the higher grades of plants it will be evident that the several
tissues that compose their bodies are not found in the root, stem, and
leaf without definite order and purpose, but that they are grouped
into systems for the performance of different kinds of work. In all
flowering plants at least three different systems may be clearly
distinguished. These are the epidermal or boundary tissue system,
the fundamental or ground tissue system, and the fibro-vascular or
conducting system. All three systems of tissue originate from meristem
cells, located at the growing point of the stem and root.
Although these systems characterise the higher types of plants, the
elementary tissues (represented in Plate XIII. and in other figures)
enter alike into the several component parts of nearly all plants.
The stem, the branch, and the root, are alike constituted of an outer
coating which affords a mechanical support, and once formed takes no
further share in the economy of the plant, excepting that of assisting
to convey fluid from the roots to the branches and leaves, an action
more of a capillary nature than vital. The nourishment of the plant is
brought about by other material structures, as the pith, the cortex,
the cambium, and so forth, all of which greatly assist in the formative
process. The woody portion of the plants is especially concerned in
furnishing support to the softer pulpy textures, while the tissues
of leaves and flowers are chiefly composed of cells compactly held
together by protoplasmic or albuminoid matter. Water, of course, enters
largely into the constituents of all plants. Beneath the epidermis is
another layer of importance, the parenchymatous, which becomes more
or less solid with the growth of the pith and cellular wall. In the
pulpy substance of some leaves the epidermis presents a thin lamina
of palisade-tissue, the bulk of the mesophyll consisting of spongy
parenchyma or sclerenchymatous fibres (seen in Fig. 310), which also
serve to show the disposition of the several layers about to be brought
under notice.
_Development of the Tissue Systems._--In the growing plant the
embryonic cells soon become differentiated into three primary meristem
layers, known as dermatogen, periblem, and phloem, from which are
developed respectively the primary cortex, epidermis, and the stele or
vascular cylinder. The dermatogen forms the outermost layer of cells at
the growing point, and when present always develops into true epidermal
tissue. In stems the dermatogen is always single-layered, while in
roots it consists of several layers and develops a many-layered
epidermis.
[Illustration: Fig. 310.--Section of Leaf of Piper.
_c._ Cortex; _ep._ Epidermis; _pal._ Palisade-tissue; _scl f._
Sclerenchymatous fibres of pericycle; _o._ Oil gland.]
The periblem occurs immediately beneath the dermatogen, forming a
hollow cylinder of tissue, which surrounds the phloem. From the
periblem is developed the fundamental tissue of the primary cortex.
When no dermatogen is present in the growing-point (stems of vascular
cryptogams) the external layer of the periblem develops cells which
perform epidermal functions. The phloem occupies the centre of the
growing-point, and consists of a solid mass of somewhat elongated
cells. From the phloem are developed the fibro-vascular and fundamental
tissues of the vascular-cylinder or stele.
[Illustration: PLATE XIII.
ELEMENTARY PLANT TISSUES.]
_Epidermal or Boundary Tissue System._--This system constitutes the
external covering of the plant, and is commonly called the epidermis.
It includes, besides the ordinary epidermal cells, the guard-cells
of the stomata and water pores, the plant hairs or trichomes, and
the epidermal or external glands. The epidermal tissues are chiefly
protective in function, serving to prevent excessive evaporation from
the interior tissues of the plant.
[Illustration: Fig. 311.
_a._ Epidermis, reticulated ducts, and conjunctive palisade cells; _b._
Vertical section of alder root, woody layer, and boundary ducts.]
In stems the external layer of cells, whatever its origin, is known as
the epidermis, while in roots it is called the epiblema. The epidermis
usually consists of a single layer of cells, but in some cases it is
two or three-layered, as in the leaves of figs and begonias.
In land plants the epidermis is usually strongly cutinised, while in
submerged plants it is never cutinised. The epidermis of land plants
is also often waxy, the wax occurring on the surface as minute grains,
rods or flakes, constituting the so-called bloom of leaves and fruits,
and giving to them their glaucous appearance. Chlorophyll bodies are
usually absent from the ordinary epidermal cells of land plants, while
they commonly occur in the epidermal cells of aquatic plants.
Ordinary epidermal cells are usually thin-walled and transparent, and
contain a nucleus and colourless watery protoplasm, but are destitute
of both chlorophyll-bodies and starch-grains.
The external layers of the outer walls constitute the cuticle of the
plant, while the internal layers and the radial and inner walls are
composed of cellulose. The cells of the epidermis are always very
compactly arranged, having their walls so closely adherent that the
intercellular spaces are entirely obliterated except at the stomata and
water-pores.
[Illustration: Fig. 312.
1. Vertical section of leaf of _Iris germanica_; _a, a._ Elongated
cells of the epiderm; _b._ Stomata cut through longitudinally; _c,
c._ Green cells of the parenchyma; _d, d._ Colourless tissue of the
interior of the leaf. 2. Portion of leaf torn from its surface; _a._
Elongated cells of the cuticle; _b._ Cells of the stomata; _c._ Cells
of the parenchyma; _d._ Limiting wall of the epidermic cell; _e._
Lacunæ or openings in the parenchyma corresponding to the stomata.]
There are exceptions to this rule, as, for example, in _Cinchona
calisaya_, which shows no trace of epidermis, this being replaced by
a corky layer of tubular cells. Where this occurs in a plant to any
extent, the whole of the outer tissues are displaced, and the bark
consists exclusively of phloem tissues. This, although of constant
occurrence in _C. calisaya_, is not so common in other species, as _C.
succirubia_, the middle structure of which consists of parenchyma in
which appear more or less numerous isolated store-cells, and when these
are absent there is a formation of rhytidoma and displacement of the
tissues containing the store-cells and ducts. The chlorophyll of _C.
succirubia_ is very marked, and its spectrum presents seven distinct
absorption bands.
The epidermal system of plants in general includes other tissues
than those already named, as the guard-cells of the stomata, the
water-pores, plant-hairs or trichomes, and the external or epidermal
glands, all of which are but modifications of ordinary epidermal tissue.
_The Stomata or Breathing Pores_ are apertures in the epidermal which
lie over large intercellular spaces (Fig 312, 2, _b_). These are
usually bordered by two modified epidermal cells, called guard-cells.
Stomata are formed in the following manner: A young epidermal cell
divides into two equal portions by the formation of a septum across its
middle, each half developing into a guard-cell; the septum now splits
lengthwise and separates the guard-cells, leaving an aperture or stoma
between them.
In the higher plants the guard-cells of the stomata are crescent-shaped
and occur in pairs, the concave sides of the cells facing each other
with the aperture between, while in mosses the stomata possesses but a
single annular guard-cell which surrounds the aperture. The guard-cells
of stomata usually contain chlorophyll-bodies in addition to the
ordinary protoplasm. They have the power of increasing or diminishing
the size of the aperture under the influence of light and moisture,
thus regulating the amount of evaporation from the internal tissues of
the plant.
_Water Pores or Water Stomata_ are apertures in the epidermis, similar
in structure to ordinary stomata, but differ from them both in function
and distribution. Water-pores excrete water in the form of drops, and
have their guard-cells fixed and immovable. They always occur at the
ends of vasal bundles, and are found on the margin and at the apex of
leaves.
_Plant Hairs or Trichomes_ are modified epidermal cells prolonged
externally, and may be either unicellular or multicellular. Each hair
consists of a basal portion, or foot, which is embedded among the
ordinary epidermal cells, and an apical portion or body, which is
prolonged externally. Ordinary epidermal hairs are usually thin-walled,
the inner layers of the wall being composed of cellulose, while the
outer layer is more or less strongly cutinised. The walls may become
hardened by deposits of lime-salts or silica. Sometimes the cells
become glandular and secrete oily, resinous, or irritating matters, as
in stinging-nettle hairs (Plate XIII., No. 19), when they are known as
glandular hairs. The development of resin-passages may be observed in
transverse sections of the stem of the ivy (_Hedera helix_) cut from
a young succulent stem, and mounted in glycerine. The resin is seen
scattered through the cortex and pith, and in the soft bast which lies
outside the cambium in various stages of development, starting from a
group of four cells without intercellular spaces.
Root hairs spring from the epiblema and are never cutinised, but are
frequently more or less mucilaginous. The root-hairs are the principal
absorbing organs of the plant, and are confined to the younger
roots, occurring just above their tips. Root-hairs are never present
in aquatic plants, and are absent from the roots of certain of the
Coniferæ. It is a curious fact with regard to bell-heather growing in
higher latitudes, that the plants possess a peculiar root structure as
a protection against droughts. In most of them the sustentation of life
depends upon the formation of a number of long thin filaments on their
roots resembling root-hairs, which penetrate the root, forming nodular
masses within it. These filaments belong to a fungus entirely parasitic
to the root, and yet different from a common parasite, inasmuch as the
plant in this way obtains so much of its nourishment, and when the
fungus is not present, or is removed, the plant can no longer live
on a peaty soil. The leaf-blade of the coarse moorland grass Nardus
is likewise endowed with a singular property--that of rolling up
cylindrically and spreading out again to adapt itself to the dry and
wet weather of the moorlands of Scotland.
[Illustration: Fig. 313.
_a._ Section of the testa of Gourd Seed, showing communicating cells
filled with colouring matter; _b._ Section of stem of Clematis, three
pores separated and more highly magnified; _c._ Transverse section of
same, showing medullary rays.]
_Fundamental or Ground Tissue System._--This system constitutes the
groundwork of plants, and is the system through which the vasal bundles
are distributed. The fundamental tissues are composed largely, though
not wholly, of parenchyma, and are chiefly concerned in the metabolic
work of plant life.
Ground tissue includes, besides ordinary parenchyma, collenchyma,
selerenchymatous parenchyma, fibrous tissue, cork, laticiferous
and glandular tissues. To the fundamental system also belongs the
chlorophyll cells of leaves, the thin-walled cells of the pith and
medullary rays, the cells of the cortex of stems and roots, and most of
the soft cellular tissues in all plants.
The lower plants consist almost entirely of fundamental tissue. In
the herbaceous forms of the higher plants the ground tissues largely
predominate, while in woody plants they are present in much smaller
proportion, the vascular tissues being the most abundant. In aquatic
plants generally, the fundamental tissues constitute the principal
system.
The hypoderma occurs immediately beneath the epidermis, and consists of
several layers of cells. A collenchymatous hypoderma is found in the
stems and petioles of most herbaceous dicotyls, and frequently occurs
next the mid-rib of leaves, where it forms a strengthening tissue. A
sclerenchymatous hypoderma occurs either as a continuous layer beneath
the epidermis, as in the stems of some ferns, _Pteris aquilina_, and in
leaves of the pine; or it may form numerous isolated strands beneath
the epidermis, as in the stems of horsetails and in certain Umbelliferæ.
[Illustration: Fig. 314.
_a._ Tangential section of _Taxus baccata_ (Yew), showing the woody
fibre; _b._ Vertical section of same, spiral fibres, and ducts; _c._
Vertical section of Elm, showing ducts and dotted cells.]
The endodermis is the innermost layer of the extra-stelar fundamental
tissues, and always abuts on the stele or steles. In monocotyls it
marks the boundary between the cortex and the central cylinder, and it
is sometimes spoken of as the nucleus sheath.
In stems the endodermal cells are usually thin-walled and unlignified,
having a suberous thickening band extending round the upper, lower and
lateral surfaces, which in cross-section appears as a black dot on the
radial wall (Fig. 314, _c_.)
According to its position in the stele, the conjunctive tissue is
divided into three principal portions, viz., that portion which invests
the vasal bundles, the pericycle; that portion which lies between the
bundles of the stele, the interfascicular conjunctive tissue; and that
which occupies the centre of the stele, the medullary conjunctive
tissue. The pericycle, formerly called the pericambium, is the
outermost layer of the conjunctive tissue of the stele.
The bundle-sheath of the young stem is more easily recognised than in
the older stem. It is, in fact, a continuous layer of cells, whose
radial walls have a characteristic dark spot on each radial wall. The
bundle-sheath lies immediately outside the vascular bundles, curving
slightly towards the centre of the stem in the spaces between the
bundles. It is more prominent in the stem when very young, as the cells
are then filled with starch granules. This layer of cells will be
readily seen in sections treated with iodine.
In dicotyls and gymnosperms the medullary rays consist essentially
of interfascicular ground tissue. The medullary conjunctive tissue
occupies the centre of the stele, constituting the so-called pith, and
usually consists of parenchymatous cells, but may contain, in addition,
either stone cells, sclerenchyma fibres, laticiferous or glandular
tissues.
_The Fibro-vascular or Conducting Tissue System._--This system
constitutes the fibrous framework of the plant, and is the system by
means of which fluids are conducted from one part of the plant to
another. Its function is partly to give strength and support, but
principally to conduct the crude and elaborated juices to and from
the leaves. It is found only in the higher plants, constituting the
tough and stringy tissues in stems and roots, and the system of veins
in leaves. The fibro-vascular system consists essentially of vascular
tissue (ducts, tracheids, and sieve-tubes), and forms long strands--the
fibro-vascular bundles--which extend vertically through the fundamental
tissues of the plant. The term “fibro-vascular,” as applied to the
conducting system, is not strictly correct, since fibres do not always
accompany the vascular elements, hence this system is often spoken of
as the vascular system, and the bundles as vascular, or more briefly as
vasal bundles.
That the arrangement, and course of the vascular bundles in
dicotyledous stems are connected with those of the leaves is an
obvious fact. It may be seen in sections of Helianthus, but is
more markedly shown in plants with regularly decussate leaves, as
Cerastium, Clematis, &c. Still, the arrangement of the bundles may
differ radically from that of the leaves, and is, to a certain extent,
independent of them. This will be noticed in sections of _Iberis
amara_, where the bundles do not run longitudinally, but in tangential
spirals. These, as Nägeli pointed out, have no direct relation with
the leaves; and he recommends a series of types for investigation, in
which it will be seen how closely the arrangement of the bundles is
connected with the arrangement of the leaves, and the number of bundles
entering the stem from each leaf: _Iberis amara_, leaves alternate,
leaf-trace with one bundle; Lupinus, leaves alternate, leaf-trace with
three bundles; Cerastium, leaves opposite, leaf-trace with one bundle;
Clematis, leaves opposite, leaf-trace with three bundles; Stachys,
leaves opposite, leaf-trace with two bundles.
[Illustration: Fig. 315.
1. Transverse section of the stem of Cedar, showing xylem or wood; 2.
Section of stem of Conifer, the phloem and zones of annual growth; 3.
Section of an Ivory Nut, cells, and radiating pores; 4. Section of the
outer or ligneous portion of same, with radiating cells.]
The connection of the leaf and stem will be best seen by cutting
longitudinal sections through a _young node_ of Helianthus, so as to
include the median plane of the leaf, or of both leaves if opposite
to each other, as they often are; steep them in dilute potash for
twenty-four hours and mount in glycerine. A medium power will serve
for their examination. The course of the vascular bundles will appear
dark through the more transparent parenchyma. The continuity of the
tissues of the stem and petiole if followed will be seen to have no
definite boundary between the two parts; the bundles from the petiole
pass into the stem, and no bundle of the upper internode lies in the
same vertical plane as that which enters from the petiole between two
successive bundles of the vascular ring.
Every complete _vasal bundle_ consists of xylem or wood and phloem or
bast.
The former consists essentially of trachery tissue (ducts and
tracheids), and may contain in addition both wood fibres and wood
parenchyma. The phloem or bast consists essentially of sieve tissue,
and usually contains some ordinary parenchyma. In angiosperms
companion-cells always accompany the sieve-tubes in the phloem, while
in gymnosperms they are absent.
According to the relative positions of the xylem and phloem elements,
there are two principal kinds of conjoint bundles--the collateral and
the concentric. Of these again there are three varieties, but the
experiments with leaves bring out parallel facts; that in ordinary
stems the staining of the wood by an ascending coloured liquid is due,
not to the passage of the coloured liquid up the substance of the wood,
but to the permeability of its ducts and such of its pitted cells as
are united into regular canals; and the facts showing this at the same
time indicate with tolerable clearness the process by which wood is
formed, for what in these cases is seen to take place with dye may be
fairly presumed to take place with sap.
Taking it, then, as a fact that the vessels and ducts are the channels
through which the sap is distributed, the varying permeability of their
walls, and consequent formation of wood, is due to the exposure of the
plant to intermittent mechanical strains, actual or potential, or both,
in this way. If a trunk, a bough, shoot, or a petiole is bent by a gust
of wind, the substance of its convex side is subject to longitudinal
tension, the substance of its concave side being at the same time
compressed. This is the primary mechanical effect. The secondary is
when the tissues of the convex side are stretched, and also produce
lateral compression. In short, the formation of wood is dependent
upon transverse strains, such as are produced in the aerial parts of
upright plants by the action of the wind.
[Illustration: Fig. 316.--Termination of Vascular System.
1.--Absorbent organ from the leaf of _Euphorbia neriifolia_. The
cluster of fibrous cells forming one of the terminations of the
vascular system is here embedded in a solid parenchyma.
2.--A structure of analogous kind from the leaf of _Ficus elastica_.
Here the expanded terminations of the vessels are embedded in the
network parenchyma, the cells of which unite to form envelopes for them.
3.--End view of an absorbent organ from the root of a turnip. It is
taken from the outermost layer of vessels. Its funnel-shaped interior
is drawn as it presents itself when looked at from the outside of this
layer, its narrow end being directed towards the centre of the turnip.
4.--Shows on a larger scale one of these absorbents from the leaf of
_Panax Lessonii_. In this figure is clearly seen the way in which the
cells of the network parenchyma unite into a closely-fitting case for
the spiral cells.
5.--A less-developed absorbent, showing its approximate connection with
a duct. In their simplest forms these structures consist of only two
fenestrated cells, with their ends bent round so as to meet. Such types
occur in the central mass of the turnip, where the vascular system is
relatively imperfect. Besides the comparatively regular forms of these
absorbents, there are forms composed of amorphous masses of fenestrated
cells. It should be added that both the regular and irregular kinds are
very variable in their numbers: in some turnips they are abundant, and
in others scarcely to be found. Possibly their presence depends on the
age of the turnip.
6.--Represents a much more massive absorbent from the same leaf, the
surrounding tissues being omitted.
7.--Similarly represents, without its sheath, an absorbent from the
leaf of _Clusia flava_.
8.--A longitudinal section through the axis of another such organ,
showing its annuli of reticulated cells when cut through. The cellular
tissue which fills the interior is supposed to be removed.]
In concentric bundles one of the elements, either the xylem or the
phloem, occupies the centre, and is more or less surrounded by the
other, as seen in Fig. 310. Meristem tissue is never present, hence
concentric bundles are always closed. They, however, occur in the stems
of most ferns, and are always surrounded by a pericycle and endodermis,
and should be regarded as steles. Concentric bundles with a central
phloem occur in the rhizomes of some monocotyles, as Calamus, Iris,
Convallaria, &c.
[Illustration: Fig. 317.--Vertical section of Sugar-cane Stem showing
parachyma and crystalline cells, × 200 diameters.]
_The Stele, or Vascular Cylinder_, is developed from the phloem of
the growing plant, and consists of one or more vasal bundles imbedded
in fundamental tissue, the whole being enclosed by a pericycle and an
endoderm. The typical _stele_ includes all the tissues evolved by the
endodermis, which, however, forms no part of the vascular cylinder
itself, but merely surrounds it. The pericycle is always the outermost
layer of the tissues of the stele, while the endodermis is the
innermost layer of the extra-stelar tissues.
The arboreus type of stem can be best followed by making sections
of a twig of the elm (_Ulmus campestris_), which will be found to
be cylindrical hirsute, green or brown according to age, the latter
colour being due to the formation of _cork_. Small brown excrescences
are scattered over its surface; these are termed _lenticels_. The cork
will be seen to lie immediately below the epidermis, and to consist
of cubical cells, with little or no cell contents; they are arranged
in radial rows, without intercellular spaces. The walls of these cork
cells will stain yellowish-brown with Schultze’s solution. Treat a
thin section with sulphuric acid and the walls will swell out and
gradually lose their sharpness of outline, with the exception of the
cuticularised outer wall of the epidermis and the _cork_. This material
is occasionally found developed in the twigs of the elm, so that it can
be separated as thick radial plates of tissue.
“By comparing sections of twigs cut of various ages, the following
information may be gleaned: That cork cambium, or _phellogen_, appears
as a layer of cortical cells below the epidermis, and that these divide
parallel to the surface of the stem. The result of successive divisions
in this direction is the formation of secondary tissue, which develops
externally as cork, internally as phelloderm. The true cork cambium
consists of only a single cell in each radial row, from which, by
successive division, all these secondary tissues are derived--_i.e._,
cambium of vascular bundles. As stems grow older, layers of cork appear
successively further and further from the external surface; not only
the cortex, but also the outer portions of the phloem are thus cut off
from physiological connection with the inner tissue. The term _bark_
is applied to tissues thus cut off, together with the cork which forms
the physiological boundary. The stem of Vitis affords a good example of
such successive layers of cork.”
[Illustration: Fig. 318.
1. Laticiferous Tissue; 2. Vertical section of a Leaf of the
India-rubber Tree, with a central gland; 3. Vertical cast of spiral
tubes of Opuntia.]
For the study of _sieve-tubes_ take the vegetable marrow, in which they
are of extraordinary size. Cut transverse sections of the stem and
stain with eosin, and mount them in glycerine. The general arrangement
will be seen to differ from that of most other herbaceous plants.
Below the epidermis a thick walled band of sclerenchyma with lignified
walls will be seen distinct from the vascular bundles, which readily
take a stain. The vascular bundles are separate and distinct, and the
structure of the bundle is abnormal, there being in each a separate
central mass of xylem, with the phloem masses lying, the one central,
the other in the peripheral side. Between the xylem and the phloem
masses is the cambium layer. The structure being the same in both will
serve for the study of the punctate sieve-plates; these are readily
stained with eosin, as shown in Sach’s text-book.
_Laticiferous Tissues_ (Fig. 318).--In cutting sections of latex care
must be taken to at once transfer them to alcohol so as to prevent the
flow of the latex from the cells, otherwise the laticiferous vessels
will be much less easily traced. The better method is to plunge the
root of the dandelion (_Leontodon taraxacum_), after cleaning, into
alcohol, and there let it remain until it has become hardened; then cut
thin tangential sections from the phloem, and longitudinal sections
through the cambium, and mount them in potash and glycerine. The
laticiferous vessels appear circular in the transverse sections with
brown contents; these are distributed in groups round the central
xylem. Observe in such sections the presence of sphere crystals of
inulin. These are formed quite irrespective of the cell-walls.
Laticiferous cells are readily seen in the cortex of _Euphorbia
splendens_, cut just outside the vascular ring. Long tubes will be seen
to run through the cortical parenchyma, with thick cellulose walls and
granular contents. These are the laticiferous cells, the branching of
which distinguishes them from the preceding structure. Included in the
granular contents are starch grains of a peculiar dumb-bell form.
_Leaf or Petiole._--The general morphology of leaf tissue is
essentially the same as that of the stem from which it proceeds. In the
typical monostotic stem of Phanerogamæ each leaf receives a portion
of the stele or central cylinder of the stem. Such portion is termed
a meristele, and may be either entire or split up into a number of
schizosteles.
The microscopical structure of leaves should be studied in the whole
organ, and by the aid of isolating elements. The whole or portion of a
leaf should be soaked in chloral hydrate solution; this will render it
transparent, whereby the internal structure can be studied as a whole.
Sections should be prepared from fresh leaves, or dried ones softened
by soaking in water. Cut them transversely, both in the direction of
the mid-rib and at right angles to it. This is best effected by placing
the material between two pieces of elder pith or fresh carrot. Sections
of the whole are made and transferred to a dish of water. Leaf sections
are easily made for examination by macerating the leaves in solution
of caustic potash varying in strength from one to five per cent. The
epidermis on both sides may be detached, and the elements of the
mesophyll and vascular bundles isolated for separate examination.
Potassium permanganate proves to be a useful reagent. A weak solution
causes the protoplasmic structures to swell up, thus assisting in the
observation of the structure of the chromatophores. This solution may
also be employed as a macerating fluid. Beautiful preparations are
obtained in this way of the sieve-tubes of Vitis.
Special structural peculiarities are to be observed in the leaves of
various plants in which the epidermis consists of more than a single
layer of cells (_e.g._, the leaves of Ficus, Peperaceæ, Begoniaceæ,
&c.), cystoleths in the cells of the epidermis of Urtica; glandular
structure in Ruta, Psorales; the coriaceous leaves of the Cherry
Laurel, and the cylindrical leaves of Stonecrop (_Sedum acre_).
_Reproductive Organs._--The development of the rudiments of flowers is
of an extremely interesting nature, and the complete flower should be
carefully studied. Median sections are best suited for the purpose.
In the large majority of plants the calyx is developed first, then
the corolla, and next the stamens. Preparations should be made from
materials hardened in alcohol, or first fixed with a strong solution of
picric acid and then hardened in alcohol.
_Pollen-grains._--Microspores are found lying free in sections made
of the reproductive organs; these may be transferred to a glycerine
fluid and examined under a high power. They are mostly spherical, with
granular protoplasmic contents, in which with much difficulty two
nuclei can be made out. Mount and examine, as types of the various
forms of granules, the pollen of Helianthus, Althœa, Cucurbita,
Ænothera, Orchis, Mimosa, Tulipa, &c. Mount any of these pollen-grains
in a weak solution of cane-sugar (about five per cent.), examine with
a high power, and note the configuration of their walls with a medium
power under polarised light. If transverse sections be made from very
young buds, the development of the anther and the pollen may be traced.
The material should be preserved in strong alcohol, and the sections
treated with equal parts of alcohol and glycerine, and exposed in a
watch-glass that the alcohol may evaporate. By this method sections may
be prepared for illustrating the formation of the _tapetum_, special
mother-cells, and division of the nucleus.
[Illustration: Fig. 319.--Pollen Grains.
A. Pollen-grain of Clove-pink; B. Poppy; C. Passion-flower (_Passiflora
cœrulea_); D. _Cobœa scandens_.]
_Starch Granules._--One of the most universally distributed materials
found in plants is starch composed of two substances, _granulose_,
which constitutes by far the largest part, and a skeleton of
_farinose_. It is only the former of these that stains blue with
iodine solutions; the latter partially assumes a brownish colour. The
structure of starch granules is not of equal density throughout; the
hilum or nuclear portion is most conspicuous, around which the rest
of the material is deposited in layers, indicative of stratification.
The several layers next to the hilum are less dense than those
farthest from it. The position of the hilum determines the form of the
grain, a few being rounded, others oval or elongated. The grain also
contains different proportions of water; this conveys the appearance
of concentric lines or curves about the nucleus. The latter is more
conspicuous in the potato starches, as seen in Plate XIII., Nos.
6-15. Starch grains, in nearly all cases, are formed by the agency
of proteid bodies, either chloroplasts or amyloplasts, and under the
action of sunlight are gradually broken up and employed in the process
of growth. There are some plants, however, notably the Compositeæ, in
which another carbohydrate, _inulin_, takes the place of starch from
the first, and is used as a reserve food material. For this reason we
look in vain for starch in the cells of Inula, Taraxacum, &c. From the
whole group of fungi starch is absent; this seems to explain the fact
that chlorophyll, or colouring matter, is rarely met with in the fungi,
hence their inability to utilize, like green plants, carbon-dioxide as
food.
[Illustration: Fig. 320.--Swollen Potato Starch, after the application
of potassium hydrate. (Magnified 210 diameters.)]
The tissues which most commonly contain starch, or which contain it
in largest quantity, are those of the parenchymatous series, though
it sometimes occurs in the latex of laticiferous tissues, and even in
ducts and tracheids. In the stems of Dicotyledons it occurs chiefly
in the parenchyma of the middle and inner bark, in the medullary ray
cells, and in the cells of the pith. In the roots of these plants it
has a similar distribution, being for the most part confined to the
middle or inner bark and the medullary rays, pith not being present in
these organs. In succulent stems and roots, of course, it also commonly
occurs in the xylem tissues of the fibro-vascular bundles.
A study of the various kinds of starches is important, since
this material is very largely used as an adulterant. Other than
microscopical means of detecting frauds are practically useless;
assaying is tedious and expensive, while the microscope is always
available and at hand. The limits of variation should be studied in
starches from the same species of plants; the variations are not very
wide, but in most cases characteristic, so that the discrimination is
at all times an easy task. The reagents required are simply iodine and
dilute potassium hydrate, aided by polarised light.
[Illustration: Fig. 321.
_a a a._ Granules and cells of cocoa; _b b b._ Arrowroot,
_Tous-les-mois_; _c c c._ Tapioca starch. (Magnified 300 diameters.)]
The starch grains of the potato are the best to study in the first
instance on account of their large size (Fig. 320).
In arrowroot starch (Fig. 321) the stratification is almost as distinct
as in that of the potato; the grains much resemble each other. Although
somewhat smaller, the grains of arrowroot are more uniform in size. The
starches are much used as an adulterant of drugs and various articles
sold as cocoas.
Wheat-starch (Fig. 322) consists of circular flattened grains varying
much in size, the central nucleus and stratification of which are very
difficult to distinguish.
In the smaller starches the hilum becomes more indistinct, and without
stratification, as in rice-starch, the latter being angular in shape.
The hilum in other leguminous plants forms a longitudinal cleft; white
rye-starch exhibits distinct cracks. Compound grains are occasionally
met with, as in the oat. In Plate XIII. will be found small groups of
starches taken under the same medium power for the sake of comparison.
In the microscopical examination of starches first use a 2/3-inch or a
1/2-inch and then a 1/6-inch objective.
[Illustration: Fig. 322.
_a._ Husks of Wheat-starch, swollen by
reagents and heat; _b._ A portion of cellulose; _c._ Rice-starch,
magnified 420 diameters.]
The bran of the husk of wheat when broken by grinding is seen to
be composed of two coats of hexagonal cells, the outer of which
is detached by the roasting process. The hexagonal cell layer is,
however, so little altered as to be perfectly distinguishable under the
microscope. Thus even a small admixture of roasted corn with coffee
or chicory can be detected without much difficulty. As to whether
starch granules should be regarded as crystalline or colloid bodies,
a difference of opinion still prevails. There are, however, reasons
for believing that the polarisation effects produced by starch grains
are not due to crystalline structure but to stress or strain, of the
same nature as the polarisation of glass when it is subject to strain.
The polarising phenomena are precisely such as would be induced in
any transparent solid composed of layers, the inner of which being
kept in a state of stress by the compression exerted by the outer
layers. Moreover, when by use of a swelling reagent, such as caustic
potash solution, the outer wall of the starch is made to expand by the
imbibition of water, the polarisation effects immediately disappear.
Were the solid particles of crystal thus forced apart by water each
particle would still exhibit polarisation phenomena.
Want of space will not permit me to further enlarge upon other
micro-chemical substances that enter into the composition of plants;
as, for example, the oil secreting glands. These when present take the
place of starch. There is, however, one product among the cell contents
of plants of some interest to the microscopist--those extremely
fine crystals known as _raphides_, composed of calcium-phosphate
and oxalate. Mr. Gulliver insisted upon the value of raphides as
characteristic of several families of plants. Schleiden states that
“needle-formed crystals, in bundles of from twenty to thirty in a cell,
are present in almost all plants,” and that so really practical is the
presence or absence of raphides, that by studying them he has been able
to pick out pots of seedling Onagraceæ, which had been accidentally
mixed with pots of other seedlings of the same age, and at that period
of growth when no other botanical character would have been so readily
sufficient.
If we examine a portion of the layers of an onion (Plate XIV., No. 3),
or a thin section of the stem or root of the garden rhubarb (No. 4), we
shall find many cells in which either bundles of needle-shaped crystals
or masses of a stellate form occur, not strictly raphides.
Raphides were first noticed by Malpighi in Opuntia, and subsequently
described by Jurine and Raspail. According to the latter observer,
the needle-shaped or acicular are composed of phosphate, and the
stellate of oxalate of lime. There are others having lime as a basis,
in combination with tartaric, malic, and citric acids, all of which
are destroyed by acetic acid; others are soluble in many of the fluids
employed in mounting. These crystals vary in size from the 1/40th
of an inch, while others are as small as the 1/1000th. They occur in
all parts of the plant; in the stem, bark, leaf, petals, fruit, root,
and even in the pollen, and occasionally in the interior of cells. In
certain species of aloe, as _Aloe verrucosa_, we are able to discern
small silky filaments; these are bundles of the acicular form of
raphides, and probably, as in sponges, act as a skeleton support to the
internal soft pulpy mass.
[Illustration: PLATE XIV.
STELLATE AND CRYSTALLINE TISSUE OF PLANTS.]
In portions of the cuticle of the medicinal squill (_Scilla maritima_)
large cells are found full of needle-shaped crystals. These cells,
however, do not lie in the same plane as the smaller cells of the
cuticle. In the cuticle of an onion every cell is occupied either by an
octahedral or a prismatic crystal of calcium oxalate. In some specimens
the octahedral form predominates; in others, even from the same plant,
the crystals are prismatic and arranged in a stellate form, as in that
of the grass (_Pharus cristatus_). (Plate XIV., No. 6.)
Raphides of peculiar figure are found in the bark of certain trees. In
the hickory (_Carya alba_) may be observed masses of flattened prisms
having both extremities pointed. In vertical sections from the stem of
_Elæagnus angustifolia_, numerous raphides of large size are embedded
in the pith, and also found in the bark of the apple-tree, and in elm
seeds, every cell containing two or more minute crystals.
In the Graminaceæ, especially the canes; in the _Equisetum hyemale_,
or Dutch rush; in the husk of rice, wheat, and other grains, silica in
some form or other is abundant. Some have beautifully-arranged masses
of silica with raphides. The leaves of _Deutzia scabia_, No. 7, are
remarkable for their stellate hairs, developed from the cuticle of both
their upper and under surfaces; forming most interesting and attractive
objects examined under polarised light. (Plate VIII., No. 173.)
Silica is found in the structure of Rubiaceæ both in the stem and
leaves, and, if present in sufficient thickness, depolarises light.
This is especially the case in the glandular hairs on the margins of
the leaves. One of the order Compositæ, a plant popularly known as the
“sneezewort” (_Archillæ ptarmica_), has a large amount of silica in the
hairs found about the serratures of its leaves.
All plants are provided with hairs; some few with hairs of a defensive
character. Those in the _Urtica dioica_, commonly called the
_Stinging-nettle_, are glandular hairs, developed from the cuticle, and
contain an irritating fluid; in other hairs a circulation is visible:
examined under a power of 100 diameters, they present the appearance
seen at Plate XIII., No. 19.
[Illustration: Fig. 323.
A. Cotton; B. Fibres of Flax; C. Filaments of Silk; D. Wool of Sheep.]
The fibrous tissue of plants is of great value in many manufactures.
It supplies material for our linens, cordage, paper, and other
industries. This tissue is remarkable for toughness of fibre, and
exhibits an approach to indestructibility, in the use it is put to in
connection with the electric light. It is of importance, then, to be
able to distinguish it from other fibres with which it is often mixed
in various manufactures. Here the use of the microscope is found of
considerable importance. In flax and hemp, in which the fibres are
of great length, there are traces of transverse markings at short
intervals. In the rough condition in which flax is imported into
this country, the fibres have been separated, to a certain extent,
by a process termed _hackling_, and further subjected to hackling,
maceration, and bleaching, before it can be reduced to the white silky
condition required by the spinner and weaver, and finally assumes the
appearance of structureless tubes, Fig. 323 B. China-grass, New Zealand
flax, and some other plants produce a similar material, but are not so
strong, in consequence of the outer membrane containing more _lignine_.
It is important to the manufacturer that he should be able to determine
the true character of some of the textures employed in articles of
clothing; this he may do by the aid of the microscope. In linen we find
each component thread made up of the longitudinal, unmarked fibres of
flax; but if cotton has been mixed, we recognise a flattened, more or
less rounded band, as in Fig. 323 A, having a very striking resemblance
to hair, which, in reality, it is; since, in the condition of elongated
cells, it lines the inner surface of the pod. These, again, should
he contrasted with the filaments of silk, Fig. 323 C, and also of
wool, Fig. 323 D. The latter may be at once recognised by the zigzag
transverse markings on its fibres. The surface of wool is covered with
furrowed and twisted fine cross lines, of which there are from 2,000
to 4,000 in an inch. On this structure depends its _felting_ property,
in judging of fleeces, attention should be paid to the fineness and
elasticity of the fibre--the furrowed and scaly surface, as shown by
the microscope, the quantity of fibre in a given surface, the purity
of the fleece, upon which depend the success of the scouring and
subsequent operations.
In the mummy-cloths of the Egyptians flax only was used, whereas the
Peruvians used cotton alone. By the many improvements introduced into
manufacturing processes, flax has been reduced to the fineness and
texture of silk, and even made to resemble other materials.
[Illustration: Fig. 324.
1. Woody Fibre from the root of the Elder, exhibiting small pores; 2.
Woody fibre of fossil wood, showing large pores; 3. Woody fibre of
fossil wood, bordered with pores and spiral fibres; 4. Fossil wood from
coal.]
_Fossil Plants._--It is well known that the primordial forests furnish
a number of families of plants familiar to the modern algæologist. The
cord-like plant, _Chorda filium_, known as “dead men’s ropes,” from
its proving fatal at times to the too adventurous swimmer who gets
entangled in its thick wreaths, had a Lower Silurian representative,
known to palæontologists as _Palæochorda_, or ancient chorda, which
existed, apparently, in two species,--a larger and a smaller. The
still better known _Chondrus crispus_, the Irish moss, or Carrageen
moss, has likewise its apparent, though more distant representative,
in chondritis, a Lower Silurian algal, of which there seems to exist
at least three species. The fucoids, or kelpweeds, appear to have also
their representatives in such plants as _Fucoides gracilis_, of the
Lower Silurians of the Malverns; in short, the Thallogens of the first
ages of vegetable life seem to have resembled in the group, and in at
least their more prominent features, the algæ of the existing time.
And with the first indications of land we pass from the thallogens to
the acrogens--from the seaweeds to the fern-allies. The Lycopodiaceæ,
or club-mosses, bear in the axils of their leaves minute circular
cases, which form the receptacles of their spore-like seeds. And when
high in the Upper Silurian system, and just when preparing to quit it
for the Lower Old Red Sandstone, we detect our earliest terrestrial
organisms, we find that they are composed exclusively of those little
spore-receptacles.
The existing plants whence we derive our analogies in dealing with the
vegetation of this early period contribute but little, if at all, to
the support of animal life. The ferns and their allies remain untouched
by the grazing animals. Our native club-mosses, though once used in
medicine, are positively deleterious; horsetails (_Equisetaceæ_),
though harmless, so abound in silex, which wrap them round with a
cuticle of stone, that they are rarely cropped by cattle; while the
thickets of fern which cover our hill and dell, and seem so temptingly
rich and green in their season, scarce support the existence of a
single creature, and remain untouched, in stem and leaf, from their
first appearance in spring until they droop and wither under the frosts
of early winter.
The flora of the coal measures was the richest and most luxuriant,
in at least individual productions, with which the fossil botanist
has formed an acquaintance. Never before or since did our planet bear
so rank a vegetation as that of which the numerous coal seams and
inflammable shales of the carboniferous period form but a portion of
the remains--the portion spared, in the first instance, by dissipation
and decay, and in the second by denuding agencies. Nevertheless almost
all our coal--the stored-up fuel of a world--is not, as it is often
said to be, the product of destroyed forests of conifers and flora
of the profuse vegetation of the earliest periods in the history of
our globe. Later investigations show that our coal measures are the
compressed accumulations of peat-bogs which, layer by layer, have
sunken down under the superimposed weight of the next. The vertical
stems of coniferous trees became imbedded by a natural process of
decay, and were subsequently overwhelmed in the erect position in
which they are found. The true grasses scarcely appear in the fossil
state at all. For the first time, amid the remains of a flora that
seems to have had but few flowers--the Oolitic ages--do we detect, in
a few broken fragments of the wings of butterflies, decided traces
of the flower-sucking insects. Not, however, until we enter into the
great Tertiary division do these become numerous. The first bee makes
its appearance in the amber of the Eocene, locked up hermetically
in its gem-like tomb--an embalmed corpse in a crystal coffin--along
with fragments of flower-bearing herbs and trees. Her tomb remains
to testify to the gradual fitting up of our earth as a place of
habitation for creatures destined to seek delight for the mind and
eye, as certainly as for the proper senses, and in especial marks
the introduction of the stately forest trees, and the arrival of the
charmingly beautiful flowers that now deck the earth.[62]