NEMATODES AND THEIR RELATIONSHIPS.

^
ByN. A. CoBB,
Technologist in Charge of Agricultural Technology, Bureau of Plant Industry,

INTRODUCTION.

THE soils of our yards, gardens, and fields swarm with thous-
ands of kinds of minute animals and plants of which we
know little or nothing. We depend on the soil for our very
existence, and it may seem that this fact should have caused
us long ago to make ourselves thoroughly acquainted with
it and all its inhabitants; yet the truth is otherwise. Here
beneath our very feet are microbes, protozoa, fungi, and many
other kinds of small organisms, thousands of species, of which
we know hardly the first thing beyond the mere fact of their
existence. In some ways this ignorance extends even to the
higher plants and animals. Confront the ordinary botanist
with the complete root system of one of our common plants
and ask him what it is and the chances are he wiU have to
confess his ignorance. While his knowledge of the above-
ground parts of the higher plants is most systematic and ex-
tensive, in most cases the corresponding parts below the
surface of the ground are almost whoUy unknown to him.
Relatively speaking, then, in a biological sense, this sou we
daily tread under foot is almost a veritable terra incognita.
Why is this so? It is difficult to formulate a reasonable
answer. The fact is that our biological researches as a rule
do not extend below the surface of the ground. This lamen-
table fact is, of course, an answer to the question, but it does
not seem a reasonable one.
Inhabiting the soil in myriads, hidden behind this veil of
ignorance, there is a group of organisms known as nematodes.
Some members of this group have forced themselves on our
attention by the enormous damage they do to crops. These,
however, so far as numbers are concerned, constitute a very
1 The illustrations are mostly from drawings made from nature under the author's direc-
tion by Mr, W, E. Chambers. Figures 1 and 19 are from the author's drawings, and figure
10 was prepared by Miss Ella Welborn from the basis of SchepotiefE's plates.
457
 458 Yearbook of the Department of Agriculture,
insignificant minority. There are multitudes of others,
some only in a lesser degree injurious, still others that are
beneficial, and yet others, the great majority, of unknown
Lip Cephalic seta
Lateral field ..., Labial papilla
Pharyngeal bulb , Pharyngeal rib
Pharynx
, Pharynx or throat
Salivary gland
, Pharyngeal tooth
Muscular layer
Pharyngeal bulb
Cuticle Salivary gland
Hair Lateral organ
Œsophagus Œsophageal tube
Cuticle ... Eye, with lens
Nerve-ripg
Excretory pora
Duct of ventral {jland.. Ampulla
Œsophagus.... .... Median bulb
Nerve ring , Nerye-ring
Ridge in cuticle Ganglion cells
Œsophagus Duct of ventral gland
Stria of cuticle Duct of lateral glana
Sub-cuticula .... A lateral gland
Wing of cutícula
Cardiac bulb
Cuticle Valvular apparatus
Cardiac bulb
Wing or ridge in skin
Dorsal field
Cardiac coUum
Valvular apparatus Cardia
Its radial muscles Stomach or intestine
Muscular layer Tessellation of same
Wall of intestine Cuticle
Wing of cuticle, Wall of intestine
Ovum fit to fertilize Ventral gland
Striation in cuticle
Ripe ovtmi
Uterus
Unripe ovum in ovary
29. Blind end ovary . Spermatozoa
SO. Intestine Fertilised egg in uterus
81. Yolk of egg Blind end of ovary-
82. Nucleus of egg Yolk of egg
S3. Shell of egg... Vaginal gland
84. Internal lateral field .. Vulva
85. External lateral field .. Vagina
86. Intestine Vaginal gland
Lateral field
Seminal vesicle
37. Copulatory muscle Wall of same
88. Wing of cuticle Spermatozoa
89. Copulatory muscle Copulatory muscle
40. Intestinal cell Accessory male organ
41. Accessory male organ. - Ductus ejaculatorius
42. Ejaculatory duct Three pre-anal papillœ
43. Copulatory miiscle Copulatory muscle
44. Wing of cuticle Proximal end spiculum
45. Constriction in spicului Pylorus
46. £ight-hand spiculum.. Anterior ribs of bursa
47. Distal end spiculum .. Anal gland
48. Accessory piece Anus
49. Caudal gland Post-anal papillae
60 Left wing of bursa
Caudal gland
51. 1st caudal gland Median ribs of bursa
C2. 2nd caudal gland Bursa
53. 3rd caudal gland Posterior ribs of bursa
64. Muscular wall Duct of caudal glands
55. Eight wing of bursa Terminiu

FIG.26.—Diagrammatic representation of the nematode anatomy. The upper part of the fig.
ure and four corresponding circular cross sections at the left represent the anterior half of a
female nematode seen from the right-hand side. The lower part of the figure and the two
corresponding cross sections represent the same view of the posterior part of a male nema"
tode. The cross sections are placed opposite the portions of the side view to which they
relate.

economic significance. So little do we know of this vast

multitude of soil-inhabiting nematodes that the first spade-
ful of earth we lift is practically certain to contain kinds
 Nematodes and Their RelationsMps. 459

never seen before (fig. 27). Numerous as are the nematodes

inhabiting the soil, the aquatic species are even more nu-
merous. Furthermore, thousands of species of nematodes
occur as parasites in animals and plants.
At present our knowledge of this group, such as it is, is
confined to a comparatively small
number of people. The reasons for
this are not far to seek. If we con-
sider any group of animals the study
of which is popular—as, for instance,
birds—we find the individuals of that
group both large and numerous, and,
moreover, much in evidence; and it
will be found, in most cases, that they
are serviceable to man; they furnish
him food, raiment, or some other valu-
able material; or they are ornamental.
FIG. 27.—Diagram representing the upper foot of soil, and
showing the relative abundance of nematodes in each
successive 2 inches. Derived from a low-lying alluvial
soi) carrying about 3,000,000,000 nematodes to the acre.
Most of the nematodes are in the upper 3 inches. Though
shown distributed uniformly in each layer, they are really
most numerous about the roots of plants.

or attractive as pets, or useful in some

other way. Now, it is not unfortunate
for the nematodes that they are none
of these, but it is unfortunate for us,
if on that account we are prevented
from knowing much that would be
most useful to us, as well as interest-
ing and diverting.
Nematodes do not furnish hides,
horns, tallow, or wool. They are not
fit for food, nor do they produce
anything fit to eat; neither do they
sing or amuse us in any way; nor
are they ornamental—in fact, when they are displayed in
museums the public votes them hideous. Lacking in all
these respects, they fail even in furnishing any moral or
praiseworthy example; they are not known to be indus-
trious like the ant, or provident like the bee, indomitable
460 Yearbook of the Department of Agriculture.
like the spider, or frugal, or honest, or anything else that is
admirable. What claim, then, one may ask, can such
beings lay to our attention? I think I hear some outraged
naturalist exclaiming in reply: ''And must a thing be use-
ful to deserve attention? Must a thing have an assessed
money value to be worthy of study ? Is there no idea above
meat for dinner and raiment for the reception? Is every-
thing to be measured by its value in dollars and cents?
Thank God, there are some things so far removed from the
lucre system we allow to dominate our lives that it is as
impossible to measure the one in terms of the other as it
would be to buy a ticket into Heaven/^ I should have a
good deal of sympathy with the man whose wrath boiled
over in this manner and found expression in such burning
language, but, at the same time, I confess I should have
more hope of converting others to my way of thinking by
adopting another tone and different tactics. Descending
to one of the lowest planes on which an appeal could be
made, I should say that, if, for example, it could be proved
that nematodes would draw as a show to such an extent as
to cause the populace to part with half a dollar each for a
place in the balcony, a dollar or two for the dress circle, and
all the rest of it, we might feel their claims for attention to
be meeting with some little encouragement. Such a demon-
stration would be more forcible than any tirade against
even the most reprehensible prejudice.
But it is unnecessary to descend to any such appeals to
show the advisability of more fuUy acquainting ourselves
with these organisms, for although, so far as we know at
present, nematodes may not often be directly beneficial to
man, a knowledge of them would be useful to every person,
simply because, though he may not know it, he is infested
by them, either continually or from time to time. This
xmpalatable fact needs no proof to the initiated; but those
not informed will doubtless be surprised to learn that no
less than about 50 kinds of nematodes are known to infest
the human body.
Oiu* domesticated animals suffer in like manner, and
even more severely (fig. 28). Thus, in countries where the
wealth consists largely in live stock, as, for instance, in the
western part of this country, in Australia, and in the Argen-
 Nematodes and Their Relationships, 461

tine Kepublic, the monetary loss caused by nematodes is

always considerable, and sometimes very great. The para-
sitic nematodes shown on this page exist in millions in most
of the great sheep-growing countries; these are only a frac-
tion of the species of nematodes that attack sheep, and are
mentioned simply as embodying a typical case among our
domesticated animals. To every person, therefore, and
especially to medical men and veterinarians, these para-
sitic nematodes should possess no common interest.

ise
FIG.28.—Natural-size figures of some of the nematode parasites of the sheep: a, b, Found
in the duodenum; c, in the intestine; d, in the stomach, very common and injurious;
e, in the intestine, very common and injurious; /, in the anterior part of the small
intestine, Inflicting severe bites; g, in the large intestine. These are only a fraction of the
nematodes known to infest the sheep.

Nor are the crops of the farmer free from the attacks
of these creatures. Wheat, the sugar beet, coffee, and
scores of other crops are frequently decimated or even
ruined by diseases caused by the attacks of microscopic
nematodes.
Beyond doubt a complete knowledge of nematodes, if
properly applied, would enable us to save a vast amount of
life and treasure and prevent a vast amount of suffering.
But it is by no means entirely for these reasons that I
would like to see these organisms receive a greater amount
462 Yearbook of the Department of Agriculture,
of attention. They possess a powerful interest because of
the unusually clear view they give us of the various processes
relating to life, and I beUeve this fact could easily be turned
to educational account; in
fact, I know it could. In the
contemplation and discussion
of one of these organisms, I
have seen the statesman forget
his social problems, the judge
his law, the mathematician his
quantities, the artist his art,
the philanthropist his schemes,
and exclaim and question in

FIG. 29.—A very remarkable nematode, Wil-

sonema, found in soils in various parts of the
United States. How it can make its way
through the soil without damage to its won-
derfully complicated and delicate mouth
parts is a mystery. /. Lateral view of female;
II, dorso-ventral view of the head of the same
individual; ///, enlarged lateral view of the
head of same individual; IV, lateral view of
the tail end. a, Ventral appendage, which,
together with the corresponding dorsal ap-
pendage, acts as a sieve; b, lateral tactue organ
associated with the sifting apparatus; c, in-
ternal elements (supports?) of the sieves; d,
inflated valvular apparatus; e, amphid; /,
cuticle; g, pl^rynx; h, nerve ring; i, excre-
tory pore; j, esophagus; k, threefold valve of
the cardiac bulb; I, cardia; m, wall of intes-
tine; n, flexure of anterior ovary; o, vulva;
P, 6gg; q, lumen of intestine; r, rectum; s,
lateral wings; t, anus; u, caudal seta; v, spin-
neret; w, anal muscles; z, apical portion of
spinneret; y, one of the three caudal glands.

terms of enthusiasm and inter-

est concerning which there
could be no doubt.
Formerly it was my duty, as
professor of biology, to bring
before the students in my
courses, first in a secondary
school and later in the uni-
versity, a series of animals and plants for dissection and
study, and I can safely say that no organisms excited so
much interest among those young men, or furnished them
 Nematodes and TJieir RelationsTiips. 463

with a greater amount of instruction, than small species of

free-Hving nematodes. These little animals present a won-
derful complexity of organ-
ization combined with such
transparency that very little
Í3 hidden from view. Diges-
tive system, nervous system,
excretory system (fig. 29),
muscular system, sexual sys-
tem, aU are to be seen with
most instructive wealth of
detail and in fuU action in
the living animal. My ex-
perience in this matter makes
me very confident in saying
that professors of biology
could do far worse than to
introduce into their courses
a more careful examination
of these creatures.

NEMATODES AND THE NEW

SCIENCE OF HEREDITY.
If it were necessary to for-
tify these assertions, I might
call attention to the fact that
since the days of Leeuwen-
hoek and the invention of
the microscope these organ-
isms have excited a keen
interest in the minds of great
naturaHsts. Anyone will be
convinced of this by even a
casual glance at the litera-
ture relating to any one of FIG. 30,—The asymmetrical nematode, Buno-
nema. This apparently deformed little crea-
several common species—for ture presents relatively monstrous warts on
instance, the nematode so one side of its body. It belongs to a large
group, usually to be found in decaying matter.
common in table vinegar
and popularly known as the ^Vinegar eel.'^ I might also
caU attention to the fact that some of the most momentous
of all our scientific discoveries have been made through the
464 Yearbook of the Department of Agriculture.

instrumentality of nematodes; witness tlie discovery of the

formation of the polar bodies in the ovum, the coalescence
of the spermatic nucleus
with that of the ripe
ovum, and the subse-
quent cell-division phe-
nomena, leading up to a
science of heredity. The
clarity of the nematode
egg makes it particularly
suitable for investiga-
tions of this kind.
If one removes from
the small part of one of
the ovaries of the com-
mon lung worm of the
frog, Ehahdias nigroven-
osaj an egg that has not
yet begun to segment,
and places it in water
under a microscope mag-
nifying 300 to 400 diame-
ters, he may easily ob-
serve one of the most
impressive and instruc-
tive of all the phenom-
ena of life—the forma-
tion of a young animal
inside the egg. This egg,
which is at first a single
cell, becomes in the
course of a few hours
under our very eyes a^
living, actively moving
FIG. 31.—A remarkable nematode, called Heth, nota- animal This imnressive
ble on account of the numerous sieves through ' ^ . ^
which it sifts its food, and also on account of the spectaclc DCgmS by the
acute backward-pointing barbs, of relatively large formation from the Oriff-
size, with which its head is armed. «in/« i <•
mal cell oi two others of
slightly unequal size, the change being accompanied by the
karyokinetic phenomena concerning which there is now-
adays so much discussion and research. Each of these cells
 Nematodes and Their RelationsTiips. 465
again divides^ and so the rhythmic change goes on, giving rise
to tissue after tissue and organ after organ, until the baby
nematode lies before us complete and ready to begin its
active battle for existence. The observer rises at last, feel-
ing that he has indeed approached as near to the mystery
of life as it is yet given to man to approach.
We are gradually rebuilding the science of heredity, and
one of the basic facts of this renovation is the union of the
nuclear material of two parent cells to form the nucleus from
which the offspring grows. This flow or transportation of
nuclear material from parent to offspring is a matter of su-
preme interest and is now being studied with the minutest
care, and is found to foUow laws capable of such exact defini-
tion that we are able to base upon them predictions as to the
nature of the offspring. Practical results of vast importance
in the breeding of animals and plants are, therefore, now only
a matter of time.
The fundamental facts in this very important new science
came to light through observations upon the eggs of nema-
todes. It was in a nematode egg that it was first observed
that the animal male and female nuclei actually coalesce to
produce the new compound nucleus, or pronucleus, which
alone is capable under the usual normal conditions of growing
and producing a new individual.
Again, it was in the nematode egg that it was first proved
that the pronucleus when it divides sends into each of the
two new ceUs a definite portion of the matter it derived from
each parent. Both of these discoveries are absolutely fun-
damental and of enormous practical significance.
It is no wonder, therefore, that there should exist on the
part of those most familiar with these fields of research, a wish
that our knowledge of nematodes should be increased and
brought to the attention of a larger number of people.
An old adage assures us there is no royal road to knowledge.
There is certainly no royal road to a knowledge of nematodes.
The traffic in this direction has not justified the installation
of through trains and sleeping cars; so he who takes this
route must be prepared to put up with inconveniences, and
to make the best of certain disgusting passages. To the
squeamish, the lazy, the impatient, the inaccurate, about to
take'tickets for this journey, I should say, ^'Don't do it! It
75922°—YBK 1914 30
466 Yearbook of the Department of Agriculture,
won't suit you." But to him not afraid of work, brave
under discouragements, patient, cautious, and with a good

FIG. 32.—The dragon nematode,

Draconema, so called because its
form is reminiscent of that of the
dragons of fable.
ß
amount of ability and common sense, I
would wish a hearty Godspeed, and say,
'^Go ahead! it's a rough road and hard to
travel, but there are some fine bits of
scenery, unequaled in their way, which
amply make up for the inconveniences."
HISTORICAL.
NEMATODES AS A CAUSE OF DISEASE.

The history of nematodes extends back

into remote antiquity and contains many
remarkable passages. It seems hardly
credible, and yet it is true, that at one
time there was a widely prevalent ^^ worm
theory" of disease, which in its day played
a part comparable to that played in more
recent times by the germ theory of dis-
ease. This worm theory had for its basis
the fact that worms are obviously asso-
ciated with numerous and varied forms
of disease. Here was something tangible
to which the medical men of those remote
times could trace certain baneful results.
This was before the invention of
the microscope, and it was not
unnatural that, in the absence of
microscopes to reveal the enormous prevalence of micro-
organisms, there should spring up and become widely accepted
 Nematodes and Their Relationships. 467
a theory that worms were a general cause of disease, and
that even those diseases not obviously due to this cause were
in reality due to it through some means not yet understood.
Of all the organisms that inhabit the human body none are
more prominent or more likely to come under the observation
of the physician than certain of our nematode parasites—
indeed a knowledge of these neinatodes and their injurious
effects may well antedate history.
The squeamish reader will not enjoy a paragraph which
tells of the details and abundance of these parasites in human
beings and the lower animals. I have removed from the
stomach of a wallaby weighing not over 50 pounds no less
than 40,000 nematodes varying in length from a few milli-
meters to several inches. As for human beings, there are
regions where the inhabitants are pot-bellied owing to nema-
tode infestation and where it is not an unheard-of thing for
an infested individual to vomit a pint or more of ^'stomach
worms''—parasitic nematodes infesting the human stomach.
Such conditions are most prevalent among savages and igno-
rant people, and it is readily behevable that in ancient times
the conditions were more favorable to these parasites than
they are to-day. Hence, it is easy to understand how a
theory of disease based on the prevalence of nematodes
could have arisen and gained a wide currency.
In modern times we have come to know much more about
such diseases and that parasitic species of nematodes cause
many diseases of mankind not formerly recognized. The
dreaded ^^hookworm" is a nematode. So is that scourge of
the Tropics, the ^^guinea worm.'' Trichina,^ costing civilized
nations hundreds of thousands of dollars yearly for the
inspection of pork, is a nematode. If raw or insufficiently
cooked trichina-infested pork be eaten by human beings, the
result is a serious, oftentimes fatal, sickness called trichinosis,
epidemics of which have claimed victims by the hundred.
Nematodes have recently been suspected, with good show of
of reason, of being carriers of cancer. So the list of more or
less serious human nematode diseases and ailments might
be increased until practically half a hundred had been
enumerated.
^ Trichinella spiralis.
468 Yearbook of the Department of Agriculture,
No less serious are the nematode diseases of plants and of
the lower animals. The common gallworm has been found
infesting the roots of several hundred different species of
plants, among them most of our cultivated crops, and causes
an annual loss amounting to millions of dollars. There is
another nematode that has at times completely checked the
growing of sugar beets in certain regions. The list of serious
diseases of this character could
easily be increased to dozens. The
same is true of animals. Every do-

FiG, 33.—One of the injurious parasitic nematodes

found in and on the roots of citrus trees, Tylen-
chulus semipenetrans. This is a male. It is the
females of this species that are most injurious to
the citrus roots. This parasite has been spread
throughout the world on the roots of citrus nursery
stock. Its original habitat is probably southeast-
ern Asia, whence came our cultivated species of
citrus. The females are much larger and penetrate
for half their length into the citrus rootlets, leaving
the posterior saccate part, devoted to the production
of eggs, outside the roots. A new generation is pro-
duced every few weeks. These nematodes are de-
voured by another nematode shown in figure 20, a
beneficial species, which hunts and devours this
citrus parasite. This is a lateral view of a full-
grown male. The spear is usually very inconspic-
uous—always deteriorated. Note also the deterio-
rated median bulb, sometimes apparently absent.
In the male, in contrast with the female, the anus
develops, a, Lip region; b, spear; c, 3-bulbed base
of spear; d, esophageal lumen; e, median esophageal
bulb; /, nerve ring; ö*, cardiac esophageal bulb; h,
beginning of the intestine; i, terminus; 7, large intes-
tinal granule; k, small intestinal granule; Z, anus;
m, spicula; n, excretory pore; 0, spermatocyte; p,
vas deferens; q, spermatozoon.

mestic species, and doubtless every

wild, has a number of specific nema-
tode parasites sapping its vitality.

THE VAST NUMBER OF UNKNOWN SPECIES.

The number of nematode species in existence must be

enormously greater than is commonly supposed. Since
most species of vertebrates are infested by one or more
species of nematode, and with relatively few exceptions
a given parasitic nematode infests but one host, it may be
 Neraatodes and TJieir Relationships, 469
estimated that more than 80^000 nematode species infest
the forty-odd thousand species of vertebrates. Insects,
much infested, will add many thousands of other species.
The mollusks, crustaceans, and various groups of worms are
also infested, and investigation continues from these sources
also to augment the number of known species of parasitic
nematodes.
Numerous as the parasitic species are, it is certain that the
species of nematodes living free in soil and in water far out-
number them; and the number of free-living individuals is

FIG.34.—This little serpentine monster, the dagger nematode (Xi'phinema) is able to coil itself
about the rootlets of plants in such a way as to use eiñciently its very long and slender spear,
which can be thrust far into the root tissues. The spear is clearly to be seen; its more slen-
der anterior part is exserted. This species is found in all parts of the United States and is a
representative of an injurious genus, found in many parts of the world. It is able to pierce
tougher and more corky roots than those nematodes which are armed with less formidable
spears.

SO great that they probably constitute one of the important

mechanical as well as biological factors in soil and in the
bottoms of lakes and oceans. Estimates based on the
writer's investigations show that in the upper foot of an arable
soil the number of nematodes runs to thousands of millions
per acre. Aquatic nematode species exist in enormous num-
bers in both fresh and salt water, while the number of indi-
viduals is past computation.
Of course these large figures are the results of estimates,
but the estimates are based carefully on past experience,
and they are, moreover, being continually justified by inves-
470 Yearbook of the Department of Agriculture.
tigations now in progress. A short time ago the writer
agreed to prepare a nematode chapter for a college text-
book devoted to North American fresh-water organisms.
On looking up the literature it was found that there was not
a single adequately described North American species of
fresh-water nematode. There was little time or oppor-
tunity to search specially for these fresh-water forms^ and
yet it has been possible in this short time to discover about
a hundred hitherto unknown kinds representing 30 genera.
This ready assembling of so many different genera and species
is fresh evidence of the enor-
mous abundance and mul-
tifarious nature of these
organisms. No doubt there
is a horde of species await-
ing discovery in North
American waters.
Something over a year
ago a new genus of nema-
todes was found attacking
citrus roots in California.
Within six months this
newly discovered citrus
parasite was located in such
widely scattered regions as
FiG. 35.—The banded nematode, Desmoscolex, California^ Florida, Spain,
one of a rather extensive group, not until 1889 Malta, Palestine, and Aus-
definitely recognized by anyone as nematodes.
Their internal organization is concealed be- tralia, showing, inciden-
neath a rather opaque cuticle. Their internal tally, that the parasite is
anatomy is of a typical nematode character.
one that has probably long
infested the citrus species and inadvertently has been carried
in nursery stock to many different parts of the world from
its original habitat, which is most probably the original
habitat of the cultivated citrus species themselves—south-
ern Asia. Here again the investigations have disclosed a
large number of new species, toward 100 in fact, assembled
on and about the roots of citrus trees in various parts of the
world.
These are not isolated and exceptional instances, but
typical cases. It will be seen, therefore, that it is something
more than a mere reconnoissance that leads me to the con-
 Nematodes and TTieir Relationships. 471

elusion that over nine-tenths of the nematode species still

remain unknown, a greater disproportion between the known
and the unknown than exists in almost any other class of
organisms.
There must be hundreds of thousands of species of nema-
todes. Of this vast number only a very few thousand have
been investigated, and of these, comparatively few with any
degree of thoroughness.

WONDERFUL VARIETY OF HABITAT.

Not the least interesting thing about nematodes is the as-
tounding variety of their habitats. They occur in arid
deserts and at the bottoms of lakes and rivers,, in the waters
of hot springs and in polar seas where the temperature is
constantly below the freezing point of fresh water. They
were thawed out alive from Antarctic ice in the far south by
members of the Shackleton expedition. They occur at enor-
mous depths in Alpine lakes and in the ocean. As para-
sites of fishes they traverse the seas; as parasites of birds
they float across continents and over high mountain ranges.
Their eggs and larvœ, invariably of microscopic size, are car-
ried from place to place by an exceedingly great variety of
agencies. Almost any visible thing that moves is capable
of transporting nematode eggs or larvae. Sometimes the
eggs and larvae are so resistant to dryness that long after
conversion to dust they may revive when moistened. This
revival of '^mummified^' nematodes may take place after as
long a period as a quarter of a century.
Nematodes are found in queer places. The wildest im-
agination could hardly outpicture the facts. One species
is found in the vermiform appendix of man; another has
its adult form only in the seeds of wheat. A third form
occurs in the felt mats on which the Germans are accus-
tomed to set their mugs of beer, and has thus far been
found in no other habitat. On the feet of birds and insects
the eggs, larvae, and adults of certain nematodes are carried
to the tops of the tallest trees. The sour sap issuing from
the wounds of a tree, often many feet above the ground,
not infrequently contains nematodes that are specific to the
wounds of that particular kind of tree. The tap water of
472 Yearbook of the Department of Agriculture,
even well-conducted cities often contains nematodes, for
nematodes are common in the potable water of our lakes and
rivers.
ABUNDANCE OF NEMATODES.

Nematodes, therefore, are extremely widespread, and to

be found in most unexpected places; they are also incon-
ceivably abundant. We little realize the enormous num-
bers in which these organisms exist all about us, wherever
we go, by land or sea. A thimbleful of mud from the bot-
tom of river -or ocean may contain hundreds of specimens.
The nematodes from a 10-acre field, if arranged single file,
would form a procession long enough to reach around the
world. A lump of soil no larger than the end of one's thumb
may contain hundreds, even thousands of nematodes, and
yet present few points that would distinguish it from a
lump of soil destitute of these organisms. As nema-
todes are usually very prolific, a single female ofttimes
producing thousands of eggs, the number of eggs vastly
exceeds that of the adults.
In short, if all the matter in the universe except the nema-
todes were swept away, our world would still be dimly
recognizable, and if, as disembodied spirits, we could then
investigate it, we should find its mountains, hills, vales,
rivers, lakes, and oceans represented by a film of nematodes.
The location of towns would be decipherable, since for
every massing of human beings there would be a correspond-
ing massing of certain nematodes. Trees would still stand in
ghostly rows representing our streets and highways. The
location of the various plants and animals would still be
decipherable, and, had we sufficient knowledge, in many cases
even their species could be determined by an examination of
their erstwhile nematode parasites.
We must therefore conceive of nematodes and their eggs
as almost omnipresent, as being carried by the wind and by
flying birds and running animals; as floating from place to
place in nearly all the waters of the earth; and as shipped
froiïi point to point throughout the civilized world in vehicles
of traffic.
It is interesting to speculate on what would happen if any
great group of plants or animals were utterly destroyed.
 Nematodes and Their Relationship, 473
and such a speculation may serve to throw Hght on the rela-
tive or economic importance of the group. We find micro-
scopic plants living in the sea deriving their sustenance
from material they find dissolved in the sea water. These
microscopic plants become the food of microscopic animals.
The microscopic animals become the food of others of larger
sizC; and so the series continues until it culminates in the
great monsters of the deep. These in turn when they die
may become dissolved in the sea water, and the material that
constituted their bodies may again be taken up by the micro-
scopic plants.
Tiny diatomaceous plants derive their sustenance from sea
water, nematodes feed upon the diatoms, fishes upon the
nematodes. What would happen if any link in such a chain
were destroyed ? It is quite conceivable that the destruction
of the nematodes might partially depopulate the ocean.
Could something of this sort possibly happen in the soil if
its nematode millions were suddenly destroyed ? It is con-
ceivable, but we do not know.
PREVALENT ERRORS WITH REGARD TO NEMATODES.
The prevalent conception of the nematode organization,
even that prevalent among scientific men, is fuU of errors.
These errors glare at us even from the pages of practically
all our most modern textbooks and encyclopedias. These
misconceptions have arisen in a natural way, because our
attention has been largely focused upon the parasitic nema-
todes. These parasitic species are simple in their external
structure, and while vastly larger in body than most of the
free-living species, are yet so small that the difficulties of
dissection are great if not insuperable. Accordingly, studies
have been carried on largely with the aid of thin slices made
with the microtome, a method both tedious and diSicult.
Progress has therefore been relatively slow, so that our
knowledge even of these deteriorated parasitic species is
often unsatisfactory.
It is a general law that parasitic species become more or
less degenerate. Depending as they do upon their host for
food and other things, they tend to lose the various organs
that would be necessary to them if they led a more active
existence. Thus it is that we have parasitic insect forms so
474 Yearbook of the Department of Agriculture,
degenerate that they possess neither wings nor legs. Their
eyes are rudimentary, as are also their antennae and mouth
parts. They are reduced to comparatively motionless worm-
like or saccate nearly colorless structures with few of the.
appendages that form so conspicuous a part of the anatomy
of an ordinary
insect.
What would
be our concep-
tion of the in-
sect group as
a whole if our
knowledge was
largely con-
fined to these
simple and de-
generate para-
sitic forms ? It
is easy to see
that we should have derived a
wholly erroneous conception.
Now, something of this sort has
actually happened in the case
of nematodes. While the more
complicated free-living species
outnumber the
parasitic spe-
cies ten to one,
our knowledge
of them has
FIG. 36.—Head (0) and tail end (&) of two difíerent nematodes, Ulus- lagged far be-
trating thé complicated anatomy of free-living species. Compare
these with the relatively simple parasitic forms shown in figure hind that of
37. It is by a study of free-living forms that we shall finally the parasites,
arrive at a just conception of nematode morphology.
partly because
they are minute and hidden away in unexplored locations,
and partly because of certain difficulties due to such a pecul-
iarly impenetrable cuticle that it is very difficult to prepare
satisfactory material for technical examination.
We have seen that the nematode organism is comparatively
complex; at the same time it is almost impossible to determine
 Nematodes and Their Relationships. 475
the function of some of the organs because of our inabihty to
make observations under natural conditions. The problem
of studying nematode habits and functions is in some re-
spects similar to that which confronts the geologist in his
attempts to form mental pictures of organic activities which
took place in remote ages. The geol-
ogist must study the relationship of
more or less fragmentary elements
in a dead condition, and imagine
what would happen if these fragments
were endowed with life similar to that
which comes within our experience in
the world as at present constituted.
It is somewhat the same with the
nematode anatomy. We have before
us for study a structure, sometimes
living, but more often dead, whose FIG. 37.—TWO parasitic nema-
functions we have to surmise on the todes, a male and a female.
These are typical parasitic
basis of analogous structures which we nematodes and represent the
may fairly assume to act in accordance ordinary conception of what
with methods to which we are accus- abeen nematode is like. We have
misled with regard to
tomed in a general way, in living organ- nematodes; they are by no
isms of another character and belong- ameans so simple in form as
consideration of the para-
ing to another group. sitic species would lead us
to believe. Compare this
Small wonder then that such con- illustration with figure 36
ceptions as we have should often prove which represents free-liviug
nematode forms. The com-
grossly incorrect. The prevalent idea plicated free-living forms
of the external appearance of a nema- more nearly represent the
tode, even among scientists, it is safe typical nematode structure.
to say, approximates to that illustrated in figure 37. As a
matter of fact, however, the illustrations on the opposite page
are quite as near the truth.
WHAT NEMATODES ARE LIKE; HOW TO FIND AND RECOG-
NIZE FREE-LIVING NEMATODES.
In answer to the question ^^ What are nematodes like,^^ we
can only reply that they do not closely resemble any other
organisms. While they constitute a group more widely
spread than almost any other, and are numbered by count-
less milHons, their relationship to the rest of the organic
world yet remains more or less of a riddle. We may compare
476 Yearbook of the Department of Agriculture,
the nematodes to an isolated oceanic island whose relation
to distant islands and continents remains problematical.
Studying the affinities of insects, for example, it is easy to
show that they are more or less closely related to spiders,
centipedes, and crabs, and these in their turn to other great
groups. Not so with nematodes. There is not another
group, great or small, to which they are known to be very
closely allied. This isolation in the present organic world is
one of the facts that may be used as an argument for the
great antiquity of the group. Nevertheless, there is an entire

FIG. 38.—Head end of the double-digger, or Diploscapter. This remarkable nematode nooks
its way through the sou and tissues of diseased plants by means of a solid piece of armor
on the front of its head. With this weapon it can chop both ways, first down and then up.
The extreme positions of the diggmg apparatus are shown by the dotted portions of the
figure. With each swing of its head the double-digger hooks its way forward.

absence of evidence that would afford direct proof of great

antiquity. No fossil nematodes are known.
HOW TO RECOGNIZE NEMATODES.
Practically any collection of soil, or any collection of sand,
mud, or débris from standing or from running water, in any
part of the world, will yield, on examination in water with a
hand lens, small slender organisms which whip themselves
about by means of more or less rapid contortions of the whole
body. This type of movement identifies them as nematodes
and differs from the movements of other small organisms of
similar form in that, though often vigorous'and conspicuous,
 Nematodes and Their RelationsJiips. 477
it is in one plane only, the dorso-ventral plane of the body,
and in that, unlike worms, the length and proportions of the
body meanwhile remain unchanged. In a clear liquid, more-
over, this thrashing about seems, as a rule, to produce no
locomotion; the nematode remains in about the same spot
unless it works its way in among vegetation, débris, or particles
of soil. It needs the friction of external matter of this char-
acter in order to accomplish its normal locomotion. When
quieted by stupefying, or killing, these nematodes are seen
to be more or less cylindrical, unsegmented organisms, with-
out locomotor appendages.
COLOR.
An internal examination shows that nearly aU the tissues
• of nematodes are comparatively colorless and transparent, and
whatever decided color the body possesses is usually confined
to the intestinal region. The cells of the intestine itself are
sometimes colored by the presence in them of organic granules
of a yellowish, greenish, or brownish tint, and the middle por-
tion of the body is thus made to appear yellowish, greenish, or
brownish. The color of the ingested food, showing through
the tissues of the body, is also sometimes a color factor; and
as the food varies in color from nearly black to colorless, so
the nematode is correspondingly tinted. Species feeding on
the juices of plants are usually nearly colorless, e. g., species
of Tylenchus and Aphelenchus. A considerable number of
species possess colored eye spots near the head. In some
species the œsophagus contains yellowish or brownish
pigment.
HOW NEMATODES WORK.
In answer to the questions How is it that nematodes do
so much harm? What are their methods? How do they
work ? it may be said they bite, puncture, gnaw, suck, and
dig as do insects, for instance, but they do all these things
with organs of an entirely different character.
The mechanisms with which they accomplish some of these
results are very interesting and the study of them constitutes
an important branch of nematology. Some of the forms are
shown'in figures 38, 39, 40, etc. When a nematode is pos-
sessed of definite jaws, these are usually three in number,
478 Yearbook of the Department of Agriculture,
instead of two, as in most other animal groups, and act radi-
ally somewhat as do the jaws of a lathe chuck. The jaws
are moved by relatively powerful muscles and often are
armed with ferocious looking teeth, which can be used in a
very effective way. (Figs. 42 and 44.)
Sometimes the mouth is armed with a sting or spear with
which to puncture the tissues of the victim, preparatory to
sucking away its vital fluids. In such cases, behind the spear
and constituting a portion of the gullet, there is a relatively

FIG. 39.—Head end of a spear-bearing nematode, showing how

the spear mechanism works. The spear is shown with its
acute point located in the midst of the lips, and appears in the
illustration to be about one-fourth of an inch long. The spear
is a hollow tube, and is connected with a very narrow chiti-
nous duct which passes back through the esophagus and
connects with the sucking bulb, which is in reality a kind of
force pump. This ellipsoidal sucking bulb is shown near the
bottom of the figure. The radiating lines shown in the bulb
represent muscular fibers. When these fibers contract they
open the central valve of the bulb and thus create a vacuum
cavity; this in turn creates suction, which extends forward
through the narrow tube just mentioned, and so on to the
spear, and hence to the lips. If the lips are placed against a
moist surface, such as the rootlet of a plant, and the suction
pump be then brought into action, the lips become fastened
to the moist surface by suction. The lips being now firmly
attached, the spear can be brought into play by the contrac-
tion of the muscles attached to its base. These are shown
surrounding the spear, and are attached to the base of the
spear at one end and extend obliquely forward to the base of
the lips, where they have their anterior attachment. The
contraction of these relatively powerful muscles drives the
spear forward and thrusts its point into the tissues fastened
against the lips. The suction of the bulb next causes the
fluid contents of the cells pierced by the spear *to now through
the spear into the esophagus and thence into the pump, which,
being e force pump, forces it on into the intestine not shown
in the figure.

powerful pump or sucking bulb. The nematode apphes its

lips to the object to be punctured, exerts suction by means
of its muscular pump, thus attaching its Ups firmly, and then
thrusts its spear through the mouth opening so as to puncture
or batter down the tissue containing its food, or through
which it wishes to pass. (Fig. 39.)
There is a series of remarkable species inhabiting the sous
of our meadows and swamps which possess a powerful spear
but no pump. They have developed another and very
 Nematodes and Their Relationships. 479

efficient method of using their

spears. These little Iotas, as I
have called them, are covered
with projecting, retrorse scales,
or prickles, so that it is with
difficulty that they move in any
other direction than forward.
Every movement of their bodies
drives them in a more or less
forthright way through the soil.
Coming against the root of a
plant, their muscular move-
ments push the head firmly
against the surface of the root.

FIG. 40.—One of the [scaly little monsters fre-

quenting the soils of our meadows and swamps.
These are known as Iotas. They are covered
with retrorse scales, or bristles, so that it is
practically impossible for them to move in.
any other direction than forward. Near the
head the remarkably large and powerful spear
can be seen through the skin. When, in
order to make punctures, this spear is thrust
out, the nematode is not pushed backward,
because of the friction which its scales offer to
surrounding soil particles. There are many
kinds of Iotas, and all of them appear to be
injurious.

so that the spear when thrust

forth acts from a well-supported
base, namely, the friction of the
surface of the body against the
surrounding soil material.
Sometimes the secretions or
excretions of nematodes are
irritating or poisonous, so that
their presence in the tissues
of animals or plants causes
abnormal swellings, or galls.
Their presence in the tissues of
animals may cause anemia and
lassitude, or muscular pains
and fever, to say nothing of
480 Yearbook of the Department of Agriculture.

special swellings, abscesses, punctured blood vessels, paralysis,

and insanity.

RELATIONSHIP TO FERTILITY AND BIOLOGY OF THE SOIL.

Notwithstanding the enormous number of nematodes ex-

isting in every acre of arable soil, the actual volume and
weight of the material composing the nematodes is relatively
small. We can not yet positively assert that they assist
materially in the fertilization of the soil; it is, however,
easily conceivable, in fact, there is a certain amount of evi-
dence for the idea, that, indirectly, some species may be of
considerable importance in maintaining or assisting to main-
tain a fertile condition. Unfortunately, however, we are
here driven to speculate on data that are anything but full
and satisfactory.
We must have some general notion, not only of the extent
of the nematode population of the soil, but also of its com-
position, before we shall be in a position to do more than
reason in a vague and unsatisfactory way upon this subject.
We need hundreds of investigators where now there are none,
and if these investigators should devota their time for years
to come to this subject alone it woidd only be after a con-
siderable lapse of time that our knowledge of these nematode
legions would be sufficient to enable us to reach reliable
general conclusions.
Since the great majority of the nematodes inhabiting the
soil are armed with an oral spear, whose sole function, as far
as we know, is to puncture other organisms, either animals
or plants, presumably plants, we may be strongly convinced
that they are on the whole injurious to the roots of the higher
plants. In specific cases we have positive proof that this
general belief is entirely correct. There are species of
spear-bearing nematodes which, by their injurious effects on
the roots of plants, cause an annual loss amounting to millions
of dollars. It is quite reasonable to suppose that the nema-
tode punctures, even when themselves not a serious setback
to the plant that is attacked, nevertheless form a highway for
the entrance of other organisms, such as injurious microbes
and fungi. In some such way we may account for the enor-
mous mortality of roots. It is an interesting fact that crop
 Nematodes and Their RelationsJiips. 481
plants in what we call good condition are found ofttimes to
have a large fraction of their roots in a decayed and useless
state. In such cases, where a root is destroyed, often the
plant throws out another root higher up on the same axis,
and in this way continues to derive its nourishment from the
soil. If the plant were freed from the necessity of constantly
supplying new roots in place of those killed off, to what ex-
tent would this release affect the aerial paxt of the plant ?
Cases are on record, and they are increasing in number,
in which it has been observed that if the soil be thoroughly
sterlized, and therefore freed, among other things, of its
nematode population, and be afterwards inoculated with
those microorganisms which are known to be necessary to
the health of growing plants, plants reared in it flourish
remarkably. May not this luxuriance be due to freedom
from injurious underground conditions, prominent among
which, we may easily imagine, are the attacks of certain
nematodes ?
Though nematodes are small they are scattered through
the soil in countless myriads in such a way that they must
constitute an important mechanical factor. From the time
they are hatched until death ensues nematodes seem to be
in constant motion. There is no evidence that they sleep,
and they rarely remain stationary for more than a few
seconds at a time, at least under the conditions in which
they come to our notice. Such active organisms, existing
as they do in every acre of arable soil in thousands of mil-
lions, must exert a more or less powerful mechanical influence.
MYSTERIOUS ORGANS—THE AMPHIDS.
Attention has already been called to the fact that nema-
todes possess organs the use of which we have been unable
to guess. The organs which I have called amphids, occur-
ring as rights and lefts like ears, one on either side of the
head, are almost universally present in free-living nematodes.
These amphids vary remarkably in size and form (fig. 16),
but not much in position. Their regular occurrence in such
a prominent location makes it exceedingly probable that
they serve some important purpose, but what that purpose
is we have not yet discovered. It is suggested by one that
75922°—YBK 1914 31
482 Yearbook of the Department of Agriculture,
they are organs of hearing, and by another that they are
breathing organs. It is quite conceivable that they are
organs of orientation; that is, organs by which the animal
knows what its position is with reference to the action of
gravitation—knows when it is right side
.V .^^ ^p Such organs of orientation occur in
higher animals, and are absolutely essen-
tial to their well being.
4^-1 ir P^rà It is possible that the amphids have to
4- ---i do with sensations or functions of which we
:j can form no clear idea. Mark the diflâ-
culties that accompany the investigation
y-^4: of such a matter. How are we to inter-

-vudOUtJOOOOOOOOOOUOCM
FIG. 41.—Six different forms of the
peculi^ organs called amphids, al-
^2^oqoooooooooo9flOQûûûD most universally present on the
heads of the free-living nematodes.
These organs are one of the mys-
teries of nematology. Nobody knows
what they are for. They occur like
ears, one on each side near the head
end. Sometimes they are of rela-
tively large size.

prêt the actions of a nem-

atode when trying to de-
termine its sensations
and motives, especially
if, as is not at all impossible, it is capable
of sensations so different from our own that
we can not even conceive of them? We
have no trouble in proving that a dog hears
with his ears as we do. Dissection shows
the structure of his ears to be closely like
that of our own. We observe how he re-
acts to various sounds—how he pricks up
his ears at a familiar call, or how he fails to
hear when his ears are in an unhealthy
state. In these observations we are guided by the dog's
actions, which we interpret in accordance with our knowledge
of our own feelings and actions under similar circumstances.
But how can we prove that a nematode hears ? How will it
act if it does hear ?
 Nematodes and Their Relationships, 483

It is true we can show after a fashion that nematodes, or

at least some of them, feel, see, taste, and smell, though in
these last two cases there is some doubt as to particulars;
that is to say, they act under appropriate circumstances as
we imagine they would act if they could feel, see, taste, and
smell somewhat after the human fashion. Moreover, we
have located and studied the organs through which these
sensations may probably be received. But no one has proved
that nematodes hear.
As might be surmised from the fact that nematodes live
under such a great variety of conditions, their food is ex-
tremely varied. Among the parasites, some feed upon living
blood, others upon various animal serums and secretions.
The plant parasites feed upon the sap and protoplasm found
in the tissues of the host plants. The free-Uving sorts some-
times feed upon small organisms of various kinds—microbes,
algse, the mycelium and spores of fungi, etc., each species
having its own particular preferences. Few are omnivorous.
Some species found in mud and slime extract food and nutri-
ment therefrom, somewhat as earthworms do, by swallowing
material indiscriminately.
NEMATODES IN TREES; HOW DO THEY GET THERE?
The nematode population of trees is worthy of account.
It is not generally known, in fact it is practically unknown,
that nematodes are found in trees, even to their very tops.
Their microscopic eggs and larvae are carried thither by cur-
rents of air, by insects, by birds, and by climbing animals;
or, keeping pace with the growth of the tree, the nematodes
may reach these high locations by their own efforts. Each
kind of tree may have its own peculiar fauna. It is well
known that nematodes are the cause of swellings or galls on
the aerial parts of plants as well as on their roots. These
galls have been a subject of interest for a long time, and have
been described and figured in scientific and horticultural
publications for half a century or more, but it is only recently
that we have come to understand how extensive is the
nematode tree population apart from the producers of gaUs.
The crevices in the bark, the natural clefts in the axils of
branches and leaves, aU furnish suitable lodgment for
nematodes, and the nematodes are there, no doubt often
484 Yearbook of the Department of Agriculture,
harmless, beyond doubt also sometimes detrimental. A few
examples may be cited to illustrate this fact.
Not long ago I had occasion to examine some diseased
apples which grew high up on an old apple tree, some 20 to
30 feet from
the ground.
I found the ap-
ples infested by
nematodes. By
experiment I
proved that
wasps and other
insects carried
the eggs and lar-
vae of the nem-
atodes from ap-
ple to apple on
the tree, and
from apples on
the ground to
others on the
tree. The in-
tensity of the
infestation may
be judged from
the fact that a
single apple
contained nem-
atodes to the
FIG. 42.—Front view of anematodemonsterwhichfeedsupon others number Ol about
of its kind. The drawing of the head is correct in every particu- QQ QQQ These
lar. The body has been sketched by the artist to assist the imagi- ^ '
nation in picturing how one of these organisms looks when seen Were OI Several
from in front. ThB artist's conception is undoubtedly very ¿iffOT»OTIf grjecies
nearly correct.. It will be seen«that this little monster
. possesses
^^ « ^ i'^ J
i
three jaws, as do most nematodes. This gives to the mouth SOmC of whlch
opening and pharynx of these organisms a peculiar three-angled -^^7-^1^^ well adaüt-
or three-sided appearance. The jaws are nearly always of prac- Y
tically equal size, although one of them is morphologically ed for forCing
equivalent to the other two. their wav U n -

aided into the apples, that is to say, were armed with mouth
parts specially adapted to puncturing and battering down veg-
etable tissue. Others belonged to genera characterized by
the secretion or excretion of material which appears to aid
in dissolving vegetable tissues.
 Nematodes and Their Relationships, 485

On another occasion, an examination of the "sap'^ issuing

from a wound in a chestnut tree, high above the ground,
showed that this liquid or semiliquid substance was swarming
with nematodes.
Finally, an examination of galls of unknown origin on a
chestnut oak showed them to be regularly inhabited by no
less than four different species of nema-
todes, some of which were of so peculiar

FIG. 43.—The devastating nematode of the onion and other

bulbous crops ( Tylenchus devastatrix). This nematode has
caused enormous damage in the Netherlands and other
parts of Europe, where it attacks the onion, hyacinth, and
numerous other plants. It has been known for a long time
in Europe and Australia; no means have yet been devised
by which it can be eradicated. Recently it has been found
also in the United States. It is one of the species which
puncture the tissues of plants by means of a long, narrow,
tubular sting, or spear, located in the mouth, I, A female;
//, head of the same more highly magnified, the spear is
most clearly shown; III, tail of a male; IV, vulva from
below; V, cross section of the neck passing through the
sucking bulb; VI, front view of the spicula and accessory
parts; VII, cross section through the middle of a female,
showing how the body cavity is filled completely by
tlieovary(ty)andtheintestine(z). a. Lip region; &, tipof
spear; c, medium sucking bulb; d, nerve ring; e, excretory
pore; /, muscles for protruding the spear; g, posterior eso-
phageal swelling; ii, excretory gland; ¿,hind end of spear; j,
loop in ovary; k, spiculum, or penis; I, muscles for opening
the vulva; m, the vulva; n, glandular (?) bodies; o, bursa;
p, hind end of ovary; q, uterus containing spermatozoa
and a segmenting egg (at r); r, segmenting egg; s, vagina;
t, the vulva or female sexual opening; u, blind end of pos-
terior rudimentary ovary; v, intestine, showing its cellular
structure; w, cross section of an egg; x, anus; y, wings of
the cuticle; z, cross section of the intestine.

a character as to strongly suggest that

these galls are their normal habitat.
Such cases, selected more or less at random from those
under investigation, are entirely typical and show conclu-
sively that our cultivated and forest trees from their roots
to their tops furnish lodgement and food for nematodes
peculiar to the situation.
EGGS ALWAYS MICROSCOPIC.
A most striking and important matter connected with the
life history of nematodes is the fact that the eggs are always
microscopic. There are nematodes which reach a length of
486 Yearbook of the Department of Agriculture.

several feet, yet their generative cells are always exceed-

ingly minute, comparable indeed with the spores of fungi,
or even with the larger microbes. This fact, together with
the enormous number of the eggs, is one of the most impor-
tant things to remember in connection with sanitary meas-
ures directed« toward diseases caused by these organisms.
We have to remember that they do not necessarily indicate
their presence by anything that we can see. One contracts
hookworm disease as one contracts typhoid, through an
invisible agency com-
posed of microscopic
elements capable of
causing the disease.
A field of beets,
onions, or potatoes
becomes infested, it
may be, from invisi-
ble eggs or larvse
planted with the
seed. The eggs or
larvae of nematodes
may be transferred
in all the numberless
ways rendered possi-
ble by their micro-
scopic size.
FIG. 44.-The shark nematode, Selachinema. This rapa- ^^^ ailierence DC-
cious little monster preys on other nematodes. The twCCn the size of the
specimen from which the illustration was derived may , A '^ fl?
have had an accident, as one of the jaws, that on the parent anu. llS Oil-
far side, appears to be missing. This nematode has been gpring is often VCrV
called the shark nematode on account of the rows of -'^^ ^ . *^
ferocious-looking, backward-pointing teeth with which great. There IS a
its jaws are armed. j-^^j^ UCmatodc COm-
mon in our brooks and ponds that has a total length of
about a millimeter. These little specks are the source of
another generation in which each individual is a thousand-
fold larger than either of its parents.
LENGTH OF LIFE.
How long does a nematode live ? Easy to ask, difläcult to
answer. How long does a vertebrate live ? It depends on
the vertebrate, whether it be a mouse or an elephant. The
 Nematodes and Their Relationships. 487

nematodes constitute a group probably ten times more

numerous in species than the vertebrates, and are corre-
spondingly varied in their size, habits, and life history, and,
naturally enough, vary also in the age to which they live. It
is certain that the eggs and larvae of some species of nematodes
will remain alive under suitable conditions for a very long
time. The larvae of certain nematode parasites have been
known to remain alive in a dry condition for approximately a
quarter of a century.
Certain forms parasitic in animals require more than one
host for their fidl development. They enter the first host
in a larval condition and after certain changes encyst them-
selves in the tissues of this host and enter upon a period of
quiescence, which no doubt may last in some cases for several
years, though just how long we do not know. These encysted
larval forms will not continue their development until their
primary host is devoured by a second host, as a wolf devours
a pig, or a bird an insect. On entering the second host the
encysted larval forms' develop, produce young, and die.
Here we have a series of events which normally occupies sev-
eral years and represents the lifetime of the parasite.
REMARKABLE INSTINCTS.
Certain other nematodes escape from their host in the form
of eggs or larvae and in this condition wiU remain alive for a
considerable length of time, it may be months, it may be
years. During all this time they undergo very little change,
and yet upon the arrival of suitable conditions they immedi-
ately spring into activity. I may illustrate this point by an
account of experiments I once made with eggs of the ordinary
Oxyuris of man. The eggs of this parasite on leaving the
host often contain immature embryos. These embryos
will grow slowly in water and other liquid media, but will
not rupture the egg shell. I endeavored to assist their
escape by cultivation in a great variety of artificial digestive
fluids such as pepsin, but all to no purpose; and yet when
these eggs were placed in a capsule and swallowed under such
conditions that the action of the stomach would cause the
capsule to be promptly supplied with fresh normal human
digestive fluid at body temperature, the eggs hatched in a
488 Yearbook of the Department of Agriculture.

X280 I
FIG. 45.—This savage little monster is found on or near the roots of plants, where it moves about
actively thrusting its head hither and thither with an almost inconceivably rapid motion,
hunting other nematodes, which it swallows whole. In the individual illustrated the
remains of several victims can be seen through the rather transparent walls of the intestine.
This is a beneficial nematode, the Mononchus papillatus of Bastian. It has been found
feeding upon Tylenchulus, another nematode injurious to the roots of citrus trees. The
illustration shows a rather immature femal espécimen which has been feeding upon Tylen-
chulus semipenetrans. The remains of three or four Tylenchuli are to be seen in the
intestine, a, Two of the innervated papillae existing on one of the six mobile lips; b, one
of the lips; c, dorsal pharyngeal tooth; d .one of the three longitudinal chitinous ribs of
the pharynx; e, pharyngeal cavity;/, esophagus; g, muscular layer of the body; h, cuticle;
i, one of the cells of the intestine; j, j,j, oral spears of three ingested Tylenchuli, the spear
in the intestine near the vulva is accompanied by an undigested portion of the lining of the
esophagus of the Tylenchulus; fc, nerve ring; Z, blind end of the anterior ovary, which,
being behind the intestine, shows less clearly than the posterior ovary; m, nucleus of one
of the intestinal cells; n, vulva; o, anus; p, terminus; q, anal muscles; r, rectum; s, cardia;
t, spicula of an ingested male Tylenchulus, the outlines of the undigested tail end of which
are to be seen faintly.
 Nematodes and Their Relationships. 489
few hours. So it is with many other nematode parasites.
They promptly recognize the specific conditions necessary to
their further development, but until those particular condi-
tions are fulfilled they remain quiescent. These quiescent
periods, of course, are an element in the age to which the
individual lives. Thus it is that while some nematodes
mature and die in a very few days, others may live for many
years.
Attention has already been called to the fact that nema-
todes feed upon other organisms. In this way they may be
either injurious or beneficial; the net result from an economic
standpoint depends on whether the organism devoured is
itself harmful or beneficial. There is a nematode that feeds
upon the roots of citrus trees and is therefore injurious.
(Fig. 33.) Living in the orchards with this first and harmful
species is a second nematode that regularly feeds upon the
first, and is therefore beneficial, doing the orange grower a
good service. (Fig. 45.) Many injurious insects passing a
part of their life in the soil, such as wireworms and cutworms,
are infested with nematode parasites. These parasites in
their larval stage inhabit the soil, where they are picked up
by succeeding generations of the insect larvae. In so far as
these parasitic nematodes are injurious to the cutworms they
are helpful to the farmer.
Some nematodes therefore are beneficial. We do not
know much about these beneficial species, but enough to
render the prospect alluring. We know that some of them,
especially those living in the soil and in water, feed upon
baneful microorganisms such as injurious microbes and fungi,
and even upon other nematodes. Doubtless the relation-
ships among these various microorganisms of the soil are as
varied and intricate as we know them to be among larger
organisms.
Of course it does not necessarily follow that because a sou-
inhabiting nematode feeds upon injurious microorganisms
it is, therefore, necessarily beneficial, for while feeding on the
micro-organisms it may be the means of transporting and
transplanting them, just as birds and animals feeding upon
obnoxious plants may become a means of distributing the
seeds. The subject is full of intricacies.
490 Yearbook of the Department of Agriculture,
NEMATOLOGY.
The foregoing fragmentary sketch may indicate to the
student, as well as to the general reader, the vast number of
nematodes that exist, the enormous variety of their forms,
and the intricate and important relationships they bear to
mankind and the rest of creation. They offer an exceptional
field of study; and probably constitute almost the last
great organic group worthy of a separate branch of biological
science comparable with entomology—^nematology.