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 Fig. 183.—Saccharomyces mycoderma.
The “Ferment of Wine” (Saccharomyces ellipsoideus) produces
wine in the juice of grapes. Uncultivated yeast-cells are always
present on grapes; an addition of this species to the “must” is not
necessary to secure fermentation. A large number of other
“uncultivated” yeast-cells appear in breweries mixed with the
cultivated ones, and cause different tastes to the beer (S.
pastorianus, etc.). S. ludwigii, found, for instance, on the slimy
discharge from Oaks, produces abundant cell-chains on cultivation.
S. apiculatus is very frequently met with on all kinds of sweet fruits, it
has orange-like cells. S. mycoderma has cylindrical cells, often
united together in chains (Fig. 183): it forms a whitish-gray mass
(“fleur de vin”) on wine, beer, fruit-juice, etc., standing in bottles
uncorked or not entirely filled. It is thought that this Fungus causes
decomposition and oxydises the fluid in which it is found, but it
cannot produce alcoholic fermentation in saccharine liquids, and it
does not form endospores; hence it is uncertain whether it is true
Saccharomyces.

Fig. 184.—Oidium lactis: a branched hypha commonly met with; b a

hypha lying in milk and producing aerial hyphæ which give rise to oidia; c
a branch giving rise to oidia, the oldest (outermost) oidia are becoming
detached from one another; d a chain of divided cells; e germinating oidia
in different stages (slightly more magnified than the other figures).
The “Dry-yeast” used in baking white bread is “surface-yeast.” In
leaven, a kneaded mixture of meal, barm and water, which is used
for the manufacture of black bread, Saccharomyces minor is
present, and a species allied to this produces alcoholic fermentation
in dough with the evolution of carbonic acid, which causes the dough
to “rise.”
2. Oidium-forms. Of many Fungi only the Oidium-forms are
known, which multiply in endless series without employing any
higher form of reproduction. Oidium lactis (Fig. 184) is an imperfectly
developed form which frequently appears on sour milk and cheese. It
can produce a feeble alcoholic fermentation in saccharine liquids.
Thrush or aphthæ (O. albicans) appears as white spots in the
mouths of children. Several similar Oidium-forms are parasites on
the skin and hair of human beings, and produce skin diseases, such
as scurvy (O. schoenleinii) and ringworm (O. tonsurans).
3. Mycorhiza. These Fungi, which have been found on the roots
of many trees and heath-plants, particularly Cupuliferæ and
Ericaceæ, consist of septate hyphæ, and belong partly to the
Hymenomycetes, partly to the Gasteromycetes. It has been shown
that the Mycorhiza enters into a symbiotic relationship with the roots
of higher plants.
 DIVISION II.

MUSCINEÆ (MOSSES).
In this Division a well-marked alternation of generations is to be
found. The development of the first or sexual generation
(gametophyte),[16] which bears the sexual organs, antheridia and
archegonia, commences with the germination of the spore, and
consists, in the Liverworts, of a thallus, but in the true Mosses of a
filamentous protonema, from which the Moss-plant arises as a lateral
bud. The second or asexual generation (sporophyte), developed
from the fertilised oosphere, consists of a sporangium and stalk.
The sexual generation, the gametophyte. The protonema in the
Liverworts is very insignificant, and not always very sharply
demarcated from the more highly developed parts of the nutritive
system. In the true Mosses the protonema is well-developed, and
consists of a branched, alga-like filament of cells, the dividing cell-
walls being always placed obliquely. In the parts exposed to the light
it is green, but colourless or brownish in those parts which are
underground (Fig. 186). The protonema is considered to be a lower
form of the stem, and grows in the same manner by means of an
apical cell; at its apex it may directly develope into a leaf-bearing
stem, or these arise from it as lateral branches (Fig. 186 k).
The more highly differentiated part of the vegetative system, the
“Moss-plant,” which is thus developed from the protonema, is in the
“thalloid” Liverworts generally a dichotomously-branched thallus
without any trace of leaf-structures (Fig. 194); in Marchantia (Fig.
197) and others, scale-like leaves (amphigastria) are found on the
under surface. The higher Liverworts and the Leafy-Mosses are
differentiated into a filamentous, ramified stem with distinct leaves
arranged in a definite manner, resembling the stem and leaves of the
higher plants (Figs. 186, 195, 200).
True roots are wanting, but are biologically replaced by rhizoids.
These are developed on the stems or thallus: in the Liverworts they
are unicellular, but in the Leafy-Mosses generally multicellular and
branched. In the latter group they are considered identical with the
protonema, and may become true protonema, and new plants may
be developed from them (Fig. 186 b).

Fig. 186.—A Lower portion of a Moss-plant with rhizoids (r), one of which bears
a reproductive bud (b). The dotted line indicates the surface of the ground; the
portions projecting above this become green protonema (p); k is a young Moss-
plant formed on one of these. B Germinating spore of Funaria hygrometrica, with
exospore still attached. C, D Older stages of the protonema.
The internal structure of the sexual generation is very simple. The
leaves in nearly all cases are formed of a single-layered plate of
cells; in the Leafy-Mosses, however, a midrib is very often formed,
and sometimes, also, marginal veins; and along these lines the
leaves are several layers of cells in thickness. The stem is
constructed of cells longitudinally elongated, the external ones of
which are narrower and sometimes have thicker walls than the more
central ones. Vessels are not found, but in several Mosses there is in
the centre of the stem a conducting strand of narrow, longitudinal
cells, which represents the vascular bundle in its first stage of
development. This strand contains elements for conveying water as
well as sieve-tubes. Stomata are entirely wanting in the sexual
generation of the Leafy-Mosses; they are found in a few Liverworts
(Marchantia), but their structure is not the same as in the higher
plants.
Vegetative reproduction takes place by gemmæ or buds
which arise on the protenema, the rhizoids, the thallus, or the shoots,
and become detached from the mother-plant; or else the protonema
and the older parts of the plant simply die off, and their branches
thus become independent plants. This well-developed vegetative
reproduction explains why so many Mosses grow gregariously. In
certain Marchantiaceæ special cupules, in which gemmæ are
developed, are found on the surface of the thallus (Fig. 197 A, s-s).
Again, protonema may also arise from the leaves, and thus the
leaves may act as reproductive bodies. Certain Mosses nearly
always reproduce vegetatively, and in these species the oospheres
are seldom fertilised.
Fig. 187.—Marchantia
polymorpha: a mature
antheridium.
 Fig. 188.—Spermatozoids.
The first generation bears the sexual organs; both kinds are
found either on the same plant (monœcious), or on separate plants
(diœcious). In the thalloid Liverworts they are often situated on the
apex of small stems (gametophores), springing from the surface of
the thallus. In the Leafy-Liverworts and true Mosses the leaves
which enclose the sexual organs often assume a peculiar shape,
and are arranged more closely than the other leaves to form the so-
called “Moss-flower.” The male sexual organs are called antheridia.
They are stalked, spheroid, club- or egg-shaped bodies whose walls
are formed of one layer of cells (Fig. 187), enclosing a mass of
minute cubical cells, each one of which is a mother-cell of a
spermatozoid. The spermatozoids are self-motile; they are slightly
twisted, with two cilia placed anteriorly (Fig. 188), while posteriorly
they are generally a trifle club-shaped, and often bear at that part the
remains of the cytoplasm, the spermatozoid itself being formed from
the nucleus. In the presence of water the ripe antheridium bursts,
and its contents are ejected; the spermatozoids, being liberated from
their mother-cells, swarm about in the water in order to effect
fertilisation.
 Fig. 189.—Marchantia polymorpha. A A young, and B a ripe
archegonium with open neck. C An unripe sporangium enclosed
by the archegonium a: st the stalk; f the wall of the sporangium.
Elaters are seen between the rows of spores.
The female sexual organs are termed archegonia. They are flask-
shaped bodies (Fig. 189), the lower, swollen portion (venter) having
a wall, in most cases from 1–2 cells thick, enclosing the oosphere
(Fig. 189 B, k): the long neck is formed of tiers of 4–6 cells,
enclosing a central row of cells—the neck-canal-cells (Fig. 189 A).
When the archegonium is fully developed, the walls of the neck-
canal-cells become mucilaginous and force open the neck of the
archegonium. The mucilage thus escapes, and, remaining at the
mouth of the archegonium, acts in a somewhat similar manner to the
stigma and conducting tissue of a carpel, by catching and conducting
the spermatozoids to the oosphere (Fig. 189 B, m), with whose cell-
nucleus they coalesce. With regard to the formation of the oosphere,
it may further be remarked that the lower part of the archegonium
originally encloses the so-called “central cell”; but shortly before the
archegonium is ripe, this cuts off a small portion, the ventral-canal-
cell, which lies immediately beneath the neck, and the larger, lower
portion becomes the oosphere.
The organs mentioned here, antheridia and archegonia, are present in the
Cryptogams (Pteridophyta) and the Gymnosperms. They have always the same
fundamental structure, but with slight modifications of detail. These plants are
therefore known as the Archegoniata.
The fertilisation of the Mosses cannot be effected without water.
Rain and dew therefore play a very important part in this process,
and for this end various modifications of structure are found.
 Fig. 190.—Andreæa rupestris. Longitudinal
section through a sporangium at the time when
the mother-cells of the spores are dividing: p
pseudopodium; f foot; v vaginula; h neck; c
columella; w wall of the sporangium; e external
row of cells; s the spore-sac; t the spore-
mother-cells; r the calyptra with the neck of
archegonium (z).
 Fig. 191.—Andreæa rupestris. Transverse section
through a ripe sporangium. In the middle is seen the four-
sided columella, surrounded by the numerous spores, drawn
diagrammatically. Surrounding them is seen the wall of the
sporangium, whose outer layer of cells is thickened and
coloured. The layer of cells is unthickened in four places (x),
indicating the position of the clefts (see Fig. 193).
Among the sexual organs, paraphyses—filamentous or club-
shaped bodies—are to be found.
The asexual generation, the sporophyte (Moss-fruit or
sporogonium). As the result of fertilisation the oosphere surrounds
itself with a cell-wall, and then commences to divide in accordance
with definite laws.[17] The embryo (Fig. 189 C) produced by these
divisions remains inside the wall a-a of the archegonium (Figs. 190,
199 D, E), and developes into the sporogonium, which remains
attached to the mother-plant, often nourished by it, as if the two were
one organism. The lower extremity of the sporogonium, the foot
(Figs. 190 f; 199 D), very often forces its way deep down into the
tissue of the mother-plant, but without an actual union taking place.
The central portion of the sporogonium becomes a shorter or longer
stalk (seta), while the sporangium itself is developed at the summit.
At a later stage, during the formation of the spores, the sporangium
very often assumes the form of a capsule, and dehisces in several
ways characteristic of the various genera (Figs. 192, 193, 194, 195,
200). The basal portion of the archegonium grows for a longer or
shorter period, forming a sheath, the calyptra, in which the capsule is
developed, but eventually it ceases to enlarge, and is then ruptured
in different ways, but quite characteristically, in each group.
Anatomically, the asexual generation is often more highly
differentiated than the sexual; thus, for instance, stomata are present
on the sporangia of the true Mosses, but are absent in the sexual
generation.
As the capsule developes, an external layer of cells—the
amphithecium—and an internal mass—the endothecium—are
differentiated. As a rule the former becomes the wall of the capsule
while the latter gives rise to the spores. In this Division, as in the
Pteridophyta, the name archesporium (Fig. 190 t) is given to the
group of cells inside the sporangium which gives rise to the mother-
cells of the spores. The archesporium is in general a unicellular
layer; in Sphagnum and Anthoceros it is derived from the most
internal layer of the amphithecium, but with these exceptions it arises
from the endothecium, usually from its most external layer. In the
true Mosses and in Riccia only spore-mother-cells are produced
from the archesporium, but in the majority of the Liverworts some of
these cells are sterile and become elaters (cells with spirally
thickened walls, Figs. 196, 189), or serve as “nurse-cells” for the
spore-mother-cells, which gradually absorb the nutriment which has
been accumulated in them. In Anthoceros, and almost all the Leafy-
Mosses, a certain mass of cells in the centre of the sporangium
(derived from the endothecium) does not take part in the formation of
the archesporium, but forms the so called “column” or “columella”
(Figs. 190, 191).
 The spores arise in tetrads, i.e. four in each mother-cell, and are
arranged at the corners of a tetrahedron, each tetrahedron assuming
the form of a sphere or a triangular pyramid. The mature spore is a
nucleated mass of protoplasm, with starch or oil as reserve material.
The wall is divided into two layers: the external coat (exospore)
which is cuticularized and in most cases coloured (brown, yellowish),
and the internal coat (endospore), which is colourless and not
cuticularized. On germination the exospore is thrown off, the
endospore protrudes, and cell-division commences and continues
with the growth of the protonema (Fig. 186, B-D).
 Fig. 192.—Andreæa petrophila. A ripe
sporogonium: a an archegonium which has
been raised with the pseudopodium; p the foot;
b the neck; d-e the dark-coloured portion of the
sporangium, whose outer cell-walls are
considerably thickened; c-c the thin-walled
portions where the dehiscence occurs; o the
 lower extremity of the spore-sac; f calyptra; g
the apex of the sporangium. (Mag. 25 times.)

Fig. 193.—Andreæa petrophila. An empty

capsule; the calyptra has fallen off. (Mag. 25
times.)
The morphological explanation which Celakovsky has given of the
sporogonium, and which is not at all improbable, is, that it is homologous with an
embryo consisting of a very small stem-portion and a terminal spore-producing
leaf. This will be further explained in the introduction to the Flowering-plants (p.
236).
In the Liverworts the young sporogonium lives like a parasite,
being nourished by the sexual generation (only in Anthoceros has it
a slight power of assimilation). In the Leafy-Mosses, on the other
hand, with regard to the power of assimilation, all transitions are
found from abundant assimilation (Funaria, Physcomitrium) to almost
complete “parasitism” (Sphagnum, Andreæa). In the majority of the
operculate Mosses the sporogonium has a more or less perfect
system of assimilation, and is able itself to form a large portion of the
material necessary for the development of the spores, so that it
chiefly receives from the sexual generation the inorganic substances
which must be obtained from the soil. The more highly developed the
assimilative system of the sporogonium, the more stomata are
present.
Apospory. In some operculate Mosses it has been possible to obtain a
protonema with small Moss-plants from the seta, when severed from its Moss-
plant, and grown on damp sand.
The Mosses are the lowest plants which are provided with stem
and leaf. They are assigned a lower place when compared with the
higher Cryptogams, partly because there are still found within the
Division so many forms with a mere thallus, partly because typical
roots are wanting and the anatomical structure is so extremely
simple, and partly also because of the relation between the two
generations. The highest Mosses terminate the Division, the
Muscineæ and Pteridophyta having had a common origin in the
Algæ-like Thallophyta.
They are divided into two classes:—
Hepaticæ, or Liverworts.
Musci frondosi. True Mosses or Leafy-Mosses.

Class 1. Hepaticæ (Liverworts).

The protonema is only slightly developed. The remaining part of
the vegetative body is either a prostrate, often dichotomously-
branched thallus, pressed to the substratum (thalloid Liverworts),
with or without scales on the under side (Figs. 194, 197); or a thin,
prostrate, creeping stem, with distinctly-developed leaves, which are
borne in two or three rows (Figs. 195, 198), viz., two on the upper
and, in most cases, one on the under side. The leaves situated on
the ventral side (amphigastria) are differently shaped from the others
(Fig. 198 a), and are sometimes entirely absent. In contradistinction
to the Leafy-Mosses, stress must be laid on the well-marked
dorsiventrality of the vegetative organs; i.e. the very distinct contrast
between the dorsal side exposed to the light and the ventral side
turned to the ground. Veins are never found in the leaves.
The ventral part of the archegonium (calyptra) continues to grow
for some time, and encloses the growing embryo, but when the
spores are ripe it is finally ruptured by the sporangium, and remains
situated like a sheath (vaginula) around its base. The sporangium
opens, longitudinally, by valves or teeth (Fig. 194, 195, 197 b), very
rarely by a lid, or sometimes not at all. A columella is wanting
(except in Anthoceros, Fig. 194); but on the other hand, a few of the
cells lying between the spores are developed into elaters (Fig. 196),
i.e. spindle-shaped cells with spirally-twisted thickenings, which are
hygroscopic, and thus serve to distribute the spores. (They are seen
in Fig. 189 C, not yet fully developed, as long cells radiating from the
base of the sporangium. They are wanting in Riccia).

Fig. 194.—Anthoceros lævis (nat. size): K-K

capsules.
 Fig. 195.—Plagiochila asplenioides: a
unripe, and b an open capsule; p involucre.
The ventral edge of each leaf is higher than its
dorsal edge, and covered by the dorsal edge of
the next one.