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 Fig. 232.—Lycopodium clavatum. A
tetrahedral spore seen from above, where the
three borders join; and a tetrad of bilateral
spores, still lying in the mother-cell.
The fertile leaves are collected upon definite regions of the stem.
They are either similar to the barren ones, and then the fertile
portions of the stem pass gradually, without any break, into the
barren portion (L. selago); or they differ from the barren leaves, and
are then collected into special apical cones (Fig. 230 a). The spores
are tetrahedral or bilateral (Fig. 232).
About 100 species, chiefly tropical.
Five species of Lycopodium are found in Great Britain. L. clavatum and L.
selago are common in mountainous districts. L. annotinum is common in the
Highlands of Scotland. The other genus of the order is Phylloglossum, with one
species, P. drummondi (Australia, Tasmania, and New Zealand), a small plant only
a few centimetres high, with two tubers, and about eleven linear leaves at the base
of the stem which is terminated by a cone of sporophylls.—Fossil Lycopodiaceæ
in the Carboniferous period.
Officinal: “Lycopodium,” the spores of L. clavatum.
 Family 2. Psilotaceæ. The sporangia are placed on the apex of short, two-
leaved stems, as 2–3, seldom four, small capsules. Small herbs, with angular
stems; leaves small, simple, and one nerved. Only four species.—Psilotum
(Madagascar, Moluccas, Sandwich Islands, etc.) is destitute of roots, their place
being supplied by special underground stems which bear a few modified leaves,
very much reduced, especially when buried deeply in the soil. Three species.—
Tmesipteris (Australia), one species.

Sub-Class 2. Selaginelleæ (Heterosporous Lycopodinæ).

Micro-and macrospores. The prothallia are very much reduced,
especially the male; the female does not leave the spore. The leaves
are ligulate.

Fig. 233.—Germination of the microspores of Selaginella:

A the spore rendered transparent, seen from above. In the
interior is seen the prothallium (f), and the first divisions of
the antheridium (a, b, c, d); in B the spore-wall is removed
and all the spermatozoid-mother-cells formed; in C, the
microspore has opened and the spermatozoids and the
mother-cells are escaping together.
The sexual generation. In the MICROSPORES are formed: (1) a
very small “vegetative” cell, representing the vegetative part of the
prothallium (f in Fig. 233 A, B), and (2) a cell many times larger and
which divides into a number (4–8) of primordial cells, each of which
divides into four spermatozoid-mother-cells, though all of these may
not develope spermatozoids. On germination, when the spore-wall is
ruptured, the spermatozoids and spermatozoid-mother-cells are
ejected into the water.
The SPERMATOZOIDS in Selaginella are elongated and club-
shaped, with two cilia (Fig. 234); but in Isoëtes lacustris they are
spirally-twisted threads which differ from all other spermatozoids by
having a bunch of cilia at each end; the other species of Isoëtes
have cilia only at the anterior end.
The macrospores. Shortly after the macrospores have been set
free, or in Selaginella, while still enclosed in the sporangium of the
mother-plant, they germinate and soon become filled with the cellular
tissue of the prothallium, and even in Selaginella the archegonium
begins to be formed before the rupture of the spore-cell-wall has
commenced (Fig. 235 A).

Fig. 234.—Spermatozoids of Selaginella: b with a

remnant of cytoplasm.
 Fig. 235.—Macrospore of Selaginella: A longitudinal
section, before the rupture of the wall, six weeks after being
sown. The endosperm (e) has not yet filled the entire
chamber. Cell-formation is still proceeding in the lower part
of the spore. The endosperm and prothallium (f f) are
separated by a distinct line (diaphragm). B Germinating
macrospore seen from outside: s wall of the spore; æ
archegonia.

Fig. 236.—Archegonia of Selaginella: A unripe, in

longitudinal section; c the central cell; k neck-canal-cell,
which is wedged in between the two-storied neck-cells; B
ripe; u ventral canal-cell; C seen from above, open. It will be
noticed that the neck is formed of two tiers of four cells each.
 The archegonia are constructed on the same plan as those of
the other Archegoniatæ, but are quite embedded in the prothallium
(Figs. 235 æ, 236).
The asexual generation varies very much in the different orders.

Fig. 237.—Isoëtes lacustris

(slightly diminished): st the stem;
r roots; b leaves.
 Fig. 238.—Isoëtes lacustris. Longitudinal
section through the base of the leaf with a
microsporangium. The edge of the groove, in
which the microspangium is placed, is
continued as a thin covering which envelopes
the sporangium. The inferior edge of the ligular
groove (L) forms a lip (J); t sterile cell-rows
(trabeculæ) which divide the sporangium into
compartments; l vascular bundle.
 Fig. 239.—Selaginella inæqualifolia. Cone in
longitudinal section; microsporangia are seen
on the left side, macrosporangia on the right
(most frequently each with four macrospores).
Order 1. Isoëtaceæ (Quill-worts). The only known genus,
Isoëtes (Quill-wort), has an extremely short, tuberous, unbranched
stem with very short internodes (Fig. 237). The stem is remarkable
as being the only one among the Vascular Cryptogams which
increases in thickness (see page 202). The meristematic cells are
situated round the axial cylinder, and form, especially,
parenchymatous tissue in two or three directions, giving rise to 2–3
grooves in which the dichotomously-branched roots are produced.
The leaves are arranged spirally in a close rosette. They are awl-
shaped and have at the base a semi-amplexicaul sheath, with a
groove (fovea), in which a sporangium is situated (Fig. 238). The
ligule is a foliar outgrowth from the upper edge of the groove.—The
macrosporangia (each with a number of macrospores), are
situated on the outer leaves, the microsporangia (Fig. 238), on the
inner ones. Between each cycle of fertile leaves there are a number
of imperfect or barren ones as in the case of the female plant of
Cycas. The spores are liberated by the decay of the sporangium.
The two kinds of sporangia develope at the commencement in the
same way. The archesporium is, at first, a hypodermal layer of cells
which grow out in the direction perpendicular to the surface of the
leaf, and divide by a number of walls parallel to this direction,
forming a sporogenous mass of cells. Some of the cell-rows of this
sporogenous mass lose their rich protoplasmic contents, and are
arrested in their growth; thus incomplete divisional walls of sterile
cells, “trabeculæ” arise in the sporangium, dividing it into a number
of compartments one above the other (Fig. 238 t). (The trabeculæ,
according to Goebel, play the same part as the nutritive cells of the
sporangium of Riella; the tapetal cells, as in the Ferns, are in a great
measure dissolved at a later period.) The sporogenous cell-rows, in
the microsporangia, give rise to a large number of spore-mother-
cells, but in the macrosporangia only one spore-mother-cell, with
tapetum, is developed from each fertile archesporial cell.
The two native species, and several others, are aquatic plants,
the remaining species are land plants, or are amphibious. About 50
species. In temperate and tropical regions.—Fossil species in the
Tertiary period.
Order 2. Selaginellaceæ. This order contains only one genus,
Selaginella. The stem, in the majority of species, is dorsiventral, long
and slender, and apparently branches dichotomously, but in reality
monopodially, with well developed lateral shoots. The leaves are
small, round, or ovate, in the majority of species arranged in whorls
of two leaves each; these whorls, however, are not decussate, but
are considerably inclined towards each other, an arrangement by
which four rows of leaves are produced, each whorl having one large
and one small leaf. The two leaves in each whorl are of unequal
size, the smaller one being placed on the upper surface and the
larger on the lower surface of the stem (Fig. 240). Some species
have spirally-arranged leaves, more resembling the arrangement in
the Lycopodiums.
 The FERTILE LEAVES most frequently differ from the barren ones,
and are collected into spike-like cones (a kind of flower; Fig. 239).
Micro-and macrosporangia are found in the same cone (Fig. 239).
Each sporangium arises from a group of superficial cells of the stem,
directly over the leaf on which it will be situated later on. Each
sporangium has a hypodermal, unicellular archesporium, and
contains a layer of tapetal cells; these are dissolved later, when the
spores are ripe, and not before as in the Ferns. In the very early
stages of their development, the micro-and macrosporangia are
precisely similar, and the differences between them arise later on. In
the microsporangium all the spore-mother-cells divide, and each
forms four tetrahedrically-arranged microspores (Fig. 204); but in the
macrosporangium only four macrospores are formed, by the division
of a single mother-cell, while the remaining spore-mother-cells are
aborted. It is rarely that the macrosporangia contain 2 or 8
macrospores.

Fig. 240.—Selaginella martensii: s

lower leaves; r upper leaves.
 For the GERMINATION OF THE SPORES, see pages 228, 229. The
prothallium arises in the macrospore (f-f, in Fig. 235 A), probably by division of the
meniscus-shaped protoplasmic mass, which is marked off at the apex of the spore;
primordial cells are thus formed which later on are surrounded by a cell-wall. In six
to seven weeks after sowing, the spore-wall is ruptured by the growing prothallium,
which already has developed archegonia (Fig. 235 œ-œ). The prothallium so
formed does not occupy the entire cavity of the spore, but four to five weeks after
sowing, the large-celled parenchyma is developed in the lower portion of the spore
by free cell-formation; this has been termed by Pfeffer, “endosperm,” since it is
similar to the endosperm of Flowering-plants. Goebel, however, has termed it
“secondary prothallium,” as the homology with the endosperm of the Angiosperms
is very doubtful.
The FERTILISED OOSPHERE divides into an upper (hypobasal) and
a lower (epibasal) cell; from the latter alone the embryo is developed
with its root, stem, foot, and two cotyledons, and the former gives
rise to an organ which appears in this instance for the first time, but
which occurs in all Flowering-plants, viz. the suspensor. This forces
the embryo down into the “endosperm,” which is entirely or partially
absorbed by the embryo. In the case of the Flowering-plants the
embryo is developed with its longitudinal axis in the elongation of the
suspensor, but in Selaginella the embryo is situated transversely to
it.
Selaginella (300–400 species), is essentially tropical, only one
species living in the North (S. spinulosa), but others grow in Central
and South Europe.
Order 3. Lepidodendraceæ are extinct, tree-like Lycopods, which are found
especially in the Lower and Middle Carboniferous. Vegetatively they are most
nearly related to Lycopodium, but the stem attained much larger proportions
(about eleven metres in height and one metre in thickness), and had a cambium
by which it increased in thickness. It was regularly dichotomous, and closely
studded with spirally-placed leaves, which left behind them peculiar rhombic scars.
The large cones resemble Pine-cones, and bore sporangia much larger than any
which are now produced (the male ones as much as 2 cm.’s in length). The
macrosporangia were situated at the base, and the microsporangia at the apex.
Order 4. Sigillariaceæ. These are, presumably, another group of extinct tree-
like Lycopods (especially in the Middle Carboniferous). The name has been
derived from the seal-like scars, which the fallen leaves have left behind in
longitudinal rows on the grooved stem. The rhizomes of these plants were formerly
termed Stigmaria, and placed in a separate genus.
 Order 5. Sphenophyllaceæ form an entirely extinct group. They do not
definitely belong to any of the three large classes of Vascular Cryptogams, but it is
perhaps best to place them in juxtaposition to these. They were herbaceous plants
with verticillate, wedge-shaped leaves, with nerves branching dichotomously into
equally strong branches. Micro-and macrosporangia were formed in the same
cone; and were situated in the axils of the leaves, as in the Lycopods.
 The Transition from the Cryptogams to the Phanerogams.
All the plants considered in the preceding chapters are included in
the term Cryptogams; all in the following chapters under the head
of Phanerogams (see page 3). Hofmeister’s pioneer works (1851,
Vergleichende Untersuchungen der höheren Kryptogamen, etc.) and
the numerous researches published later by other investigators,
have closed the gap which was formerly thought to exist between
these plants; so that we now, in the series: Bryophyta—Pteridophyta
—Gymnospermæ—Angiospermæ see the expression of a single line
of development in accordance with a definite plan. The forms
through which this gradual development has taken place have in
course of time, however, to a great extent died out, and only single
links of the chain connecting the lowest to the highest still remain.
The alternation of generations, which we found indicated in
certain Thallophytes, can be proved with the greatest clearness in all
the higher Cryptogams, from the Mosses upwards; it is also found in
the Phanerogams, but not in such a pronounced degree, because
one of the generations is so far reduced that it has almost given up
its independence. For the sake of greater clearness, we will begin
with the comparison of the sporophyte, asexual (second) generation.

The asexual (2nd) generation of the Cormophytes.

The asexual generation which follows from the further
development of the fertilised oosphere, is, in the Mosses, only the
sporogonium (according to one theory it is perhaps homologous with
a spore-bearing leaf, situated upon a short stem, see p. 187); in
Filicinæ, Equisetinæ, and Lycopodinæ, on the other hand, it is a
highly developed plant differentiated into stem, leaf, and true root,
and bearing the sporangia on its leaves. The MODIFICATION OF THE
SHOOT is very slight in Filicinæ. The first leaves of the embryo are
very simple in form (Fig. 205), but after a certain age all the leaves
which arise are essentially alike. The fertile leaves do not differ from
the barren ones, and are found associated with them, and their
formation does not limit the growth in length of the stem. It is only in
a few of the true Ferns, and in the Hydropterideæ, that the fertile
leaves differ considerably from the barren ones. A division of labour
in which certain leaves are set apart for nutrition, and others for
reproduction, is found more pronouncedly in the Equisetinæ and
Lycopodinæ, for in these groups, with a few exceptions, the fertile
and barren leaves are very dissimilar; the former are collected in
special ear-like cones, which terminate the further growth of the
short stems on which they are borne. In connection with the cone,
leaves are sometimes developed which form a transition from the
barren to the fertile ones (the “annulus” in Equisetaceæ), and in
these cases the first indication of a flower with perianth or floral-
leaves is to be traced. Among the Cryptogams the highest division of
labour is found in Selaginella and Isoëtes, which have the two kinds
of sporangia borne on different leaves. The division of labour
(modification) is, however, still more pronounced in the
Phanerogams: the leaves which bear the microsporangia (“pollen-
sacs”) have quite different forms from those which bear
macrosporangia (the “nucellus” in the ovule), the former are termed
stamens, the latter carpels; in certain instances, too, there is even a
contrast between the “male plants” and the “female plants.”
Moreover, a division of labour, in a much greater degree, takes place
in the leaves which do not directly take part in reproduction, and it is
thus possible in many plants to draw a sharp line not only between
stamens and carpels, but also between four or five distinct kinds of
leaves, which differ in form, structure, and corresponding functions,
and which appear in regular sequence on the shoot: namely,
between “scale-leaves” and “foliage-leaves,”[21] both of which occur
in the Cryptogams, and the “floral-leaves,” including the bracts and
leaves of the “perianth,” which latter often differ from each other in
form and colour, and are then separated into sepals and petals. The
leaves—stamens and carpels—which bear the sporangia are termed
sporophylls, and the shoot, or extremity of a shoot, whose leaves are
modified into sporophylls, is terminated in its further growth by their
production, and is known as a flower. The flower which is most
completely furnished has calyx, corolla, stamens, and carpels
arranged in this order. If the only sporophylls present are stamens,
then it is said to be a male (staminate) flower, and if only carpels,
then a female (pistillate) flower, and in both these cases the flowers
are unisexual, or diclinous. If stamens and carpels are both present
in the same flower, it is termed hermaphrodite. Diclinous plants in
which the female flowers are situated on one plant, and the male
flowers on another, are termed diœcious; and those in which the
same plant bears the two kinds of flowers are termed monœcious.
When the male, female, and hermaphrodite flowers are found in the
same species, the plant is said to be polygamous.
The sporangia-bearing leaves—Sporophylls. In the Mosses
the asexual generation is only represented by the sporogonium, and
if the theory is correct which considers the sporogonium to be an
embryo consisting of a rudimentary stem and terminal leaf, then the
spores are produced on the leaves in these plants. The sporangia in
the Filicinæ are situated in groups (sori) on the back or on the edge
of the leaves. The number of sporangia in the sorus diminishes very
greatly in the Marattiaceæ and Gleicheniaceæ (three to four in the
latter, Fig. 213). In the Equisetinæ the sporangia are situated in a
small number on the underside of shield-like leaves, and in
Lycopodinæ, singly, in the axils of the fertile leaves, which are alike
and bear either micro- or macrosporangia. In the Phanerogams there
is a great difference between the stamens and carpels.
 Fig. 241.—Cycas: a stamen (nat.
size) seen from the under side; b four
pollen-sacs, not yet open, forming a
“sorus”; c three open pollen-sacs; d a
pollen-grain.
 Fig. 242.—Stamens of
Araucaria (pollen-sacs long and
pendulous).
Fig. 243.—Male flower of Taxus.
 Fig. 244.—A Cross section through a quadrilocular anther in
different stages of development: s the seam where it bursts open; vf
vascular bundle; k connective. B A stamen. C Another stamen seen
from the front (f) and from the back (b).
Stamens. In the lowest Phanerogams (Cycadeæ) there are many
indications of relationship to the Ferns. The stamens are flat and
broad, and have on the back many pollen-sacs (microsporangia)
arranged in small groups (true sori), which even have a small
“placenta,” similar to the one possessed by the Ferns, and open
towards the inside by a longitudinal cleft (Fig. 241, compare Fig.
213). A section of the Coniferæ agree more closely with the
Equisetaceæ, in having a few (three to eight) pollen-sacs arranged
on the underside of more or less shield-like leaves (Figs. 242, 243,
compare with Fig. 224 a, c, d). In the Abietaceæ the number of
sporangia is diminished to two, which are placed also on the lower
side (Fig. 267) of a stamen. The number of pollen-sacs
(microsporangia) in the Angiosperms is nearly always four to each
stamen; they are longitudinal projections which are placed in pairs
on each side of the central line of the stamen, two on the edge, and
the other two generally on the side which is turned inwards; the
pollen-sacs generally dehisce longitudinally (quadrilocular anthers,
Fig. 244). A few, for instance Orchidaceæ and Asclepiadaceæ, have
only two pollen-sacs (bilocular anthers); and in others, such as
Solanum and the Ericaceæ, they open by pores; in Lauraceæ and
Berberidaceæ, by valves. The part of the stamen which bears the
pollen-sacs is termed the anther. Most frequently this is supported by
a stalk known as the filament.

Fig. 245.—A carpel of

Cycas revoluta with 5 ovules
(s), at half to one-third nat.
size.
 Fig. 246.—Carpel with 2 ovules of
Ceratozamia robusta (1/1).
Carpels. The simplest forms of carpels are found in Cycas. In this
genus both the foliage and fertile leaves are pinnate, and hence
present great similarity; the ovules (macrosporangia) are situated on
the margin of the central portion, just as the sporangia are placed on
the edge of the fertile leaf of Ophioglossum (Fig. 245, compare with
Fig. 209). The carpels of the other Cycadeæ present greater
divergence from the foliage-leaves, being peltate, for instance, in
Zamia and Ceratozamia (Fig. 246). The ovules in the Coniferæ are
situated on the upper side and near the base of the ovuliferous
scales, almost in the same position as the sporangia in the
Lycopodinæ (Figs. 269, 272, 273 H, compare Figs. 230, 239). In
Taxus the uninclosed ovule is placed on the apex of a shoot (Fig.
264). In all these plants the ovules are not enclosed by the carpels,
that is, they are not enclosed in chambers formed by the turning in of
the walls of the carpel, and hence the name Gymnospermæ is given
to them. In the higher Flowering-plants, the Angiospermæ, the
ovules are distinctly situated on the edge, the upper surface, or base
of the carpel; but the carpel closes round the ovules which are