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from the surface.* Each mass of germinal matter increases
in size by the absorption of nutrient pabulum, which, as
in all other cases, passes through the layer of formed
' material. But at the same time, a portion of the germinal
matter undergoes conversion into formed material, which
accumulates upon the surface within that already formed,
and as each new layer is deposited upon the surface of the
germinal matter, those layers of formed material already pro-
duced are stretched, and with them the last developed are
more or less incorporated. (PI. VIII, fig. 28, p. 60.) For a
time, the germinal matter increases, while new-formed
material is being produced. In other words, both the
constituent parts of the entire cell increase in amount
up to a certain period of its life. (PI. IV, b.) But as
new cells continue to be produced below, those already
formed are gradually removed farther and farther from
the vascular surface, while at the same time their formed
material becomes more condensed and less permeable to
nutrient matter. From this point, each entire cell ceases
to increase in size, while the germinal matter actually
diminishes, because it undergoes conversion into formed
material ; at the same time, owing to the increased density
of the formed material, and its greater distance from the
vessels, little new pabulum is taken up to compensate for this.

* The description here given is not strictly accurate, inasmuch as
the new masses of germinal matter do not all move in a direction to-
wards the surface. Some tend in the opposite direction, towards the
subcuticular tissue, but this need not be discussed here, as it would
complicate the description without helping in any way to elucidate the
question now being considered.

2



52 PROTOPLASM.



The germinal matter (nucleus) becomes smaller as the cell
advances in age. So that it is possible to judge of the age
of a cell, irrespective of its size, by the relative amount of
its component substances. In old cells, there is much
formed material in proportion to the germinal matter, while
young cells seem to be composed almost entirely of the latter
substance. In very old cells, the small portion of germinal
matter still unconverted into formed material, dies, and the
cell having by this time arrived at the surface, is cast off, a
mass of perfectly passive, lifeless, formed material.

The facts here described are illustrated in the figure repre-
sented in PL IV, p. 48, which should be carefully studied.

Of the so-called Intercellular Substance. In cartilage
and some other tissues, there is no line of separation
between the portion of formed material which belongs
to each mass of germinal matter, as is the case in epi-
thelium, but the formed material throughout the entire
tissue forms an uninterrupted mass of tissue, matrix, or, as
it has been termed, connective substance. (PI. V, fig. 15).
From the apparent essential difference in structure, it has
been supposed that tissues of this character were developed
upon a principle very different to that upon which epithelial
structures were produced. It has been maintained by some
that in cartilage a cell wall, distinct from the intervening
transparent material, existed around each cell, and it has
been very generally concluded that the matrix was depo-
sited between the cells, and hence this was called " inter-
cellular substance." But it must not be supposed that
epithelium is in all cases to be distinguished from cartilage
by the existence of separate cells. In many forms of epi-



PLATE VI.
CARTILAGE SHOWING MODE OF PRODUCTION OF FORMED MATERIAL.

I-ig. 16. _ Flg , Yl.




ocyfiEj

Cartilage., frog ; showing germinal i
formed material, x 700.



:::::::: .;; liiLliiilir .:: ,:;;; \-\^..^-.~~ :: i^susiia
ig rartilage, kitten, showing tlie CON-






Fig. 19.



'<-: at different :\ ' 9. At\ ".attirth;

6. six weeks old ; c, nearly full grown ; d, adult

cat. x 'J15. Showing alteration in the lelative

proportions of germinal matter and formed

material at different ages



.' '-i-ninal m;



ilage inti



page 6-2



INTERCELLULAR SUBSTANCE.



53



thelium at an early period of formation, the formed material
corresponding to the several masses of germinal matter is
continuous throughout, and presents no indications of divi-
sion into separate cells. This is well seen in the lower
part of the specimen represented in PI. IV, but in fig. 13,
PI. V, an unusually striking example is given. The spe-
cimen was taken from the deeper portion of the con-
junctival epithelium of man. Not only is there no indi-
cation of division into distinct cells, but the structure
would be described as a matrix exhibiting spaces occu-
pied by the masses of germinal matter. The arrange-
ment exactly corresponds with that existing in the case of
cartilage, and the masses of germinal matter with a thin
investment of formed material may be removed just as in
that tissue. It is, therefore, clearly erroneous to consider
cartilage and epithelium as representatives of different classes
of tissues. The analogy between them will be at once under-
stood by a glance at fig. 13, and fig. 15, which have been
carefully copied from actual specimens. In fig. 14, a portion
of older epithelium from the same surface is represented.
In this, each mass of germinal matter is invested with its
own layers of formed material, and these are distinct from
neighbouring portions. A " cell," or elementary part of
fully-formed cartilage and tendon, consists of a mass of
germinal matter, with a proportion of formed material
around it. A line passing midway between the several
masses of germinal matter would mark roughly the limit of
the formed material, corresponding to each particular mass
of germinal matter, and this would correspond with the
outer part of the surface or boundary of the epithelial cell.



54 PROTOPLASM.



In order to understand the true relation of the so-called
intercellular substance of cartilage or tendon to the masses
of germinal matter, it is necessary to study the tissue at
different ages. At an early period of development, these
tissues appear to consist of masses of germinal matter only.
As development advances, the formed material increases,
and the masses of germinal matter become separated farther
and farther from one another. (PI. VI, fig. 16.) The appear-
ances of a cell wall around the germinal matter in the
fully-formed tissue, and other alterations which occur, and
anomalous appearances which often result as age advances,
can be even more readily understood upon the view here
advanced, than upon the intercellular-substance theory
which has been so strongly supported by some observers.
See PI. VI, figs. T 6 to 22.

Of the Formation of the Contractile Tissue of Muscle.
A muscle "cell," or elementary part, will consist, like that
of cartilage and tendon, of the so-called nucleus, with
a portion of the muscular tissue corresponding to it. In
general arrangement it closely resembles what is seen in
tendon. The contractile material of muscle may be shown
to be continuous with the germinal matter, and oftentimes
a thin filament of the transversely striated tissue may be
detached with the oval mass of germinal matter still con-
nected with it, showing that, as in tendon, the germinal
matter passes uninterruptedly into the formed material.
This contractile tissue is not, like the germinal matter
which produced it, in a living state. In the formation
of the contractile tissue, the germinal matter seems to
move onwards, and at its posterior part gradually under-



WHAT IS A CELL?



55



goes conversion into the tissue. At the same time it
absorbs nutrient material, and thus, although a vast amount
of contractile tissue may have been produced, the germinal
matter which formed it may not have altered in bulk.
(PI. VII, fig. 25.) The fibres of yellow elastic tissue are
formed in the same manner, and each fibre is thickened by
the formation of new material from germinal matter, which
lies upon the external surface of each fibre (fig. 26.).

The Formation of Nerve Fibres. The nerve fibre is
composed of formed material, which is structurally con-
tinuous with the formed material of the nerve cells of the
nerve centres. A nerve fibre at an early period of develop-
ment consists of a number of oval masses of germinal
matter linearly arranged. As development proceeds, these
become separated farther and farther from one another, and
the non-living tissue which is thus spun off as they become
separated, is the nerve. (PI. VII, fig. 27.)

What is essential to the Cell? All that is essential
to the cell or elementary part is matter that is in the
living state germinal matter, and matter that has been in
the living state formed material. With these is usually
associated a certain proportion of matter about to become
living the pabulum or food. So that we may say that
in every living thing we have matter in three different
states matter about to become living, matter actually
living, and matter that has lived. The last, like the first,
is non-living, but unlike this it has been in the living state,
and has had impressed upon it certain characters which
it could not have acquired in any other way. By these
characters we know that it has lived, for we can no more



5 6 PROTOPLASM.



cause matter artificially to exhibit the characters of the dried
leaf, the lifeless wood, shell, bone, hair, or other tissue, than
we can make living matter itself in our laboratories.

Cells are not like B 'ricks in a Wall. Cells forming a
tissue have been compared with bricks in a wall, but the
cells are not like bricks, they have not the same con-
stitution in every part, nor are they made first and then
embedded in the mortar. Each brick of the natural
wall grows of itself, places itself in position, forms and
embeds itself in the mortar of its own making. The whole
wall grows in every part, and while growing may throw
out bastions which grow and adapt themselves perfectly to
the altering structure. Even now it is argued by some
that because things, like fully formed cells, may be made
artificially, the actual cells are formed in the same sort
of way an argument as forcible as would be that of a
person, who after a visit to Madame Tussaud's Exhibition,
seriously maintained that our textures were constructed
upon the same plan as the " life-like " wax figures he had
seen there.

Every one who really studies the elementary parts of
tissues and investigates the changes which occur as the
germinal matter passes through various stages of change
until the fully developed structure results, will be careful
not to accept without due consideration the vague generali-
sations of those who persist in authoritatively declaring that
the changes occurring in cell growth are merely mechanical
and chemical, although they are unable to produce by any
means at their disposal a particle of fibrine, a piece of carti-
lage, or even a fragment of coral. They avoid the difficulty



PL A'!

NUTRITION AND MOVEMENTS OF G5.RMJNAL MATTER.
6



supposed, pabulum, fl
i !>v the arrow*



iterial and passing into the gr-
eat formed rna:
c*, oldest portion ot



a. terminal matter:
ping in the direction
iversing the formed.




linaL matte
it produced ;



A minute particle of germinal matter
its component spherules of living ma
thin layer of so.'t formed, material on
undergoing change.



tier and



Fig. 25.




nal matter and formed material (contractile tissue) of muscle. The iermhial m

.: 111 the direction Of the arrow. Ir, ' = "~ v^t^,-, .. arwl /, V,ur. ITMH rnov f ><l from tht



11 the. direction of the arrow. Ir, is now between a. and b, 1 ed frcm U.

position between 6 and c and contractile tissue there seen has been fon



Fig. 26.




ellow elastic tissue from the lamb. The germinal matter is moviin; in thci
, and forming the yellow elastic tissue as it proceeds It ha* 1 en said that.
-,. but a grat ruiruber u; DO
specimena.













NUTRITION OF A LIVING CELL. 57

as regards the germinal matter by ignoring its existence, and
attribute to a "molecular machinery" which the mind cannot
conceive, and which cannot be rendered evident to the
senses, all those wonderful phenomena which are really due
to vital power. Moreover, resemblances to living organisms
of the most fanciful kind are adduced apparently for the
purpose of leading people to believe that non-living matter
behaves like that which is alive.*

On the Nutrition of a Living Cell. In nutrition, the
active changes are exclusively confined to the germinal
matter. The formed material is passive, and probably acts
like a filter, permitting some things to pass and interfering
with the passage of others. In nutrition, pabulum becomes
germinal matter to compensate for the germinal matter
which has been converted into formed material. Now let
us consider the order of these changes, and endeavour to
express them in the simplest possible manner.

Let the germinal matter which came from pre-existing
germinal matter be called a; the non-living pabulum, some

* Professor Tyndall describes (" Proceedings of the Royal Society,"
vol. xvii, No. 105) the changes resulting from the influence of light on
the vapour of an aqueous solution of hydriodic acid. His rhapsodical
description, which extends over an entire page, contains the following
curious allusions and comparisons : A cloud was developed like an
organism from a formless mass to a marvellously complex structure ;
spectral cones with filmy drapery; exquisite vases with the faintest
clouds, like spectral sheets of liquid, falling over their edges ; clouds
like roses, tulips, sunflowers, and bottles one within the other ; a cloud
like a fish, with eyes, gills, and feelers, and like a jelly fish, with the
internal economy of a highly complex organism, exhibiting the twoness
of the animal form ; as perfect as if it had been turned in a lathe ; and
likely to prove exceedingly valuable to pattern designers !



5 8 PROTOPLASM.



of the elements of which are about to be converted into
germinal matter, shall be b; and the non-living formed
material resulting from changes in the germinal matter, c.

It is to be remarked that b does not contain c in solu-
tion, neither can c be made out of b unless b first passes
through the condition a, and a cannot be formed artificially,
but must come from pre-existing a.

In all cases b is transformed by a into a, and a under-
goes conversion into c. Can anything be more unlike
chemical and physical change ? Neither a, nor #, nor c can
be made by the chemist ; nor if you give him b can he
make a or c out of it ; nor can he tell you anything about
the "molecular condition" or chemical constitution of a,
for the instant he commences his analysis a has ceased to
be a, and he is merely dealing with products resulting from
the death of a, not with the actual living a itself. The
course which the pabulum takes in the nutrition of the
germinal matter of a cell is represented by the arrows in
fig. 23, pi. VII.

The nature of the process of nutrition is more fully
discussed towards the end of the next section, " OF LIFE."

Of the Increase of Cells. Several distinct modes of cell
increase or multiplication have been described, but in all
cases the process depends upon the germinal matter only.
It is this which divides ; and it is the only part of the cell
which is actively concerned in the process of multiplication.
It may divide into two or more equal portions, or give off
many buds or offsets, each of which grows as a separate
body as soon as it is detached. (PI. VIII.)

The formed material of the cell is perfectly passive in



CHANGES IN THE CELL IN DISEASE. 59

the process of increase and multiplication. Even the
apparently very active contractile tissue of muscle has no
capacity for increase or formation. If soft or diffluent, a
portion of the formed material may collect around each of
the masses into which the germinal matter has divided, but
it does not grow in or move in and form a partition, as has
often been stated. When a septum or partition exists, it
results not from "growing in," but it is simply produced by a
portion of the germinal matter undergoing conversion into
formed material of which the partition is composed. (PI. V,
fig. 15 a and b.)

Of the Changes in the Cell in Disease. I have en-
deavoured to show that of the different constituents of
the fully formed cell, the germinal matter is alone con-
cerned in all active change. This is in fact the only
portion of the cell which lives, while at an early period
of development, some of the structures usually regarded as
essential to cell existence are altogether absent, and the cell
is but a mass of germinal matter. But it must be borne in
mind that at all periods of life, in certain parts of the
textures and organs, and in the nutrient fluids, are masses
of germinal matter, destitute of any cell-wall, and exactly
resembling those of which at an early period the embryo
is entirely composed. White blood and lymph corpuscles,
chyle corpuscles, many of the corpuscles in the spleen,
thymus and thyroid, corpuscles in the solitary glands, in
the villi, some of those upon the surface of mucous mem
branes, some in connection with muscle, nerve, bone, carti-
lage, and some other tissues, are of this nature, and consist
of living germinal matter, with mere traces of soft formed



60 PROTOPLASM.



material around each mass. There is no structure through
which these soft living particles, or small portions of living
matter detached from them, may not make their way. The
destruction of tissue may be very quickly effected by the
growth and multiplication of such masses of germinal
matter. Many of the changes in disease result from the
undue growth of this substance, and indeed there is no
operation peculiar to living beings in which germinal or
living matter does not take part. Any sketch of the struc-
ture of the cell would be incomplete without an account of
some of the essential alterations which occur in it in disease.
I propose, therefore, to refer very briefly to the general
nature of some of the most important morbid changes.

Within certain limits, the conditions under which cells
ordinarily live may be modified without any departure from
the healthy state, but if the conditions be very considerably
changed, disease may result, or the cell may die. For
instance, if cells, which in their normal state grow slowly,
be supplied with an excess of nutrient pabulum, and increase
in number very quickly, a morbid state is engendered. Or
if, on the other hand, the rate at which multiplication takes
place be reduced in consequence of an insufficient supply
of nourishment, or from other causes, a diseased state may
result. So that, in the great majority of cases, disease or
the morbid state essentially differs from health or the healthy
state in an increased or reduced rate of growth and multi-
plication of the germinal matter of one or more particular
tissues or organs. In the process of inflammation, in the
formation of inflammatory products, as lymph and pus, in the
production of tubercle and cancer, we see the results of in-



PLATE VIII.

FROM THE NUTRITION OF HEALTH, TO THE INFLAMMATION O?
DISEASE.



*



The production of formed material from germinal matter in epithelial cells. See also Plate IV.

Fisf. 33.
?ig. 30. Fie. 31. Fig. 32.




Rupturp of forr

[jcrmiLtiuA fre-.

pabuli



of.ger-
ol adult c-'-ii i
of pabulum. In this
foi-med

from the germinal matter of epi-
thelium.




Multiplication of pus corpuscles.



Fig 36.














CELL IN DISEASE. 6 1

creased multiplication of the germinal matter of the tissues
or of the germinal matter derived from the blood, con-
sequent upon the appropriation of excess of nutrient pabulum.
In the shrinking, and hardening, and wasting which occur
in many tissues and organs in disease, we see the effects of
the germinal matter of a texture being supplied with too
little nutrient pabulum, in consequence sometimes of an
alteration in the pabulum itself, sometimes of an undue
thickening and condensation of the tissue which forms the
permeable septum, which intervenes between the pabulum
and the germinal matter.

The above observations may be illustrated by reference
to what takes place when pus is formed from an epithelial
cell, in which the nutrition of the germinal matter, and
consequently its rate of growth, is much increased. And
the changes which occur in the liver cell in cases of wasting
and contraction of that organ (cirrhosis] may be advanced
as an illustration of a disease which consists essentially in
the occurrence of changes at a slower rate than would be
the case in the normal condition, consequent upon the
normal access of pabulum to the germinal matter being
interfered with.

The outer hardened formed material of an epithelial cell
may be torn or ruptured mechanically, as in a scratch or
prick by insects (PI. VIII, figs. 32 to 35); or it may be
rendered soft and more permeable to nutrient pabulum by
the action of certain fluids which bathe it. In either case
it is clear that the access of pabulum to the germinal matter
must be facilitated, and the latter necessarily "grows" that
is, converts certain of the constituents of the pabulum that



62 PROTOPLASM.



come into contact with it into matter like itself at an
increased rate. The mass of germinal matter increases in
size, and soon begins to divide into smaller portions, fig. 33.
Parts seem to move away from the general mass, fig. 34.
These at length become detached, and thus several separate
masses of germinal matter, which are embedded in the
softened and altered formed material, result, figs. 34, 35.
These changes will be understood by reference to the figures
in Plate VIII. In this way the so-called inflammatory
product pus results. The abnormal pus-corpuscle is pro-
duced from the germinal or living matter of a normal
epithelial or other cell, or elementary part, the germinal
matter of which has been supplied with pahilum much
more freely than in the normal state. In all forms of in-
flammation, the germinal matter of the parts inflamed
increases very much, and the same change occurs in every
kind of fever, fig. 36, pi. VIII, but not proceed to the same
extent. In both conditions there is increased development
of heat due to the increase of the germinal matter. In-
flammations and fevers are so very closely related that an
inflammation may be spoken of as a local fever, and a
fever as a general inflammation.

It will be seen how easily the nature of the changes
occurring in cells in inflammation, fever, and other morbid
changes, can be explained, if the artificial terms, cell-wall,
cell-contents, nucleus, be given up. In all acute internal
inflammations and in fevers a much larger quantity of in-
animate pabulum is taken up by certain cells and con-
verted into germinal matter than in the normal state.
Hence there is, at least in the parts affected, increase in



GERMINAL MATTER IN DISEASE. 63

bulk. Cells of particular organs, which live very slowly in
health, live very fast in certain forms of disease. More
pabulum reaches them, and they grow more rapidly in
consequence.

It is by this process of increased multiplication and
reproduction of certain kinds of germinal matter of the
organism, under altered conditions, that the germs which
constitute the material particles of contagious diseases
result. These living particles (contagium) having acquired
during multiplication new and peculiar properties not pos-
sessed by the germinal matter from which they originally
sprung, retain these properties and reproduce their kind a
million fold whenever placed under conditions favourable
to the process, though the operation may be fatal to the
organism in which it occurs.

In cells which have been growing very rapidly and are
returning to their normal condition, in which the access of
nutrient pabulum is more restricted than in the abnormal state,
as is also the case in normal cells passing from the em-
bryonic to the fully-formed state, the outer part of the
germinal matter undergoes conversion into formed material,
and this last increases although the supply of pabulum is
reduced.

From these observations it follows that disease may re-
sult in two ways either from the cells of an organ growing
and multiplying faster than in the normal state, or from
their doing so more slowly. In the one case, the normal
restrictions under which growth takes places are diminished ;


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