THE RELATIONSHIPS BETWEEN POPULATION INCREASE AND

POPULATION DENSITY IN PLANT PARASITIC NEMATODES

II. Sedentary nematodes

BY

J. W. SEINHORST
Institute of Phytopathological Research, Wageningen, Netherlands

The areas of the host tissue exploited by individual sedentary nematodes overlap. For
sedentary nematodes with one generation at a time (e.g. Heterodera rostocbiensis)and penetrating
randomly into but not damaging the host tissue, the relation between the densities of subsequent
generations Pi and Pf is given by:
Pf = a (- ln q)-1 (1 - qPi) in which a = the rate of multiplication when Pi --->0 (maximum
rate of multiplication), q = 1 - p, p = the proportion of the total area available to the nematode
population, that is exploited for food at a density Pi = 1. As not all eggs hatch or die, a density
bPi must be added to find the total density after multiplication has been completed (b = the
proportion of the eggs that do not hatch or die during the experiment). Results of an experiment
by den Ouden & Seinhorst (1965) are in accordance with this equation but in most experiments
severe damage caused by the nematodes makes the relation between density and rate of multi-
plication more complicated.
Nematode.rmultiplying continuously
A model for the relation between density and rate of increase of sedentary and migratory
nematodes multiplying continuously and based on the same hypotheses as that for nematodes
with one generation at a time can be expressed as:
dx/dt = r (i - { (1 -I)inwhichx = P/E, u = qE, E = the
equilibrium density, P - population density, q = 1 - p, p = the proportion of the total avail-
able area exploited at a density Pi = 1, and t = time.
The ratios between initial and final densities derived from this equation for values of Xi
(= Pi/E) between 0 and 1 do not differ materially from those of the equation of the logistic
curve if as> 0.1.

SEDENTARY NEMATODESMULTIPLYING IN SEPARATE GENERATIONS

A model for the relation between the densities of two subsequent generations
of a nematode species multiplying in separate generations like Heterodera rosto-
clriensis can be based on three assumptions : 1) The "average nematode" is the
same at all densities. 2) The nematodes invade the host tissue randomly i.e. they
do not attract or repel each other. 3) The size of the second generation is propor-
tional to the amount of food taken up by the first generation. The first two
assumptions lead to Nicholson's competition model upon which Seinhorst (1965)
also based a model for the relation between nematode density and damage. They
are the simplest assumptions that can be made. The first could be tested by measur-

1) In = natural logarithm.
 158

ing rates of multiplication of inviduals from populations which differ in density

but have lived under the same external conditions. The second hypothesis could
be tested by observing the distribution of nematodes in or on roots. The third
assumption is simple mathematically but not biologically. However, it represents
the most efficient use of the food available to the population. The proportion f
of the available space exploited as a food source at a density P of the nematode
is then

in which p is the proportion of the available space exploited at a density P = 1.

If the capacity of the plant to produce food for the nematode population is in-
dependent of the density of the parent generation (i.e. the plant is not damaged
by the nematodes) the size of the second generation, if proportional to f (eq.
(12) ), is .

At very low densities (P - 0), f/lJ approaches to a maximum value as

(Rule of de l'H6pital)
As P f = cf, lim P f/Pi = - c In q. If the maximum rate of multiplication of
Piezo
the parent generation is called a, then a = - c In q, c = a / -In q (15)
'
and

The shape of the curve for the relation between log Pi and log P f according
to eq. (16) is given in fig. 1, which also contains a curve according to eq. (3).
Apparently the curves are very similar in shape.

This is the maximum value P f could have according to eq. (16). In further dis-
cussions it will be represented by the symbol M.
As P f / Pi < 1 if Pi is very high there is an equilibrium density E when
Pf / P, > 1 at low values of Pi, for which eq. (16) becomes

This equation cannot be solved algebraically but the value of L' can be determined
graphically.

1) Eqs. (1) to (11) are to be found in Seinhorst (1966).

 159

Eq. (16) can also be written

( y = -a-iPf In q, x = PtIn q, e = the base of natural logarithms).

If in eq. (16) and (18) u is substited for qE, xi for Pi / E and x f for P f / E then
In q = E-1 ln u, a = - ln u (1 - u)-1 and x f = (1 - u)-1 (1 - UXi).
The relation between the densities of two subsequent generations is therefore
determined entirely by the proportion 1 - u of the available food that is taken
up by the population when at the equilibrium density. The shape of the curves
obtained when plotting log Pf against log Pi (fig. 1) is therefore independent of
the values of a and q.

Fig. 1. Shape of curves according to equations (3) and (16). Pi and Pf on logarithmic scales
and in arbitrary units.

Eq. (16) does not take into account an influence of the degree of competition
for food on the growth and mortality in the parent generation, before the egg
producing stage is reached. This competition could have three effects :

a) the mortality of juvenile animals is not influenced by their density. Then,

if eq. (16) applies, the size and egg production of the adults of the parent genera-
tion are smaller the larger the density of this generation. The number of eggs
produced per animal n = c (1- gP2) / Pi.
160

b) the proportion of the available space exploited for food is in accordance

with eq. (12), but of the animals penetrating only so many survive, that all
adults are more or less of the same size irrespective of the initial density of the
population. In Heterodera the relation between egg and cyst density of the parent
-
generation Pi and C would then be C = c (1 and numbers of eggs per
cyst would be independent of initial density.
c) numbers of animals reaching adulthood and the size and egg production
of the adult females are smaller the larger the initial density of the population.
The food sources used by animals that die prematurely are not necessarily within
reach of those staying alive. The greater the initial density the more the final
density will then lag behind that expected according to eq. (16).

Two or more generations

The relation between the densities of three subsequent generations Pi, P2 and
P3 is given by

This equation cannot be reduced to one identical to eq. (19) and with x = cxPi.
Eq. (13) therefore only applies for the densities of two subsequent generations.
If not all nematodes attack the plant at the same moment, conditions for
attack and multiplication may be different for different groups of nematodes. The
final nematode density is then a sum of densities related to initial densities as
expressed by eq. (16) with different values of a and q:

This equation only reduces to one identical to (19) if qk and (P,)k have the same
values for all k's. This means that the number of nematodes that enter per unit
weight of newly formed roots should always be the same but that it does not
matter whether these nematodes produce the same number of eggs or not.
Terminating an experiment before all larvae have penetrated into a root or have
developed into cysts with eggs does not affect the applicability of eq. (16), but
variation in speed of root development without equivalent variation in rate of
hatching of eggs, penetration of larvae into the roots and development of these
nematodes does. Therefore the model also applies to Heterodera species with more
than one generation per year and Meloidogyne species during the period between
sowing or planting of a host plant and the invasion of the roots by the first larvae
of the second generation.

Influence of eggs zvhich do not hatch on the results of experiments

Equation (13) only applies to numbers of newly formed eggs. Not all eggs of
Heterodera populations hatch under the influence of a host plant. Therefore, if
 161

the initial population was not added to the soil as egg suspensions but was added
or already present as eggs in cysts, the final population will consist of a portion
according to eq. (10) and one b times the initial population (the eggs that did not
hatch). Therefore if the initial population Pi was present as eggs in cysts

By substituting q for q'(1-b), a' (-ln becomes a' (1-b) (-ln q)-x and by
substituting a for a' (1-b)

As b is often small (of the order of 0.2) the addition of b Pi becomes of importance
only when Pi is much greater than Pf e.g. when resistant potatoes are grown on
soil heavily infested with H. rostochiensis.

.
Comparison between theory and experiment
There are very few available data for testing the applicability of equations (13)
and (16) because of the severe damage that high densities of Heterodera species
usually inflict upon good hosts. This reduces rates of multiplication. In an ex-
periment reported by den Ouden & Seinhorst (1964) this damage was negligible,
the lengths of the plants and the yields of tubers being the same at all densities.
The rates of multiplication found at different initial densities are in reasonable
agreement with those according to eq. (13) (Fig. 2).
To avoid the complications caused by injury to the plant by nematodes,
investigations on the relation between initial densities of Heterodera and rates of
multiplication could be done with host plants having a low equilibrium density
or a high tolerance limit. Another way is to incorporate the influence of damage
on the rate of multiplication into the equation relating the densities of subsequent
generations and to compare the available data with this theoretical relat-
ion.

Influence of initial densiry on cyst content in the new generation

In the experiment by den Ouden & Seinhorst (1964) mentioned above the
soil was inoculated with suspensions of eggs of H. rostochiensis. All cysts found at
the end of the experiment were therefore new. The numbers of eggs per cyst
found at the different initial densities are given in fig. 2. They decrease only little
with increase of initial densities. Population increase was therefore largely
regulated by exclusion of so large a proportion of invading larvae from reaching
maturity that the remaining nematodes could develop to adults of about normal
size and reproductive capacity.
162

Fig. 2. Above: relation between initial and final densities, Pi and Pf, of Heterodera rostochiensis
according to den Ouden & Seinhorst (1964). Pi and Pf in eggs per 100 g of soil.
Below :relation between initial density of the nematodes and numbers of eggs per cyst in the
new generation.
 163

SEDENTARY NEMATODESMULTIPLYING CONTINUOUSLY

Of the sedentary nematode species only few have one active generation at a
time. In most of them generations overlap, often resulting in a practically con-
tinuous multiplication. In Meloidogyne and Heterodera species the generations are
separated spatially. The larvae only penetrate into root tips. Only the last speci-
mens of one and the first of a later generation could penetrate into the same roots
and compete there for food. Therefore eq. (21) applies to portions of the root
system during a certain period. The total population of new eggs is the sum of
P f's according to this equation over all root portions diminished by the number
of newly formed eggs that have already hatched. This means that there is a maxi-
mum rate of multiplication at low initial densities and an equilibrium-density.
Further, the relation between initial and final densities in this group of nematodes
depends on the values of the constants and variables in the equations to be sum-
med. Moreover this relation is almost always complicated by damage done by the
nematodes to the plant.
Hemicycliopbora species only attack and remain at root tips. The nematodes
move through the soil with the growing tips. All stages and different generations
occur mixed and compete for food. The nematodes may stay on one root tip for
at least several days, and we may therefore expect that the regions from which
different individuals withdraw their food overlap to a certain extent. Hemicyclio-
phora species may therefore be the least complicated examples of the group of
sedentary nematodes that multiply continuously. A model will be derived for this
case.
Of several species (e.g. Tylenchulus sp., Nacobbus sp., Sphaeronema sp.) certain
juvenile stages are migratory, the adult females sedentary. They multiply con-
tinuously. A model would have to consider both groups and their interactions.

Derivation of a model for the relation between population density and rate of increase of
sedentary nematodes multiplying continuously and of which all stages and generations attack
the same parts of the plant.
If nematodes move only occasionally from one place to another and do not
attract or repel each other when feeding, Nicholson's competition model again
applies for the relation between the density of the nematode and the proportion
of the food source from which food is withdrawn. The amount of food taken up
is according to (12) f = c (1-qP). As in migratory nematodes this food serves
primarily to maintain the animals already present. What is taken up in excess of
what is necessary for that purpose is used for increase of the population. If E is
the equilibrium density, then E animals take up a quantity of food c (1-qE) and a
quantity c (I-qE)E-i is needed to maintain a density 1. The rate of increase is
therefore
164

Dividing both sides of the equation by E and substituting x for PE-1 and u for
qE simplifies (22) to

Fig. 3. Relation between x (= P/E) and rt (in arbitrary units) according to eq. (2) (curve 1)
and as calculated from eq. (24) for
u= 3 X 10-1 (curve 2)
_ u = 1.3 X 10-1 (curve 3)
u = 2.3 X 10-3 (curve 4)
u = 3.5 x 10-5 (curve 5)
u= 7 X 10-1° (curve 6)

Values of dx/dt according to eq. (1) and to eq. (24) for r = 1 and different
values of u were compared. The smaller are the values of u the larger is the
deviation of the calculated rates of increase according to eq. (24) from those
 165

according to eq. (1). The same holds for relations between x and t and between
xand x; (Fig. 3,4). As eq. (24) cannot be solved algebraically the
relations between x and t were calculated on a computer for four values of u and
r = 1. The relations between xi and x f were derived graphically from the relations
between x and t, choosing a maximum rate of multiplication for all cases of 20
times. The curve according to eq. (3) and those derived from eq. (24) for values

Fig. 4. Relation between xi (= Pi/E) and xf (= Pf/E) in experiments of fixed duration accord-
ing to eq. (3) (curve 1), as derived from eq. (24) (curves 2 to 6, values of u as in fig. 3) and
according to eq. (16) (curve 7). For all curves a = ert = 20.

of u of 0,3 and 0,13 (fig. 4) could not be distinguished in experiments with nema-
todes (largest differences between values of x f 20% of that according to eq. (3) ).
The maximum area that could be exploited by one animal (in an infinitely large
- xx) x-1 or -In u times the area available
space containing food) is lim (1
.
166

per animal at the equilibrium density. The ratio between this maximum area that
can be exploited by one animal and the area exploited by one animal at the equili-
brium density is -In u (1 - For a > 0, (relation between x f and xi derived
from eq. (24) not distinguishable experimentally from that according to eq. (3),
Fig. 4) it is 2.7 or smaller.
The values of r and u are independent of each other except that r > 0 (increase,
when x < 1) for u < 1, and r < 0 (decrease at all densities) for u > 1.

Decrease of nematode density because of overpopulation

a) When r > 0 (a > 1)

Eq. (24) is based on theoretical considerations assuming that x < 1 and r > 0.
If x is very large the amount of food available per animal is negligeable and mor-
tality is independent of density. The curves for the relation between xi and x

Fig. 5. General relation between Pi and Pf for a> 1 (A), a < 1, .r < 1 (B) and a < 1, .r > 1 (C).
Drawn lines: no reduction of lifc span of nematode because of starvation; dashed lines: density
dependent reduction of life span of nematode because of starvation. Pi and Pf on logarithmic
scales. The places of the asymptotes Pf = aPi and are arbitrary within the limits
indicated for the values of a and set by the conditions c < a and c < 1.

plotted on double logarithmic scales have therefore two asymptotes parallel to

the 45 line (Fig. 5). The reproduction rate at very low densities and the mortality
rate at very high densities are unrelated. The curve diners fundamentally
- in shape
from one according to eq. (3) at high values of x.
The food taken up by the nematodes is used partly for non reproductive func-
 167

tions, partly for the production of offspring (development of juveniles and pro-
it
duction of eggs). If is assumed that the life span of the nematode is not reduced
as long as the quantity of food taken up suffices for the non reproductive functions,
eq. (24) will apply up to a value of x at which no offspring is produced any more.
If a proportion s of the quantity of food taken up by one nematode living in a
very large space is necessary for non reproductive life processes, then a proportion
x s (-ln u) of the space available for a population must be exploited for the food
necessary for these processes at a density x of the animal. There will be no off-
spring when 1 - ux = x s (-ln u).
If u = 0.3 and s = 0.5, x is about 1.5; if s is 0.25, x is about 3.5. At higher
values of x the life span of the nematode may be shortened because of an in-
sufficient food supply and the relation between density and rate of decrease
depends on that between food shortage and rate of mortality.

b) when r < 0
When r < 0, u > 1. Because of the definition of x it has negative values in this
case. At low population densities (high (negative) values of x) the rate of decrease
will again be independent of density. Decrease occurs because fewer young
animals are produced than animals die (s < 1, fig. 5B) or because no young animals
are produced at all, and the life span of the nematodes is shortened by food short-
age (s > 1, fig. 5C). In the first case eq. (24) applies again up to population densities
(down to values of x) at which no offspring is produced any more. At still higher
densities again the influence of food shortage on mortality determines the rate of
decrease.
Because of slight differences in the food source from place to place and irregular
distribution of the nematodes there will be no sharp boundary between the
densities at which offspring are still produced where the life span of the nematodes
is not shortened by food shortage and those densities at which no offspring is
produced where the life span of the nematode is reduced by food shortage.

Applying the model to migratory nematodes

The only difference between sedentary and migratory nematodes lies in the
time they stay in one place. If the nematodes do not attract or repel each other
the pattern of overlapping remains the same whatever distance specimens travel
between feedings. Eq. (24) is therefore a better model for the relation between
density of migratory nematodes and their rate of increase than eq. (1) (which can be
written dx/dt = r x (I-x) ). No simple set of logical assumptions about the
utilisation of food for maintaining the density already present and for increasing
the number of animals will result in eq. (1). If, nevertheless, the rates of increase
of animal populations found in
experiments !are in accordance with this equation
(Seinhorst, 1966) this can only be due to the fact that the constant u in eq. (24)
had values in these experiments that make the differences in the numerical
outcome of (1) and (24) of the same order or smaller than the statistical variations
168

of the results of the experiments, or that there are influences that change curves of
the shape of numbers 2 to 6 (Fig. 4) towards that of numbers 1 and 7. One, the
presence of temporarily inactive animals in the population, is discussed below.
Another could be that the nematodes do not act entirely independently of each
other. They may avoid feeding at the same place with another individual or
distinguish places which have been depleted of food from those which have not.
Feeding of two or more animals in one place may urge some to leave it sooner
than they would have done if alone. This could increase the efficiency of the
feeding and the rate of increase especially at the medium densities. At very low
densities there is very little overlapping of exploited areas, at densities close to
equilibrium a very large proportion of the food source is exploited and only little
increase is possible.
Eq. (24) is based on the same assumptions as Seinhorst's (1965) theory on the
relation between nematode density and the damage done to the plants attacked
and therefore retains uniformity in theoretical approach.
As the results of experiments are in agreement with both eq. (1) and eq. (24)
if u > 0.1 all that has been derived from eq. (1) (Seinhorst, 1966) still applies
if x is not much larger than 1. More generally it can be said that the relation
between initial and final densities in an experiment of a certain duration is always
represented by an S shaped curve with unequal branches and that two cases can be
distinguished; nematodes multiplying in separate generations which do not
overlap in time (eq. (21) ) and nematodes multiplying continuously (eq. (24) and
relations between xi and x f derived from it). The latter are represented sufficiently
accurately by eq. (3) (Seinhorst, 1966). Eq. (21) applies at all densities, eqs (3)
and (24) up to densities slightly above the equilibrium density. Fig. 5 of Seinhorst
(1966) shows that eq. (3) may also apply when a < 1.
Application of eq. (24) presupposes that the effect of population increase on
the rate of increase is instantaneous. This would mean that eggs hatch very shortly
after being laid and that the larvae take up as much food as the average individual
of the population. This is certainly not the case. Moreover numbers of eggs
produced per animal are most probably smaller the greater the density of the
population. The rate of multiplication will therefore be larger than is in accordance
with total population density (eggs + active animals). Curves as derived from eq.
(24) for the relation between initial and final density of nematodes in an experiment
may be changed by this to curves more in accordance with eq. (3). A period of
inactivity during moulting has the same effect as eggs that have not yet hatched.
The effects of time lags between production of offspring and their becoming
competitive are more fully discussed for the case of logistic population growth
by Wangersky & Cunningham (1957). They show that when the time lag exceeds
a certain limit, population density fluctuates with time. In experiments of fixed
duration a maximum final density is then reached at an initial density < E. With
nematodes this has been found when the host plant was severely damaged
(Chitwood & Feldmesser, 1948, Peters, 1961).
 169

Oostenbrink (1966) proposes the application of Fujita & Utida's equation

to nematode populations multiplying continuously. According to this equation

xf first increases and then decreases with increase of The only logical meaning
of the part bxi could be that a number of nematodes, proportional to the initial
density dies after population change (according to the first part of the equation)
has been completed, i.e. after the death or removal of the plants from the pots or
field plots or after the nematodes have stopped their reproductive activity. If the
nematodes multiply continuously the first part of the equation gives the final
density of the population until reproductive activity stops if the initial population
and its offspring do not differ materially in characters influencing the rate of in-
crease. The only difference then could be a different rate of mortality of parent
generation and offspring after reproduction ceased. This is an unlikely hypothesis
for nematodes multiplying continuously. Moreover, only in experiments of short
duration would a sizeable part of the final population still consist of animals
already present at the beginning. In Heterodera species the parent generation and
the offspring do not compete for food in the same way as migratory nematodes
even if the generations are not so well separated as in H. rostochiensis. If it is
nevertheless assumed that eq. (3) (the first part of eq. (25) ) gives an adequate
estimation of the total population at a given moment, the proportion of sedentary
stages (which are lost when the host plant dies or is removed) is not proportional
to the total density at an earlier moment. The surviving population could there-
fore not be found by subtracting a density proportional to that at an earlier mo-
ment from the total density (sedentary stages + eggs and free larvae).
In nematodes multiplying continuously the initial population could differ from
the offspring in longevity and reproductive capacity. There are two possibilities
then. If u (qE, page 164) is the same for the initial population and its ofFspring,
then the influence of the differences on the relation between initial and final
densities can be introduced in eqs (3) and (24) by substituting P'i = CPi (c = a
constant) for Pi and ckPi for the proportion k of the final population that consists of
animals which are already present in the'initial population (P' f = (1 - k + ck) P f).
This does not lead to an equation identical to eq (25). If the animals of the
original population live for a much shorter time or reproduce much more slowly
than their offspring, the curve for the relation between initial and final density
would have a maximum at a value of Pi < E. If u is diff'erent for the initial popu-
lation and the offspring a still more complicated situation arises. As there are no
data available to indicate that such a situation exists no further effort is needed in
working out their consequences. The data to which Oostenbrink (1966) applies
eq (25) are too scanty for fitting any curve (only three points). In Fhis other data
the existence of maxima of P f are either not statistically reliable (based on the
final density at only one initial density) or could be explained by severe damage
to the host (Seinhorst, in litt.). This limits the discussion on population dynamics
170

in plant parasitic nematodes to curves of the type given in figs. 4 and 5 and the
influence that damage could have on them.
Fig. 4 shows that all discussed relations between initial and final densities in an
experiment or observation period can be expected to be so similar that they can
only be distinguished in very accurate experiments. If one type of deviation
from a model is partly compensated by another deviation, no deviation could
possibly be demonstrated. For a discussion of the further consequences of the
theories proposed above it is therefore not necessary to distinguish between
different cases or elaborate on more complicated situations. In experiments of
fixed duration the rate of multiplication always approaches to a maximum a
(= ert) at very low densities. If a > 1 there is a true equilibrium density (E > 0).
If a < 1 the relation between initial and final densities still is represented by an S
shaped curve (Fig. 5). The smaller a is, the lower the densities will be at which
the two bends of the S are found and the closer the two asymptotes of the curve
will be to each other.
Measured rates of multiplication do not only depend on the ability of animals
to increase in number on a food source below a certain population density. If the
food source is a plant some time may elapse between the beginning of an experi-
ment and the moment the food source has reached a certain size. At the end of the
experiment some time may elapse between the removal of the plants and the
measurement of population densities. Mortality of nematodes during these
periods will lead to a reduction of the rates of multiplication measured in an
experiment.

The author wishes to thank Mr. C. A. van den Anker for his advice on some
mathematical problems, Mr. M. L. Potters and Mr. L. A. A. J. Eppink for the
numerical integration of eq. (24) on a computer, and Mr. den Ouden for supplying
unpublished data.

ZUSAMMENFASSUNG
DIE BEZIEHUNGEN ZWISCHEN VERMEHRUNGSRATE UND
POPULATIONSDICHTE BEI PFLANZENPARASITÄREN NEMATODEN

II. SedentäreNematoden
Sedentäre Nematoden mit Vermehrung in getrennten Generationen
Die Teile des Wirtspflanzengewebes, aus denen einzelne Exemplare einer sedentären Nema-
todenart Nährstoffe entziehen, überschneiden sich. Wenn die Beziehung zwischen Populations-
dichte und Volumen des beanspruchten Gewebes dem Nicholsonschen Konkurrenzmodell
entspricht und die Grösse der Nachkommenschaft diesem Volumen proportional ist, dann wird
die Beziehung zwischen den Populationsdichten von zwei aufeinander folgenden Generationen
Pi und Pf einer sedentären Nematodenart, bei der nur eine Generation auf einer Pflanze vor-
kommt (wie bei Heterodera rostorhiensis),dessen Tiere willkürlich ins Wirtsgewebe eindringen,
aber dieses nicht schädigen, wiedergegeben durch die Gleichung:
Pf = a (- In q)-1 (1 - qPi), worin a = die Multiplikationsrate bei Pi → 0 ist, a = 1 - p
und p = der Teil des den Nematoden zur Verfügung stehenden Gewebes, woraus bei einer
Bevölkerungsdichte Pi = 1 Nahrung entzogen wird. Weil nicht alle Eier schlüpfen oder ab-
 171

sterben, muss - um die gesamte Bevölkerungsdichte zu finden - noch eine Dichte bPi hinzu-
gefügt werden, worin b = der nichtgeschlüpfte oder abgestorbene Teil der Eier ist. Die Resultate
eines Versuches von den Ouden & Seinhorst (1965) stimmen gut mit dieser Gleichung überein,
aber in den meisten aus der Literatur bekannten Versuchen ist durch Schaden an der Pflanze
die zweite Generation bei grösseren Anfangsdichten kleiner als nach der obengenannten
Gleichung.
Nematoden mit kontinuierlicher Vermehrung
Ein Modell für die Beziehung zwischen Populationsdichte und Vermehrungsrate von Nema-
toden, die sich kontinuierlich vermehren, kann auf denselben Hypothesen gegründet werden
wie das Modell für sedentäre Nematoden, bei denen zur gleichen Zeit nur eine Generation auf
einer Pflanze vorkommt:

Hierin ist x = P/E, u = qE, E = die Gleichgewichtsdichte, P = die Bevölkerungsdichte,

q = 1 - p, p = der Teil des den Nematoden zur Verfügung stehenden Gewebes, woraus bei
einer Bevölkerungsdichte Pi = 1 Nahrung entzogen wird, und t = Zeitmass. Das Verhältnis
zwischen Bevölkerungsdichten am Anfang und Ende eines Versuches nach dieser Gleichung
weicht für Werte von xi zwischen 0 und 1 nur wenig ab von dem nach der Gleichung (3) in
Seinhorst (1966) (xf = a Xi { (a - 1)xi + 1 } -1, Xi = Pi/E, Xf = Pf/E), wenn u > 0,1.
Fujita & Utida's Gleichung (Oostenbrink, 1966) trifft nicht zu für Nematodenpopulationen.
Ein Maximumwert von Pf bei einem Wert von Pi < E ist bisher nur einwandfrei festgestellt
worden, wenn die Wirtspflanze stark durch die Nematoden geschädigt war.

REFERENCES
CHITWOOD, B. G. & FELDMESSER, J., 1948. Golden nematode population studies. Proc. helminth.
Soc. Wash. 15 : 43-55.
OOSTENBRINK, M., 1966 (a). Vermeerdering van plantenaaltjes op de geschade plant. Tijdsrhr.
PlZiekt. 72: 208-209.
- 1966 (b). Major characteristics of the relation between nematodes and plants. Med.
Landb.Hogesch.Wageningen,66: 1-45.
OUDEN,H. DEN & SEINHORST, J. W., 1965. De invloed van enkele systemische nematiciden
op de vermeerdering van Heterodera rostochiensisop aardappelen van Tylenchorhynchus dubius
op stoppelknollen. Med. Landb.Hogesch.en Opzoek Stns. Gent 29 : 810-817.
PETERS,B. G., 1961. Heterodera rostochiensispopulation density in relation to potato growth.
J. Helminth. R.T. Leiper suppl., 151-150
SEINHORST, J. W., 1965. The relation between nematode density and damage to plants. Nematolo-
gica 11, 137-154.
- 1966. The relationships between population increase and population density in plant
parasitic nematodes. I. Introduction and migratory nematodes. Nematologica12, 157-169.
WANGERSKY, P. J. & CUNNINGHAM, W. J., 1957. Time Lag in Population Models. Cold Spring
Harbour SymposiaonQuantitative Biology22, 329-338.