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 Order 7. Piptocephalidaceæ. The conidia are formed
acrogenously and in a series, by transverse divisions. The
zygospore arises at the summit of the conjugating hyphæ, which are
curved so as to resemble a pair of tongs. Piptocephalis and
Syncephalis live parasitically on the larger Mucoraceæ.
Sub-Class 2. Oomycetes.
Sexual reproduction is oogamous with the formation of brown,
thick-walled oospores which germinate after a period of rest. Asexual
reproduction by conidia and swarmspores. Parasites, seldom
saprophytes.
The oospores are large spores which are formed from the egg-cell
(oosphere) of the oogonium (oosporangium, Fig. 89, 95). A branch of
the mycelium attaches itself to the oogonium and forms at its apex
the so-called “antheridium” (pollinodium[13]): this sends one or more
slender prolongations (fertilising tubes) through the wall of the
oogonium to the egg-cell.
 Fig. 85.—Empusa muscæ (Fly-mould). I. A fly killed by the
fungus, surrounded by a white layer of conidia. II. The conidiophores
(t) projecting from the body of the fly. Some of the conidia, a few of
 which have developed secondary conidia, are attached to the hairs
(mag. 80 times). III. A perfect hypha. IV. A hypha in the act of
ejecting a conidium (c), enveloped in a sticky slime (g). V. A
conidium which has developed a secondary conidium (sc). VI. A
branched hypha produced by cultivation. VII. A secondary conidium
which has produced a small mycelium (m). VIII. A conidium
germinating on the fly’s body. IX. Mycelium. X. Conidia germinating
like yeast in the fatty tissue of the fly. (III.-VII. and IX. magnified 300
times; VIII. and X. magnified 500 times.)
A fertilisation, a passage of the contents of the antheridium to the egg-cell, has
as yet only been observed in Pythium; in Phytophthora only one small mass of
protoplasm passes through the fertilising tube to the egg-cell; in Peronospora and
the Saprolegniaceæ no protoplasm can be observed to pass through the fertilising
tube, so that in these instances parthenogenesis takes place; Saprolegnia thuretii,
etc., have generally even no antheridia, but nevertheless form normal oospores.
Fertilisation of the egg-cell by means of self-motile spermatozoids is only found in
Monoblepharis sphærica.

A. Asexual reproduction by conidia only.

Family 1. Entomophthorales.
The mycelium is richly branched. The family is a transitional step
to the conidia-bearing Zygomycetes, since the oospores of many
members of this family arise, and are formed, like zygospores.
Order 1. Entomophthoraceæ. Mycelium abundantly developed.
This most frequently lives parasitically in living insects, causing their
death. The conidiophores forming the conidial-layer project from the
skin, and abstrict a proportionately large conidium which is ejected
with considerable force, and by this means transferred to other
insects. These become infected by the entrance of the germ-tube
into their bodies. The spherical, brown resting-spores develope
inside the bodies of insects and germinate by emitting a germ-tube.
Genera: Empusa has a good many species which are parasitic on flies, moths,
grasshoppers, plant-lice. The conidia emit a germ-tube which pierces the skin of
the insect; a number of secondary conidia are then produced inside its body, by
division or by gemmation similar to that taking place in yeast, each of which grows
and becomes a long unbranched hypha, and these eventually fill up the body of
the animal, causing distension and death. Each of these hyphæ projects through
the skin, and abstricts a conidium, which is ejected by a squirting contrivance. The
best known species is E. muscæ (Fig. 85), which makes its appearance
epidemically towards autumn on the common house-fly, and shows itself by the
dead flies which are found on the windows and walls attached by their probosces,
distended wings, and legs. They have swollen abdomen, broad white belts of
hyphæ between the abdominal rings, and are surrounded by a circle of whitish
dust formed by the ejected conidia.—Entomophthora sends out, at definite places,
from the mycelium hidden in the insect’s body, bundles of hyphæ, which serve the
purpose of holding fast the dead insects, the ramifications attaching themselves to
the substratum: the conidiophores are branched, the conidia are ejected by the
divisional walls between the hyphæ and the conidia dividing into two layers, those
which terminate the hyphæ suddenly expanding and throwing the conidia into the
air. E. radicans makes its appearance epidemically on caterpillars.
B. Asexual reproduction by zoospores or conidia.

Family 2. Chytridiales.
In this family the mycelium is very sparsely developed or is
wanting. The entire plant consists principally or entirely of a single
zoosporangium whose zoospores have generally one cilium. The
resting-spores arise either directly from the zoosporangium, which,
instead of forming zoospores, surrounds itself by a thick cell-wall; or
they are formed by the conjugation of two cells (in which case they
are spoken of as oospores). Microscopic Fungi, parasitic on water
plants (especially Algæ) or small aquatic animals, seldom on land
plants.
Order 1. Olpidiaceæ. Without mycelium. Swarmspores and
resting-spores.
In the Olpidieæ, the swarmspores, probably, most frequently form themselves
into a plasmodium (naked mass of protoplasm) which may become a single
zoosporangium or a resting sporangium. Olpidium trifolii occurs in Trifolium
repens.—In the Synchytrieæ the plasmodium emerging from the swarmspores
breaks up either at once, or after a period of rest, into smaller plasmodia, each of
which will become a zoosporangium. Synchytrium anemones is found on
Anemone nemorosa; S. mercurialis on Mercurialis perennis; S. aureum on many
plants, particularly Lysimachia nummularia.
 Fig. 86.—Chytridium lagenula.
Zoosporangium a before, b after the liberation
of the swarmspores.
 Fig. 87.—Obelidium mucronatum: m mycelium;
s swarmspores.
Order 2. Rhizidiaceæ. Mycelium present. Zoospores and resting-
spores.
Chytridium (Fig. 86). Obelidium (Fig. 87) is bicellular; the one cell is the
mycelium, the other the zoosporangium; found on insects. The species of
Cladochytrium are intercellular parasites on marsh plants. Physoderma.
Order 3. Zygochytriaceæ. Mycelium present. Zoospores and
oospores. The latter are the product of the conjugation of two cells
(Fig. 88).
Polyphagus euglenæ on Euglena viridis. Urophlyctis pulposa on species of
Chenopodium.
 Family 3. Mycosiphonales.
The mycelium is bladder-like or branched. Zoospores. Sexual
reproduction by oospores, which are produced in oogonia. The latter
are fertilised, in some forms, by the antheridium.
Order 1. Ancylistaceæ. The entire bladder-like mycelium is used for the
construction of zoosporangia, oogonia, or antheridia. Lagenedium is parasitic on
Spirogyra, etc.
Order 2. Peronosporaceæ. Almost entirely parasites. The
unicellular, often very long and abundantly branched mycelium lives
in the intercellular spaces of living plants, especially in the green
portions, and these are more or less destroyed and deformed in
consequence. Special small branches (suction-organs, “haustoria”)
are pushed into the cells in order to abstract nourishment from them.
Both oospores and conidia germinate either immediately, or they
develope into sporangia with swarmspores, having always two cilia.
Only one oospore is formed in each oogonium; its contents (Fig. 89)
divide into a centrally placed egg-cell and the “periplasm”
surrounding it; this is of a paler colour and on the maturity of the
oospore forms its thick, brown, external covering.
 Fig. 88.—Polyphagus euglenæ. A with smooth, B with thorny oospores; m and f
the two conjugating cells.
 Fig. 89.—Peronospora alsinearum.
Mycelium with egg-cell and antheridium.
 Fig. 90.—Phytophthora infestans (strongly magnified).
Cross section through a small portion of a Potato-leaf (the
under side turned upwards): a the mycelium; b b two
 conidiophores projecting through a stoma; c conidia; e the
spongy tissue of the leaf; g the epidermis.
The Potato-fungus (Phytophthora infestans) is of great interest. Its
thallus winters in the Potato-tuber; other organs for passing the
winter, such as oospores, are not known. When the tuber
germinates, the Fungus-hyphæ penetrate the young shoot and keep
pace with the aerial growth and development of the plant. The
conidiophores emerge through the stomata, especially on the under
side of the leaves; they branch like a tree (Fig. 90), and appear to
the naked eye as a fine mould on the surface of the plant. The
disease soon makes itself known by the brown colouring of those
parts of the plant which are attacked, and by their withering. An
ovoid conidium arises at first by the formation of a dividing wall at the
apex of each branch of the conidiophore (Fig. 90 c c), and
immediately underneath it another is formed, which pushes the first
to one side, and so on. These conidia sometimes germinate directly,
and form a mycelium, but most frequently their protoplasm divides
into many small masses, each of which becomes a pear-shaped
zoospore provided with two cilia (Fig. 91). Water is required for their
germination, and when the ripe conidia are placed in a drop of water
the swarm-cells are formed in the course of about five hours. They
swarm about in rain and dewdrops in the Potato-fields, and are
carried with the water to the Potato-plants and to the tubers in the
soil. The wind also very easily conveys the conidia to healthy Potato-
fields and infects them. The enormous quantity of conidia and
swarm-cells that may be formed in the course of a summer explains
the rapid spreading of the disease; and the preceding makes it clear
why wet summers are favourable to its existence. When the swarm-
cells germinate, they round off, and then surround themselves with a
cell-wall which grows out into the germ-tube, and pierces through the
epidermis of the host-plant (Fig. 92). Having entered the host, a new
mycelium is formed. The potato disease, since 1845, has been
rampant in Europe; it has, no doubt, been introduced from America,
which, it must be remembered, is the home of the Potato-plant.
 Fig. 91.—Phytophthora infestans: a-c conidia detached; in
c the swarm-cells are leaving the mother-cell; d two free-
swimming swarm-cells.

Fig. 92.—Phytophthora infestans. Cross section through a portion of a Potato-

stalk. Two germinating conidia (a, b) piercing the epidermis, and the mycelium
penetrating the cells.
The conidia exhibit various characters which are employed for the separation of
the genera. Pythium is the most simple form. The contents of the terminally-formed
conidia emerge as a spherical mass and divide into swarmspores. P. de Baryanum
lives in the seedlings of many different Flowering-plants, which it completely
destroys.—Phytophthora is distinguished by the circumstance that the sparsely-
branched conidiophores bear, sympodially, chains of conidia. Besides the Potato-
fungus (see above), Ph. fagi belongs to this group; it developes oospores very
abundantly, and does great harm to seedlings of the Beech, Sycamore, and Pine
trees.—Peronospora generally has conidiophores which are repeatedly forked,
and bear a conidium on each of the most extreme ramifications. Many do great
harm to their host-plants. P. viticola, on Vines, and P. nivea, on umbelliferous
plants, have swarmspores, which are absent in the following species of this genus:
P. sparsa, on Roses; P. gangliformis, on composites; P. alsinearum, on Stitchwort;
P. parasitica, on cruciferous plants; P. viciæ, on Vetches and Peas; P. schachtii, on
Beets; P. violacea, on the flowers of Scabiosa; P. radii, on the ray-florets of
Matricaria.—Cystopus (Albugo) has the conidia developed in chains, which form a
cohesive white layer underneath the epidermis of the host-plant. Cystopus
candidus, on cruciferous plants, especially Shepherd’s Purse and Brassica; the
germination commences on the cotyledons, and from this point the mycelium
developes together with the host-plant; C. cubicus, on the leaves of Compositæ.

Fig. 93.—A fly overgrown with

Saprolegnia.
 Fig. 94.—Formation of swarmspores in a Saprolegnia: a germinating
swarmspores.
Order 3. Saprolegniaceæ, Water-Fungi which live as
saprophytes on organic remains lying in water, for instance, on dead
flies (Fig. 93), worms, remains of plants; but they may also make
their appearance on living animals, being frequently found, for
example, on the young trout in rearing establishments.
 Fig. 95.—Oogonium with
two antheridia, Achlya
racemosa.
The thallus is a single, long and branched cell. It has one portion
which serves as root, and lives in the substratum, where it ramifies
abundantly for the purpose of absorbing nourishment; and another
portion projecting freely in the water, and sending out hyphæ on all
sides (Fig. 93). The asexual reproduction takes place by
swarmspores (Fig. 94), which are developed in large sporangia;
these swarmspores generally possess two cilia, and on germination
grow into new plants. The entire protoplasm in the oogonium is
formed into one or more oospheres, without any surrounding
“periplasm.” The oospheres may not be fertilised (p. 100), and then
develope parthenogenetically.
Genera: Saprolegnia, whose swarmspores disperse immediately after having
left the sporangium. S. ferax is the cause of a disease in fish (“Salmon disease”)
and in the crayfish.—Achlya, whose swarmspores accumulate in a hollow ball
before the mouth of the sporangium.—Leptomitus has strongly indented hyphæ,
causing a “linked” appearance. L. lacteus is frequent in the waste matter from
sugar factories.—Monoblepharis deviates from the others by the greater
development of its fertilising process; the oosphere, situated in an open oogonium,
becoming fertilised by self-motile spermatozoids, which are provided with a cilium
at the posterior end.

Class 2. Mesomycetes.
The Mesomycetes are intermediate forms between the
Phycomycetes and the Higher Fungi. In the vegetative organs, and
in the multicellular hyphæ, they resemble the Higher Fungi; the
methods of reproduction, however, show the characters of the
Phycomycetes, namely sporangia and conidiophores of varying size
and with varying number of spores; definite and typically formed asci
and basidia are not present. Sexual reproduction is wanting. The
Hemiasci are transitional between the Phycomycetes and the
Ascomycetes, the Hemibasidii (Brand-Fungi) form the transition to
the Basidiomycetes.
Sub-Class 1. Hemiasci.
The Hemiasci are Fungi with sporangia which, although
resembling asci, yet have not, however, a definite form and a definite
number of spores. Besides endospores, conidia, chlamydospores
and oidia are found.
Order 1. Ascoideaceæ. Ascoidea rubescens forms irregular, reddish-brown
masses in the sap issuing from felled Beeches. It has free sporangia, which
resemble asci in their structure, in the development and ejection, and in the
definite shape and size of the spores. The formation of the sporangia takes place
when the nutriment is nearly exhausted, and resembles that of the conidia, since
they are developed from the end of a hypha which enlarges, and the swelling
becomes separated by a transverse wall. Within the sporangia numerous spores
of a cap-like form are developed, which are set free through an opening at the
apex. Sporangia are formed successively at the apex of the same hypha, the
second commencing to develope as the first is dehiscing. Conidia and sporangia
are not formed simultaneously; the former may be considered as closed
sporangia.
Order 2. Protomycetaceæ. Protomyces pachydermus causes hard swellings
on the stems and leaf-stalks of the Cichorieæ (Taraxacum, etc.). These swellings
consist of chlamydospores (resting-spores), which germinate and become free,
ascus-like sporangia, with numerous small spores. In nutritive solutions the
chlamydospores form conidia with yeast-like buddings. P. macrosporus on
Ægopodium, and other Umbelliferæ.
Order 3. Thelebolaceæ. Thelebolus stercoreus, is found on the dung of deer,
hares, and rabbits, and has closed sporangia, which resemble asci in their shape
and regular construction, and in the ejection of spores. The covering encloses only
one sporangium, even where the sporangia arise close together.
This order, by reason of the covering of the sporangia, forms the
transition from the Hemiasci to the Carpoasci, while the two first
supply an intermediate step to the Exoasci.
Sub-Class 2. Hemibasidii, Brand-Fungi.
The Brand-Fungi (also known as Ustilagineæ) are Fungi with
basidia-like conidiophores, which, however, have not yet advanced
to a definite form or number of conidia. They are true parasites,
whose mycelium spreads itself in the intercellular spaces of
Flowering plants. The mycelium is colourless, quickly perishable, has
transverse walls at some distance from each other (Fig. 96), and
sends out haustoria into the cells of the host-plant.
 Fig. 96.—Entyloma ranunculi. 1. Cross section of a portion of a leaf of
Ficaria permeated by the mycelium; a bundle of hyphæ with conidia
emerging from a stoma; in one of the cells are found four brand-spores. 2.
A brand-spore developed in the middle of a hypha.
It most frequently happens that the germ-tube enters the host-
plant at its most tender age, that is, during the germination of the
seed; the mycelium then wanders about in the tissues of the shoot
during its growth, until it reaches that part of the plant where the
spores are to be formed. The spore-formation takes place in the
same way in all those species whose brand-spores are developed in
the floral parts of the host-plant. Many Brand-Fungi have, however, a
more local occurrence, and the mycelium is restricted to a smaller
area of the leaf or stem. Those organs of the host-plant in which the
brand-spores are developed often become strongly hypertrophied. In
perennial plants the mycelium winters very often in the rhizome.
 Fig. 97.—Doassansia alismatis. 1. A fruit-body, formed by a covering of
oblong hyphæ, which encloses a mass of brand-spores, and is embedded
in the leaf-tissue of the host-plant; 20 times natural size. 2. A germinating
brand-spore, 500 times natural size. 3. Three connected resting-spores,
400 times natural size. 4. Two conidia grown together, 600 times natural
size.
The brand-spores are the winter resting-spores of the Brand-
Fungi. They arise in the tissues of the host-plant, which is often
destroyed, and become free through the rupture of the epidermis;
they are thick-walled, generally brown or violet, and very often
possess warts, spines, or reticulate markings. Fruit-bodies, that is
enclosed organs of reproduction, are found in few genera
(Sphacelotheca, Graphiola; Doassansia, Fig. 97). In Tolyposporium,
Tuburcinia, Thecaphora (Fig. 102), etc., the brand-spores are united
into a ball of spores. On germination the brand-spores behave as
chlamydospores, namely, as the fundament of conidiophores, by
emitting a short germ-tube, i.e. a conidiophore (“promycelium”). The
Ustilaginaceæ (Fig. 99, 2) have a short transversely divided
conidiophore, with laterally developed conidia (comp. the basidia of
the Protobasidiomycetes). The conidiophores of the Tilletiaceæ are
undivided (unicellular promycelia), and bear the conidia terminally,
and so resemble the basidia of the Autobasidiomycetes.

Fig. 98.—Tuburcinia. 1. T. trientalis. Hyphæ, some of which bear conidia at the

apex, forcing themselves out between the epidermal cells on the under side of the
leaf; 320 times natural size. 2. T. trientalis. A ball of spores in which some of the
individual brand-spores are about to germinate; 520 times natural size. 3. T.
primulicola: various forms of conidia (500 times natural size).
In Tilletia, Entyloma, Neovossia, Tuburcinia, the brand-spores germinate and
form basidia-like conidiophores with spindle-shaped conidia; their mycelium, on
the other hand, produces later only single, sickle-shaped conidia, so that two kinds
of conidia are found, as in a few Basidiomycetes. In some species, e.g. Ustilago
hordei, the brand-spores only germinate vegetatively and form a mycelium. In
nutritive solutions (solutions of dung, etc.) where they live as saprophytes, the
brand-spores of many species emit germ-tubes, and on these, yeast-like conidia
are produced by repeated budding, which grow into mycelia only when the nutritive
solution is exhausted. These conidia have not the power of producing alcoholic
fermentation. The very numerous conidia, which are found in the dung of
herbivorous animals, are probably the yeast-conidia of Brand-Fungi. The brand-
spores, which are eaten by animals with the grain and hay, pass into the dung and
without doubt give rise to a very rich multiplication of yeast-conidia.