THE TERM Embryology, in its
widest sense, is applied to the various changes which take place during
the growth of an animal from the egg to the adult condition: it is,
however, usually restricted to the phenomena which occur before birth.
Embryology may be studied from two aspects: (1) that of ontogeny, which deals only with the development of the individual; and (2) that of phylogeny, which concerns itself with the evolutionary history of the animal kingdom. | |
In vertebrate animals the development of a new being can only take place when a female germ cell or ovum has been fertilized by a male germ cell or spermatozoön.
The ovum is a nucleated cell, and all the complicated changes by which
the various tissues and organs of the body are formed from it, after it
has been fertilized, are the result of two general processes, viz., segmentation and differentiation
of cells. Thus, the fertilized ovum undergoes repeated segmentation
into a number of cells which at first closely resemble one another, but
are, sooner or later, differentiated into two groups: (1) somatic cells, the function of which is to build up the various tissues of the body; and (2) germinal cells,
which become imbedded in the sexual glands—the ovaries in the female
and the testes in the male—and are destined for the perpetuation of the
species. | |
Having regard to the main purpose of this work, it is
impossible, in the space available in this section, to describe fully,
or illustrate adequately, all the phenomena which occur in the different
stages of the development of the human body. Only the principal facts
are given, and the student is referred for further details to one or
other of the text-books on human embryology. | |
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1. The Animal Cell |
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All the tissues and organs of the body originate from a microscopic structure (the fertilized ovum),
which consists of a soft jelly-like material enclosed in a membrane and
containing a vesicle or small spherical body inside which are one or
more denser spots. This may be regarded as a complete cell. All the
solid tissues consist largely of cells essentially similar to it in
nature but differing in external form. |
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In the higher organisms a cell may be defined as “a nucleated
mass of protoplasm of microscopic size.” Its two essentials, therefore,
are: a soft jelly-like material, similar to that found in the ovum, and
usually styled cytoplasm, and a small spherical body imbedded in it, and termed a nucleus.
Some of the unicellular protozoa contain no nuclei but granular
particles which, like true nuclei, stain with basic dyes. The other
constituents of the ovum, viz., its limiting membrane and the denser
spot contained in the nucleus, called the nucleolus, are not essential to the type cell, and in fact many cells exist without them. |
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Cytoplasm (protoplasm) is a material probably of
variable constitution during life, but yielding on its disintegration
bodies chiefly of proteid nature. Lecithin and cholesterin are
constantly found in it, as well as inorganic salts, chief among which
are the phosphates and chlorides of potassium, sodium, and calcium. It
is of a semifluid, viscid consistence, and probably colloidal in nature.
The living cytoplasm appears to consist of a homogeneous and
structureless ground-substance in which are embedded granules of various
types. The mitochondria are the most constant type of granule
and vary in form from granules to rods and threads. Their function is
unknown. Some of the granules are proteid in nature and probably
essential constituents; others are fat, glycogen, or pigment granules,
and are regarded as adventitious material taken in from without, and
hence are styled cell-inclusions or paraplasm. When, however,
cells have been “fixed” by reagents a fibrillar or granular appearance
can often be made out under a high power of the microscope. The fibrils
are usually arranged in a network or reticulum, to which the term spongioplasm is applied, the clear substance in the meshes being termed hyaloplasm.
The size and shape of the meshes of the spongioplasm vary in different
cells and in different parts of the same cell. The relative amounts of
spongioplasm and hyaloplasm also vary in different cells, the latter
preponderating in the young cell and the former increasing at the
expense of the hyaloplasm as the cell grows. Such appearances in fixed
cells are no indication whatsoever of the existence of similar
structures in the living, although there must have been something in the
living cell to give rise to the fixed structures. The peripheral layer
of a cell is in all cases modified, either by the formation of a
definite cell membrane as in the ovum, or more frequently in the
case of animal cells, by a transformation, probably chemical in nature,
which is only recognizable by the fact that the surface of the cell
behaves as a semipermeable membrane. | |
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FIG. 1– Diagram of a cell. (See enlarged image)
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Nucleus.—The nucleus is a minute body,
imbedded in the protoplasm, and usually of a spherical or oval form,
its size having little relation to that of the cell. It is surrounded by
a well-defined wall, the nuclear membrane; this encloses the nuclear substance (nuclear matrix),
which is composed of a homogeneous material in which is usually
embedded one or two nucleoli. In fixed cells the nucleus seems to
consist of a clear substance or karyoplasm and a network or karyomitome.
The former is probably of the same nature as the hyaloplasm of the
cell, but the latter, which forms also the wall of the nucleus, differs
from the spongioplasm of the cell substance. It consists of fibers or
filaments arranged in a reticular manner. These filaments are composed
of a homogeneous material known as linin, which stains with acid
dyes and contains embedded in its substance particles which have a
strong affinity for basic dyes. These basophil granules have been named chromatin or basichromatin
and owe their staining properties to the presence of nucleic acid.
Within the nuclear matrix are one or more highly refracting bodies,
termed nucleoli, connected with the nuclear membrane by the
nuclear filaments. They are regarded as being of two kinds. Some are
mere local condensations (“net-knots”) of the chromatin; these are
irregular in shape and are termed pseudo-nucleoli; others are distinct bodies differing from the pseudo-nucleoli both in nature and chemical composition; they may be termed true nucleoli, and are usually found in resting cells. The true nucleoli are oxyphil, i.e., they stain with acid dyes. | |
Most living cells contain, in addition to their protoplasm and
nucleus, a small particle which usually lies near the nucleus and is
termed the centrosome. In the middle of the centrosome is a minute body called the centriole, and surrounding this is a clear spherical mass known as the centrosphere.
The protoplasm surrounding the centrosphere is frequently arranged in
radiating fibrillar rows of granules, forming what is termed the attraction sphere. | |
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Reproduction of Cells.—Reproduction of cells is effected either by direct or by indirect division. In reproduction by direct division
the nucleus becomes constricted in its center, assuming an hour-glass
shape, and then divides into two. This is followed by a cleavage or
division of the whole protoplasmic mass of the cell; and thus two
daughter cells are formed, each containing a nucleus. These daughter
cells are at first smaller than the original mother cell; but they grow,
and the process may be repeated in them, so that multiplication may
take place rapidly. Indirect division or karyokinesis (karyomitosis)
has been observed in all the tissues—generative cells, epithelial
tissue, connective tissue, muscular tissue, and nerve tissue. It is
possible that cell division may always take place by the indirect
method. | |
The process of indirect cell division is characterized by a
series of complex changes in the nucleus, leading to its subdivision;
this is followed by cleavage of the cell protoplasm. Starting with the
nucleus in the quiescent or resting stage, these changes may be briefly grouped under the four following phases (Fig. 2). | |
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1. Prophase.—The nuclear network of chromatin filaments assumes the form of a twisted skein or spirem,
while the nuclear membrane and nucleolus disappear. The convoluted
skein of chromatin divides into a definite number of V-shaped segments
or chromosomes. The number of chromosomes varies in different
animals, but is constant for all the cells in an animal of any given
species; in man the number is given by Flemming and Duesberg as
twenty-four. 2
Coincidently with or preceding these changes the centriole, which
usually lies by the side of the nucleus, undergoes subdivision, and the
two resulting centrioles, each surrounded by a centrosphere, are seen to
be connected by a spindle of delicate achromatic fibers the achromatic spindle.
The centrioles move away from each other—one toward either extremity of
the nucleus—and the fibrils of the achromatic spindle are
correspondingly lengthened. A line encircling the spindle midway between
its extremities or poles is named the equator, and around this the V-shaped chromosomes arrange themselves in the form of a star, thus constituting the mother star or monaster. |
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2. Metaphase.—Each V-shaped chromosome now undergoes longitudinal cleavage into two equal parts or daughter chromosomes, the cleavage commencing at the apex of the V and extending along its divergent limbs. | |
3. Anaphase.—The daughter chromosomes, thus separated,
travel in opposite directions along the fibrils of the achromatic
spindle toward the centrioles, around which they group themselves, and
thus two star-like figures are formed, one at either pole of the
achromatic spindle. This constitutes the diaster. The daughter chromosomes now arrange themselves into a skein or spirem, and eventually form the network of chromatin which is characteristic of the resting nucleus. | |
4. Telophase.—The cell protoplasm begins to appear
constricted around the equator of the achromatic spindle, where double
rows of granules are also sometimes seen. The constriction deepens and
the original cell gradually becomes divided into two new cells, each
with its own nucleus and centrosome, which assume the ordinary positions
occupied by such structures in the resting stage. The nuclear membrane
and nucleolus are also differentiated during this phase. | |
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