THE FORMATION OF VEGETABLE MOULD THROUGH THE ACTION OF WORMS
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middle; and 23 per cent. by the basal part. If they had been drawn
indifferently by any point, the proportion for the apical, middle and
basal parts would have been 33.3 per cent. for each; but, as we have just
seen, it might have been expected that a much larger proportion would
have been drawn in by the basal than by any other part. As the case
stands, nearly three times as many were drawn in by the apex as by the
base. If we consider the broad triangles by themselves, 59 per cent.
were drawn in by the apex, 25 per cent. by the middle, and 16 per cent.
by the base. Of the narrow triangles, 65 per cent. were drawn in by the
apex, 14 per cent, by the middle, and 21 per cent. by the base; so that
here those drawn in by the apex were more than 3 times as many as those
drawn in by the base. We may therefore conclude that the manner in which
the triangles are drawn into the burrows is not a matter of chance.
In eight cases, two triangles had been drawn into the same burrow, and in
seven of these cases, one had been drawn in by the apex and the other by
the base. This again indicates that the result is not determined by
chance. Worms appear sometimes to revolve in the act of drawing in the
triangles, for five out of the whole lot had been wound into an irregular
spire round the inside of the burrow. Worms kept in a warm room drew 63
triangles into their burrows; but, as in the case of the pine-leaves,
they worked in a rather careless manner, for only 44 per cent. were drawn
in by the apex, 22 per cent. by the middle, and 33 per cent. by the base.
In five cases, two triangles were drawn into the same burrow.
It may be suggested with much apparent probability that so large a
proportion of the triangles were drawn in by the apex, not from the worms
having selected this end as the most convenient for the purpose, but from
having first tried in other ways and failed. This notion was
countenanced by the manner in which worms in confinement were seen to
drag about and drop the triangles; but then they were working carelessly.
I did not at first perceive the importance of this subject, but merely
noticed that the bases of those triangles which had been drawn in by the
apex, were generally clean and not crumpled. The subject was afterwards
attended to carefully. In the first place several triangles which had
been drawn in by the basal angles, or by the base, or a little above the
base, and which were thus much crumpled and dirtied, were left for some
hours in water and were then well shaken while immersed; but neither the
dirt nor the creases were thus removed. Only slight creases could be
obliterated, even by pulling the wet triangles several times through my
fingers. Owing to the slime from the worms’ bodies, the dirt was not
easily washed off. We may therefore conclude that if a triangle, before
being dragged in by the apex, had been dragged into a burrow by its base
with even a slight degree of force, the basal part would long retain its
creases and remain dirty. The condition of 89 triangles (65 narrow and
24 broad ones), which had been drawn in by the apex, was observed; and
the bases of only 7 of them were at all creased, being at the same time
generally dirty. Of the 82 uncreased triangles, 14 were dirty at the
base; but it does not follow from this fact that these had first been
dragged towards the burrows by their bases; for the worms sometimes
covered large portions of the triangles with slime, and these when
dragged by the apex over the ground would be dirtied; and during rainy
weather, the triangles were often dirtied over one whole side or over
both sides. If the worms had dragged the triangles to the mouths of
their burrows by their bases, as often as by their apices, and had then
perceived, without actually trying to draw them into the burrow, that the
broader end was not well adapted for this purpose—even in this case a
large proportion would probably have had their basal ends dirtied. We
may therefore infer—improbable as is the inference—that worms are able by
some means to judge which is the best end by which to draw triangles of
paper into their burrows.
The percentage results of the foregoing observations on the manner in
which worms draw various kinds of objects into the mouths of their
burrows may be abridged as follows:—
Nature of Drawn into the Drawn in, by or Drawn in, by or
Object. burrows, by or near the near the base.
near the apex. middle.
Leaves of 80 11 9
various kinds
—of the Lime, 79 17 4
basal margin of
blade broad,
apex acuminated
—of a Laburnum, 63 10 27
basal part of
blade as narrow
as, or
sometimes
little narrower
than the apical
part
—of the 34 ... 66
Rhododendron,
basal part of
blade often
narrower than
the apical part
—of Pine-trees, ... ... 100
consisting of
two needles
arising from a
common base
Petioles of a 76 ... 24
Clematis,
somewhat
pointed at the
apex, and blunt
at the base
—of the Ash, 48.5 ... 51.5
the thick basal
end often drawn
in to serve as
food
—of Robinia, 44 ... 56
extremely thin,
especially
towards the
apex, so as to
be ill-fitted
for plugging up
the burrows
Triangles of 62 15 23
paper, of the
two sizes
—of the broad 59 25 16
ones alone
—of the narrow 65 14 21
ones alone
If we consider these several cases, we can hardly escape from the
conclusion that worms show some degree of intelligence in their manner of
plugging up their burrows. Each particular object is seized in too
uniform a manner, and from causes which we can generally understand, for
the result to be attributed to mere chance. That every object has not
been drawn in by its pointed end, may be accounted for by labour having
been saved through some being inserted by their broader or thicker ends.
No doubt worms are led by instinct to plug up their burrows; and it might
have been expected that they would have been led by instinct how best to
act in each particular case, independently of intelligence. We see how
difficult it is to judge whether intelligence comes into play, for even
plants might sometimes be thought to be thus directed; for instance when
displaced leaves re-direct their upper surfaces towards the light by
extremely complicated movements and by the shortest course. With
animals, actions appearing due to intelligence may be performed through
inherited habit without any intelligence, although aboriginally thus
acquired. Or the habit may have been acquired through the preservation
and inheritance of beneficial variations of some other habit; and in this
case the new habit will have been acquired independently of intelligence
throughout the whole course of its development. There is no _à priori_
improbability in worms having acquired special instincts through either
of these two latter means. Nevertheless it is incredible that instincts
should have been developed in reference to objects, such as the leaves of
petioles of foreign plants, wholly unknown to the progenitors of the
worms which act in the described manner. Nor are their actions so
unvarying or inevitable as are most true instincts.
As worms are not guided by special instincts in each particular case,
though possessing a general instinct to plug up their burrows, and as
chance is excluded, the next most probable conclusion seems to be that
they try in many different ways to draw in objects, and at last succeed
in some one way. But it is surprising that an animal so low in the scale
as a worm should have the capacity for acting in this manner, as many
higher animals have no such capacity. For instance, ants may be seen
vainly trying to drag an object transversely to their course, which could
be easily drawn longitudinally; though after a time they generally act in
a wiser manner, M. Fabre states {89a} that a Sphex—an insect belonging to
the same highly-endowed order with ants—stocks its nest with paralysed
grass-hoppers, which are invariably dragged into the burrow by their
antennæ. When these were cut off close to the head, the Sphex seized the
palpi; but when these were likewise cut off, the attempt to drag its prey
into the burrow was given up in despair. The Sphex had not intelligence
enough to seize one of the six legs or the ovipositor of the grasshopper,
which, as M. Fabre remarks, would have served equally well. So again, if
the paralysed prey with an egg attached to it be taken out of the cell,
the Sphex after entering and finding the cell empty, nevertheless closes
it up in the usual elaborate manner. Bees will try to escape and go on
buzzing for hours on a window, one half of which has been left open.
Even a pike continued during three months to dash and bruise itself
against the glass sides of an aquarium, in the vain attempt to seize
minnows on the opposite side. {89b} A cobra-snake was seen by Mr. Layard
{90} to act much more wisely than either the pike or the Sphex; it had
swallowed a toad lying within a hole, and could not withdraw its head;
the toad was disgorged, and began to crawl away; it was again swallowed
and again disgorged; and now the snake had learnt by experience, for it
seized the toad by one of its legs and drew it out of the hole. The
instincts of even the higher animals are often followed in a senseless or
purposeless manner: the weaver-bird will perseveringly wind threads
through the bars of its cage, as if building a nest: a squirrel will pat
nuts on a wooden floor, as if he had just buried them in the ground: a
beaver will cut up logs of wood and drag them about, though there is no
water to dam up; and so in many other cases.
Mr. Romanes, who has specially studied the minds of animals, believes
that we can safely infer intelligence, only when we see an individual
profiting by its own experience. By this test the cobra showed some
intelligence; but this would have been much plainer if on a second
occasion he had drawn a toad out of a hole by its leg. The Sphex failed
signally in this respect. Now if worms try to drag objects into their
burrows first in one way and then in another, until they at last succeed,
they profit, at least in each particular instance, by experience.
But evidence has been advanced showing that worms do not habitually try
to draw objects into their burrows in many different ways. Thus
half-decayed lime-leaves from their flexibility could have been drawn in
by their middle or basal parts, and were thus drawn into the burrows in
considerable numbers; yet a large majority were drawn in by or near the
apex. The petioles of the Clematis could certainly have been drawn in
with equal ease by the base and apex; yet three times and in certain
cases five times as many were drawn in by the apex as by the base. It
might have been thought that the foot-stalks of leaves would have tempted
the worms as a convenient handle; yet they are not largely used, except
when the base of the blade is narrower than the apex. A large number of
the petioles of the ash are drawn in by the base; but this part serves
the worms as food. In the case of pine-leaves worms plainly show that
they at least do not seize the leaf by chance; but their choice does not
appear to be determined by the divergence of the two needles, and the
consequent advantage or necessity of drawing them into their burrows by
the base. With respect to the triangles of paper, those which had been
drawn in by the apex rarely had their bases creased or dirty; and this
shows that the worms had not often first tried to drag them in by this
end.
If worms are able to judge, either before drawing or after having drawn
an object close to the mouths of their burrows, how best to drag it in,
they must acquire some notion of its general shape. This they probably
acquire by touching it in many places with the anterior extremity of
their bodies, which serves as a tactile organ. It may be well to
remember how perfect the sense of touch becomes in a man when born blind
and deaf, as are worms. If worms have the power of acquiring some
notion, however rude, of the shape of an object and of their burrows, as
seems to be the case, they deserve to be called intelligent; for they
then act in nearly the same manner as would a man under similar
circumstances.
To sum up, as chance does not determine the manner in which objects are
drawn into the burrows, and as the existence of specialized instincts for
each particular case cannot be admitted, the first and most natural
supposition is that worms try all methods until they at last succeed; but
many appearances are opposed to such a supposition. One alternative
alone is left, namely, that worms, although standing low in the scale of
organization, possess some degree of intelligence. This will strike
every one as very improbable; but it may be doubted whether we know
enough about the nervous system of the lower animals to justify our
natural distrust of such a conclusion. With respect to the small size of
the cerebral ganglia, we should remember what a mass of inherited
knowledge, with some power of adapting means to an end, is crowded into
the minute brain of a worker-ant.
_Means by which worms excavate their burrows_.—This is effected in two
ways; by pushing away the earth on all sides, and by swallowing it. In
the former case, the worm inserts the stretched out and attenuated
anterior extremity of its body into any little crevice, or hole; and
then, as Perrier remarks, {93} the pharynx is pushed forwards into this
part, which consequently swells and pushes away the earth on all sides.
The anterior extremity thus serves as a wedge. It also serves, as we
have before seen, for prehension and suction, and as a tactile organ. A
worm was placed on loose mould, and it buried itself in between two and
three minutes. On another occasion four worms disappeared in 15 minutes
between the sides of the pot and the earth, which had been moderately
pressed down. On a third occasion three large worms and a small one were
placed on loose mould well mixed with fine sand and firmly pressed down,
and they all disappeared, except the tail of one, in 35 minutes. On a
fourth occasion six large worms were placed on argillaceous mud mixed
with sand firmly pressed down, and they disappeared, except the extreme
tips of the tails of two of them, in 40 minutes. In none of these cases,
did the worms swallow, as far as could be seen, any earth. They
generally entered the ground close to the sides of the pot.
A pot was next filled with very fine ferruginous sand, which was pressed
down, well watered, and thus rendered extremely compact. A large worm
left on the surface did not succeed in penetrating it for some hours, and
did not bury itself completely until 25 hrs. 40 min. had elapsed. This
was effected by the sand being swallowed, as was evident by the large
quantity ejected from the vent, long before the whole body had
disappeared. Castings of a similar nature continued to be ejected from
the burrow during the whole of the following day.
As doubts have been expressed by some writers whether worms ever swallow
earth solely for the sake of making their burrows, some additional cases
may be given. A mass of fine reddish sand, 23 inches in thickness, left
on the ground for nearly two years, had been penetrated in many places by
worms; and their castings consisted partly of the reddish sand and partly
of black earth brought up from beneath the mass. This sand had been dug
up from a considerable depth, and was of so poor a nature that weeds
could not grow on it. It is therefore highly improbable that it should
have been swallowed by the worms as food. Again in a field near my house
the castings frequently consist of almost pure chalk, which lies at only
a little depth beneath the surface; and here again it is very improbable
that the chalk should have been swallowed for the sake of the very little
organic matter which could have percolated into it from the poor
overlying pasture. Lastly, a casting thrown up through the concrete and
decayed mortar between the tiles, with which the now ruined aisle of
Beaulieu Abbey had formerly been paved, was washed, so that the coarser
matter alone was left. This consisted of grains of quartz, micaceous
slate, other rocks, and bricks or tiles, many of them from 1/20 to 1/10
inch in diameter. No one will suppose that these grains were swallowed
as food, yet they formed more than half of the casting, for they weighed
19 grains, the whole casting having weighed 33 grains. Whenever a worm
burrows to a depth of some feet in undisturbed compact ground, it must
form its passage by swallowing the earth; for it is incredible that the
ground could yield on all sides to the pressure of the pharynx when
pushed forwards within the worm’s body.
That worms swallow a larger quantity of earth for the sake of extracting
any nutritious matter which it may contain than for making their burrows,
appears to me certain. But as this old belief has been doubted by so
high an authority as Claparède, evidence in its favour must be given in
some detail. There is no _à priori_ improbability in such a belief, for
besides other annelids, especially the _Arenicola marina_, which throws
up such a profusion of castings on our tidal sands, and which it is
believed thus subsists, there are animals belonging to the most distinct
classes, which do not burrow, but habitually swallow large quantities of
sand; namely, the molluscan Onchidium and many Echinoderms. {97}
If earth were swallowed only when worms deepened their burrows or made
new ones, castings would be thrown up only occasionally; but in many
places fresh castings may be seen every morning, and the amount of earth
ejected from the same burrow on successive days is large. Yet worms do
not burrow to a great depth, except when the weather is very dry or
intensely cold. On my lawn the black vegetable mould or humus is only
about 5 inches in thickness, and overlies light-coloured or reddish
clayey soil: now when castings are thrown up in the greatest profusion,
only a small proportion are light coloured, and it is incredible that the
worms should daily make fresh burrows in every direction in the thin
superficial layer of dark-coloured mould, unless they obtained nutriment
of some kind from it. I have observed a strictly analogous case in a
field near my house where bright red clay lay close beneath the surface.
Again on one part of the Downs near Winchester the vegetable mould
overlying the chalk was found to be only from 3 to 4 inches in thickness;
and the many castings here ejected were as black as ink and did not
effervesce with acids; so that the worms must have confined themselves to
this thin superficial layer of mould, of which large quantities were
daily swallowed. In another place at no great distance the castings were
white; and why the worms should have burrowed into the chalk in some
places and not in others, I am unable to conjecture.
Two great piles of leaves had been left to decay in my grounds, and
months after their removal, the bare surface, several yards in diameter,
was so thickly covered during several months with castings that they
formed an almost continuous layer; and the large number of worms which
lived here must have subsisted during these months on nutritious matter
contained in the black earth.
The lowest layer from another pile of decayed leaves mixed with some
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