I I nt. Revueges. Hydrobiol.

I 70 1 1985 I 4 I 509-525 I
I;. I. LEBEDEVA and T. N. GERASIMOVA
Department of Biology, Moscow State University, Moscow, and Water Problems I nstitute of the
Academy of Sciences of the USSR, Moscow, USSR
Pcculiarities of Philodina roseola ( EHRBG.) (Rotatoria, Bdelloida) -
Growth and Reproduction under Various Temperature Conditions
keg wordfi : rotifers, individnal culturing (ontogenesis), growth, reproduction, temperature
Abstract,
J Iost investigations dealing with rotifer reproduction have been performed with populations
and not n ith individuals. At the same time the variations in the growth and reproduction rates
of rotifers are of considerable interest. This paper presents the results of investigations carried
out by culturing individuals of Philodina roseola. during the lifespan of an individual under different
temperature conditions ranging from 9 to 35 "C. The paper presents curves showing the growth
in length and weight, the periods of maximum growth rate and the maximum sizes of Philodina
as affected by temperature conditions. The reproduction rate is investigated at the same time.
The variation in egg evolution duration, in duration of the reproductive period and in the egg
laying rate are shown under all the conditions investigated. The data serve as a basis for estimating
the effect of temperature on the productivity of Philodina roseola.
Contents
1. List of Symbols . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 509
2. I ntroduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 510
3. Materials and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . 510
4. Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 511
4.1 Growth in length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 511
4.2 Growth in weight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 514
4.3 Reproduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 517
5. Discnssion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 521
6. Conclusions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 523
7. Suinniary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 524
8. Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 524
9. References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 525
1. List of Symbols
t "C temperatme, centigrades
t time, days or hours
age of an individual, days
length (weight) of the rotifer body at age t (here and below L in pin and W
(fresh weight) in mg . 10-4)
initial length (weight) of the post-embryonic rotifer body
L ( W)
Lo( W, )
extreme length (weight) achieved at 'G + - (used in connection witIl
Bertalanffy equation)
body width, pm
body volume pm3
V = 0.47 bLL
V = W, if 1 prn3=10-9 nig raw weight
specific rate of growth in length (weight) of an individual
absolute increase in weight (per day)
relative increase in weight (per clay)
duration of egg developnient up to the ~iionient of hatching
age at the first laying (prereproductive period), hours
generation time; D, = D, + D,
number of eggs
( 1 )
dou hling tinie:
1
*2 = I"dar
days
2. I ntroduction
Rotifers are an important element of freshwater ecosystems. They play an out-
standing role in the substance and energy cycles of water reservoirs. The rate of
energy transforuiation at this level and the production level are largely determined by
such an abiotic factor as temperature ( t o) .
The aim of the study was to identify growth and reproduction pecularitiee of
Philodim roseola (EHRBG.) within the whole interval of biokinetic teniperatures
during ontogenesis. According to some authors (LIPEROVSKAYA 1977, SCHAEFER and
PIPES 1973), the temperature range of tolerance for Ph. roseola is 5-35'. A t 38-40 "C
rotifers die within 2-3 days.
3. Materials and Methods
The present investigation was carried out by individual cultivation, each specimen being
contained in 0.5 ml media from birth until death. The culture of Ph. roseolu was extracted from
activated sludge at aeration stations. Individuals of a genetically homogeneous clone, the offspring
of young females in subsequent generations were used for the experiments, as young females in
their period of maximum fecundity give the most viable posterity (KING 1967). Rotifers were
adapted to the following temperatures during the lifespan of 2-3 generations: go, 14", 20, 2 6 O , 3 2 O,
35 OC. The culture media, prepared with water from B water supply, was renewed every day.
A week-old culture of ChZoreZZa vulgaris BEJ ER. with 3 pm cell diameter served as food for the
rotifers. I ts concentration was 11 . 106 cells per ml, or 0.16 mg of raw algae weight per ml. The
average number of bacteria was 1.2-1.3 . 106 cells/ml. The relation of bright and dark period
during a day was 16 : 8, the average illumination being 500 lux.
Each series started with 30 individuals; the number had decreased to 20 at 32 OC. All in all,
140 amictic feniales of Ph. roseola were used for the experiment. Each series was terminated u hen
all the individuals wcre dead.
During the experiment, L and b of each rotifer were measured at the moment of its complete
stretching, while under a microscope with 7 ~ 1 0 magnification. When reproduction began, the
eggs were removed and counted. I n addition, De, D, and t were determined for each temperature
interval.
Philodina roseola under Various Temperature Conditions
51 1
4. Results
4.1 Growth in length
Values obtained by daily measurements served as a basis for plotting the curves of
the growth in length of Ph. roseola at the temperatures investigated. The hatching
time was taken as zero and the size of a newly hatched rotifer ( 226x 34 pi ) as the
initial size. Hatching time is given with an accuracy of one hour. Figure 1 shows that
t oc La, Pm
- 74 566
- 20 588
%---x 26 588
3 2 520
&-.--A 35 41L
- Reproducti on period
o......o
200L I I I I > 3 I , , , , , , , , , , , , , , , , n
1
0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 2 2 2 3 24' 38 39 40
I ndividual length growth (L, pm) of Philodinn roseola females at different tempers-
tures. The confidence band was calculated at the 95 'J /o level of significance.
As e 171, dov5
Figure I .
the length of Ph. roseola changes greatly during the organism's lifespan and that i t is
teni p eratu re dependent.
The growth in length of Ph. roseola at the temperatures investigated and at 20 "C
(LEBEDEVA and GERASIMOVA 1981) can be approximately described by the Berta-
lanffy equation as modified for growth in length (WINBERG 1966, MINA and KLXVEZAL
1976) :
Coefficient k was determined at all temperatures from the minimal value of the rela-
tive mean-square deviation cr.
Lt = L,- (L, - L,) . e- kt
(2)
(3 )
i = i
512
L. I. LEBEDEVA and T. N. GERASIMOVA
L;=rotifer length measured on the i th day;
L: =theoretical rotifer length calciilated with the help of equation (2);
If = number of measurements.
The values of k and c at. the different experiniental teinperatures were as follows:
t "C lc n
14 0.136 0.0494
20 0.412 0.0174
26 0.363 0.0540
32 0.630 0.0389
35 0.720 0.0270
These figures show that k increases as the temperat,ure rises from 14 to 35 O. Equation
(2) can be used to calculat,e CL:
k (L,-Lo) eckt
L,-(L,-L,) e - k l
CL =
(4)
With the coefficient k and the known values of L,, and L, at the experimental teni-
peratures, we can use equations (2) and (4) to calculate the size of a rotifer at any age
and the specific rate of growth in length of Ph. roseolcc cultivated under similar
temperature and food conditions. Conversely, by analysing the composition of natural
and laboratory populations of Ph. roseoln cultured under similar conditions, we can
estimate (with a certain approxiination) their age and growth rate.
To perinit, analysis of the variations in the Philodina rate of growth in length during
their lifespan with the help of ernpirical data, the value CJ was calculated for each age
using the-following equation :
In &-In L,
t ,--t 1
CL =
(5)
where t,--t, =1 day, L, and L?=rotifer lengths (in p i ) at the beginning and the end
of a day.
The most intensive growth (according to the CL value) was observed at all the
temperatures during the first days of rotifer life. Figure 2 shows that one-day-old
rotifers had inaximum CL values for all temperature intervals (curve 1). I t is during
this period that the influence of teniperature on CL is inost evident. CL first increases
during the interval from 14 to 32C and then decreases at 35". During the second
day (curve 2) the correlations of the growth rates under different thermal conditions
change greatly. The sharp decrease in CL at the investigated temperatures occurs at
different times, hut always coincides with the beginning of reproduction. C, of breeding
females is not large and is practically constant.
It is known (BRODY 1945, VASNETSOV 1934) that the growth rate of animals is
related not only to their age but also to the sizes they achieve. Figure 3 shows that
the Philodinn speciniens 260-320 pm long have the highest CL values, the height of
the peaks increasing as the temperatlure rises from 14 to 32 "C. The earlier the onset of
puberty, the sharper was the drop in the CL of the specimens. The peculiarities of
rotifer growth at the lowest ( 1 4 O ) and highest (35 "C) temperatures manifested theni-
selves in the fact that the decrease in CL took place long before puberty in relatively
sinall females (260-270 p i ) and is, we think, an indication of the oppressive influence
of these teniperatnres.
Philodina roseola under Various Temperature Conditions
513
0,6
-
(0
x
0.5
W
+
0.4
0 14 20 26 32 35
Temperature ("CI
Figure 2. Specific rate of growth in length (C,) of individuals of different age, depending on the
temperature, for the
0- 0 first,
0- - -.second,
v---v third,
@--.*-a fourth,
0- * a - - U fifth,
B- - W sixth,
v ..***..'*. v seventh day of life.
3200.
. .
. . - .
. .
. .
26' x'
t oc
- 14
- 20
x--x 26
32
A--.-A 35
o.....o
- Reproduction
period
200 300 400 500 600
Length (L), Prn
Figure 3. Correlation between specific rate of growth in length (C,) and the average length of
an individual of Philodina roseola at different temperatures.
35 Int. Revue ges. Hydrobiol. 70 (1985) 4
514
L. I. LEBEDEVA and T. N. GERASIMOVA
4.2. Growth in weight
The investigation of factors governing growth in weight deserves special attent,ion
from the point of view of individual productivity estimation during ontogenesis, as
growth in weight directly characterizes the magnitude of somatic production. Curves
showing the growth in weight of Ph. roseola (Fig. 4) were plotted on the basis of the
t T ~~r n g . 1 0 . ~
- 14 36.6
- 20 35.3
X--X 26 35.3
O. . . . . O 32 25.3
&...-A 35
- Reproduction period
8. 2
I
I , , * ,
0 2 4 6 8 10 12 14 16 18 20 22 24 "35
Figure 4.
Age ( T I , days
Illdividual weight growth ( W) of PhiZodina roseoZu at different temperatures.
ineasureinents of individual length and width, taking into account the nonproportion-
ality of female growth, with the help of equation (I ). The figure shows that Philoclina
weight growth is most intensive at 20" (except for the 10th and 22nd days at 26 "C).
This exception is due to the fact that at 26 O reproduction is over by this time and is
accompanied by a slight acceleration growth in feniale weight. The beginning of
reproduction is accompanied hy a decrease in the somatic growth rate under all
temperature conditions, as the greater part of the energy is spent on generative growth.
When reproduction is over, a certain increase in the somatic growth rate is observed,
which stops after the extreme weight is achieved. This increase is evident at all the
temperatures except 35 "C.
I n order to analyse variations in the growth in weight of Ph. roseola,, the following
characteristics were calcnlated :
In W2- l n WI
t 2 - - t l
3w, c,v=
The results of these calculations for each 24-hour period are shown in Figures 5-8.
The value of the daily weight mass increase ( A W) during the lifespan varies consider-
ably (Fig. 5).
The general tendency of Cw variations during the individual lifespan (Fig. 6) mani-
fests itself in the fact that juvenile rotifers have the highest rates of growth in weight.
lndividuals aged one or two days had the highest CW values. The only exception was
Philodina roseola under Various Temperature Conditions
lo/ 9
il 7
0 I i
14 20 26 32 35
Temperature 1C)
515
Figure 5.
0-0 first,, a- . - . -0 third day, and
0- - -.second,
Absolute daily increase in body weight ( A W) of Philodina roseola individuals of different
ages at different temperatures, for the
V"."""" v the average values for the whole period
of growth.
at 14 "C. A rapid decrease in the growth rate at 32-35" is associated mainly with
early puberty in Ph. roseola specimens. Hence, both low ( 1 4 O ) 2nd high (33-35 "C)
temperatures inhibit the growth of young rotifers. This temperatures effect has dif-
ferent physiological causes, but the sanie "production consequences". When reproduc-
tion starts, Cw decreases sharply at all temperatures.
During the whole period of growth, the average daily C ~ V values increase as the
temperature rises. Figure7 shows that the average value of Cm at different ternperatu-
res is related to the total duration of the growth period and decreases regularly as the
duration of the growth period increases when the temperature falls from 35 to 14 "C.
This decrease is most abrupt in the temperature interval from 35 down to 26 "C, but
is less severe in the interval from 26 down to 14 "C.
Figure 8 illustrates how the average daily CFv values vary with the weight and length
of growing rotifers clearly showing that specimens weighing up to 10-4 nig had their
maximum CW values at 26-35 "C, whereas larger individuals achieved their Cjvmar at
temperatures of 14-20 "C. The Cw peak at 14-20", which is further towards the zone
of large PItilodina specimens, confirms the aforesaid regularity of Ph. roseola growth
at high temperatures, when the reduction in growth rate is connected mainly with the
onset of reproduction and not with the achieved size. Thus, rotifers weighing 8- 10~
~10- 4 rng are already reproducing at 26-32 "C, whereas at 14-20 "C they are still
juveniles.
3.5'
516 L. I. LEBEDEVA and T. N. GERASIMOVA
2 4 6 8 10 12 14 16 18 20 2:: ? *
t o c
- l i
- 20
X--X 26
32
o. . . . . <i
&- . - A 35
@ The beginning an0
8 tnp end of reproducti or, pe-icd
Age ( T I , doys
Philodinu roseolu at different temperatnres.
Figure 6. Variations in the specific rate of growth in weight ( C, ) during the whole lifespan of
1 -
8 16 2r+ 32
of the growth period at difierent temperatures.
Growth period durati on, days
The alignment chart (Fig. 8) and the rotifer size data make it easy to determine the
specific rate of growth in weight of rotifers of any size or age within the whole range
of experimental temperatures.
Philodina roseola under Various Temperature Conditions
517
Figure 8.
t oc
- 14
- 20
--+ 26
32
A-...-A 35
- Reproduction period
o......,J
L - Length, Drn, at different
temperatures
W- Weight, rng-?O"
4 8 12 16 20 24 28 32 36
w rng.10'~
t , I , I ;
-140 WO W- ? 0
NOrnPI 0
NmmC) U
W
W In
0
in in
0
0 0
5:
U
41
41 m
O
m m
0
0 Lo
5:
0 0
N O U . Y
LZOO'
+
L Z P 1 & g &
-
0 m
0 5 : m
in n
N ( ? V -? Lo
I_
00
- 4 . 3 U l nm
5: O N
L32' $
1350 -*
N N ?- 0.-
N m 0 U U
Dependence of the specific rate of growth in weight ( ( 7, ) of Philodina roseola on body
weight and length at different temperatures.
4.3 Reproduction
The data concerning the reproduction of Ph. roseola at the experimental teniperatures
are systematized in Table 1 and Figures 9 and 10. They show that all the parameters
recorded vary greatly with the temperature. When temperature rises up to 32 "C the
duration of development decreases, the acceleration in development being acconipanied
by a reduction in the total number of eggs laid during the whole lifespan ( Fr) and in
the total duration of the reproductive period ( Tr) .
Thus, a temperature rise from 20 to 35 "C leads to two contrary consequences:
on the one hand, the breeding rate is accelerated by high development rates, but, on
the other, it is decelerated due to the decrease in the total number of eggs laid ( E; ) .
Table I shows that Fr is reduced by a factor of 3.25 during the lifespan, if the tenipe-
rature rises from 20 to 32 "C. This decrease is especially sharp within the interval
86-32 "C. The value Faay& reaches a maximum at 26 "C (twice as high as at 20")
and then, as the temperature increases to 32 "C, it decreases owing to the smaller total
number of eggs laid. A further increase in temperature to 35 "C leads to the coniplete
518 L. I. LEBEDEVA and T. K. GERASIMOVA
Table 1. Philodina roseola reproduction and development under various
temperature conditions
Parameters
~~~
Temperature, "C
14 20 26 32 35
i.
2.
3.
4.
5.
6.
7.
Time of egg development
up to the moment of
hatching (De), honrs 114.8-t 9.31 24. 051. 61 21.1 t-0.85 19. 950. 72 20.7 S0.64
Age of the first laying
(prereproductive period),
Generation time
( D, ) , hours 187.0k12.13 72. 8k7. 87 51.1 k3. 03 33. 3t 0. 52 -
( Dq =DL +0,) , days 12.6 4.0 3.0 2.2 -
duration ( ! PT) , days 13. 95 3.03 16.8t1.40 5.8k1.40 2. 3t i . 21 -
Doubling time (!P2), days 7.79 2.71 1. 77 2.13 -
Reproductive period
Number of eggs
per day (Fday)
1 .G 2.9 5.5 3.1 -
Number of eggs during
the reproductive period
(FA 2 2 . 4 t 5.63 45.4k4.79 31.753.G8 7. 2k4. 41 -
Figure 9. Variation in the number of eggs (P) laid by a female per day during reproduction at
different temperatures. The confidence band was calculated at the 95 O/" level significance.
inhibition of reproduction in Ph. roseola: at this teniperature 43 o',, of the females aged
1-5 days had one or two eggs inside thein, but none of the eggs were laid.
The effect of low temperatures is accompanied by a sharp fall in the reproduction
rate (by all parameters). A t 14 "C developnient is decelerated (De, D, arid Dp increase)
and the number of eggs laid during both one day and the whole breeding period de-
creases. Thus, in spite of a sufficiently long reproductive period (T,= 13.9 days), the
total nuniber of eggs (F,) was less at 14" than at 20" or 26", but was twice as large as
at 32 "C. The average daily egg production ( pday) was lowest at 14" (as compared with
other teniperatures). The nuniber of eggs laid by a female per day ($'day/$!) at 14" is
5 times smaller than at 26 "C. A further reduction in temperature down to 9" leads to
Philodituz roseola under Various Temperature Conditions
519
4 8 12 I 6 20 24 28 32 36
Weight (Wl,rng.IO-'
0 critical body weight ( Wp) .
Figure 10. Fday dependence on body weight ( W ) during ontogenesis at different tempera,tures.
the cessation not only of reproduction but also of feeding and moving. Thus, a tenipe-
rature of 9 "C can be regarded as the biological zero for Philodina roseola.
The effect of temperatures, both high (32-35 "C) and low (14 "C) was accompanied
by disturbances in the egg laying process. The bodies of the specimens held at 1 4 O
and 32 "C were full of unlaid eggs, the number of which sometinies neared 11. I n some
cases the birth of hatched young individuals was observed, and this usually led to the
death of the females. I n UMEZAWA'S experiments (1953) Ph. roseola were kept in
distilled water at 20 "C, and viviparity was also observed under these conditions.
Rotifers were able to live without food or some time, but starvation was acconrpanied
by a reduction in their lifespan and a deceleration in their development. The transition
to viviparity is apparently a response to unfavourable conditions.
The effect of temperature on the duration of the reproductive period and on the
niiniber of eggs laid by a female during the subsequent day is evident (Fig. 9). The
reproductive performance during the whole lifespan of Ph. roseola is conipletely
different at low and high temperatures: at 26 "C we observed a quick increase of
Fd& with the peak on the 5th day; at 1 4 O we have a low, stretched curve of daily egg
production without a pronounced period of maximum egg production. The curve for
Fdayl$2 at 20 "C occupies an intermediate position. Figure 9 shows that reproduction
is inhibited at 32"; the reproductive period starts and ends at the same time as for
26 "C, but Fday is much lower.
Figure 10 presents the correlation between the weight of a rotifer body and the
Fday value. It is evident that, after puberty and the onset of reproduction, Fday in-
creases with increasing weight of the Ph. roseola body. The dependence of Fday on W
is almost linear at this period. For each temperature there is a critical weight ( Wp) of
Yh. roseola. When W reaches the value Wp, Fday becomes a maximum. The fecundity
(Fdaay) then decreases more or less sharply. The dependence of Faay on W when W = Wp
can be approxiniately described with the help of the following equation:
Fd a y =aW -t C (6)
The coefficients a and C were determined for all of the experimental teniperature
from the condition of the minimum value of the relative mean-square deviation.
The values of a, C and u were as follows:
520
L. I. LEBEDEVA and T. N. GERASI MOVA
t "C WF a lo4 C U
14 19.1 * 10-4 mg 0.182 -1.75 0.14
26 18.7 lo-* mg 0.380 0.80 0.15
32 11.7 * lO-4mg 0.362 -1.10 0.08
20 27.7 * 10-4 mg 0.223 - 1.50 0.12
As this table und Figure 10 show, coefficient a characterizes the effect of W on the
rate of Pdav increase. Thus, at 26 "C fecundity increases especially abruptly and a
reaches its niaxiniuni value. Figure I 0 shows clearly the inhibition of reproduction
at 32 "C when the WF value is ai a minimum.
Figure 11 presents the average rates of processes characterizing Pk. roseola, re-
9 11 14 20 26 32 35
Temperature ( " C )
Figure 11. Average values of rates (day-1) at different temperatures: I the doubling time (i/!P2);
2 hatching from an egg (l / De); 3 first laying (l/Di).
production and development during the whole reproduction period at each tempera-
ture. The hatching rate (curve 2, l/De) increases sharply when the temperature rises
from 14 to 20 "C. The increase is less abrupt when the temperature increases to 32",
and at 35" the l / D, value begins to decrease. The doubling time of Ph. roseola was
calculated for each day. We assume that the females lay eggs at uniform intervals
during the day over the whole reproduction period. Thus the value T2 reflects both
the duration of egg development and female fecundity. Figure 11 shows that the rate
of female doubling time (the average value for each temperature) increases uniformly
when the temperature rises to 32 "C.
5. Discussion
Somatic growth in Ph. roseola is a finite, decelerating process that takes place during
ontogenesis. The time at which L, and W , are reached varies greatly, depending on
the t,ernperature: the growth curves ascend steeply at 32-20 "C, but slope gently
at 14". Periodicit,y of growth is characteristic of Ph. roseola, as well as of other in-
vertebrates (PETCHEN and KUZNETSOVA 1966; IVLEVA 1969; BREGMAN 1971). This
periodicity manifests itself in the presence of two growth periods in Ph. roseola:
Philodina roseola under Various Temperature Conditions 52 1
prepuberal and post-puberal. The great influence of temperature on the growth rate
is very marked in the juvenile period. One and two day old Ph. roseola specimens had
the highest growth rate (1.560-1.357 . day-1). Apparently this value should be re-
garded as a physiological limit for the growth rate of Ph. roseola under the correspond-
ing conditions. The high CW value for juveniles at high temperatures (32-35 "C) is
observed even when growth and development processes are inhibited. I n itself, a high
ClTr value does not indicate the fact that this temperature range is optimum for the
vital activity of rotifers as, at these temperatures, growth ceases very quickly and W
becomes 1.4-4.3 times smaller than at 14-20 "C. The CW of adult individuals is not
high, and has approximately the same value at all temperatures. Ph. roseola should
be regarded as a quickly growing animal, judging both by its growth rate and by the
time at which i t reaches its maximum sizes. The deceleration of somatic growth at
the onset of puberty and at the beginning of the reproductive period is connected
with the limited energy budget of the individual. The higher the reproduction rate,
the more intensive is the prepuberal somatic growth of rotifers and the sharper is the
drop in CW during the second period. The reproductive effort when the temperature
rises from 26 to 32" apparently stresses the animals almost to the limits of their
abilities, because at these temperatures soniatic growth completely ceases in females
during reproduction. It should be noted that such intensive reproduction at 26-32 "C
is accompanied by an increase in female mortality during this period. This is confirmed
indirectly by the fact that, at 26", somatic growth in females is resumed after the
cessation of reproduction, until the rotifers achieve their maximum sizes, but at
20 "C somatic growth does not cease in slowly reproducing females. Apparently the
laying of 6-9 eggs per day at 26" exhausts the whole energy budget of the individual
(with the deduction of the energy spent on the metabolism), whereas the laying of
2-4 eggs per day at 20 "C does not completely exhaust the energy resources, sonie of
them being spent on somatic growth. A rise in temperature to 32" is accompanied by
a deceleration in generative growth and simultaneous cessation of somatic growth.
A further rise in temperature to 35 "C leads to homeostasis disturbances which niani-
fest themselves in the cessation of reproduction and a very short period of somatic
growth. Viviparity and rupture of the female body full of eggs are symptoms of com-
plete homeostasis break-down, the first signs of which are observed at 32 "C.
As the data show, the rates of growth, reproduction and developnient of Ph. roseola
differ greatly under different temperature conditions. Qlo-the van% Hoff I.H. tempera-
ture coefficient-is widely used to investigate the effects of temperature on the inten-
sity of the metabolism, growth and development of aquatic animals. The Qf0 values
were calculated in order to estimate quantitatively the effect of temperature on the
somatic growth rate and on the indices for the developnient and reproduction rates.
Understanding only too well that the connection between biological process rates and
temperature is more complex than in chemical reactions, we considered it expedient
to use QIo for the comparative estimation of the values at the experimental tempera-
tures, as well as for the calculation of characteristics at intermediate temperatures.
Qio was calculated with the help of the following equation:
where V, and V2 are the rates of the processes investigated at t ! and t : .
For the whole temperature interval (14-32") within which the growth and develop-
ment rates increase, QIo varies over a narrow range (Table 2). The mean Qio value was
2.6 for the development rate of Ph. roseola within the interval of 14-32 "C. It was
slightly lower for the somatic growth rate (Clv), this confirming the viewpoint of
MINA and KLEVEZAL (i976), who st,ated that, as temperature rises, the growth and
522 L. I. LEBEDEVA and T. N. GERASIMOVA
Table 2. The Qio values for the growth and reproduction rates of Philodina roseola
in various teniperature intervals
Process Temperature interval
rate 14 -20 20 -26 26 -32 32 -35 14 -32
~~
C W 6.80 1.27 1.55 0.44 2.37
1 ID, 13.58 1.24 1.10 0.87 2.65
1/D, 4.82 1 .SO 2.04 - 2.61
1 IDq 6.65 1.64 1.66 - 2.63
1I Fday 2.69 2.90
0.38 - 2.80
development rates accelerate a little sharper than somatic growth. I n order to conipare
quantitatively the effect of temperature on the rates of the processes investigated at
narrow intervals, the corresponding Qfo values were calculated for 14-20", 26-32"
and 32-35 "C. It turned out that the Qlo values are considerably'higher than the means
for most of the processes within the interval of 14-20" (Table 2). The rate of egg
development ( l/De) and the soniatic growth rate of one-day-old individuals (Cwi)
increase most sharply. The rises in temperature from 20" to 26" and from 26" to 32 "C
result in a much less pronounced acceleration of the growth and development rates,
which is in conforiiiity with the facts described in literature (WINBERG 1968, IVLEVA
1981).
A rise in temperature from 38" to 35" causes a drop in the growth (Cfv) and hatching
( i/De) rates : this temperature interval is close to the upper mortality threshold.
It corresponds to the tolerance limit in the zone of high teniperatures. The high (Ilo
values for the growth and reproduction rates of Ph. roseolu within the interval of
14-20" can be explained by the fact that t o = 14" is situated close to the lower bound
of biokinetic temperatures for Ph. roseola, as at 9" they stop growing, reproducing and
moving. The QIo values within the temperature interval of 20-26" appear to be most
significative of Ph. roseola, as here the growth and development rates increase uni-
f ormly .
The general tendency is that the increase in the growth and development rates
takes place against the background of decreasing total fecundity during the reproduc-
tive period when temperature rises from 20 to 26 "C; a further rise in temperature
results in a decrease in Fday, too-acceleration is expensive! An additional rise in
temperature from 32 to 35 "C leads to a drop in Qlo, various processes of vital activity
being inhibited to various degrees: when generative growth stops and soniatic growth
is inhibited, the development rate of eggs laid earlier is only slightly reduced. The
juveniles seein to be able to tolerate heat better than the adults. I t should be re-
membered that the reduction in the growth rate of juveniles was not as pronounced
as that of adults under the conditions of low temperatures.
The calculation of Qlo reveals certain features of the variationsin the main character-
ristics of the growth and reproduction rates of Ph. roseola in response to increases in
temperature rises froin 14 to 35 "C. The values of Qt,, for the developiiierit rates (l/De,
i/Dl, l/T, ) and the somatic growth rate of juvenile individuals (Cfv) vary most clearly
and uniformly. Variations in Ql o for the total (F,.) and daily (Fday) fecundity were least
uniform. The growth rate increases inore slowly than the reproduction and develop-
ment rates when temperature rises.
The (Ilo values obtained for Ph. roseola permit the calculation of their growth and
reproduction rates for any temperature within the interval of 14-35 "C under similar
food conditions. The Qlo values for rotifers can also be used for modelling some pro-
cesses.
Philodina roseola under Various Temperature Conditions 523
6. Conclusions
The results show that temperature greatly effects the activity, growth and repro-
duction of Philodina roseola, who survives in rather a broad range of temperature.
As the biological zero for Ph. roseola 9 "C should be regarded, as at this temperature
the rotifers cease eating and moving.
A t 14 "C some processes of the organism are slowed down more severely than could
be expected on the basis of KROGH'S (1914) publication. Thus, the Ql0 value for the
Cw of juvenile rotifers was 6.8 within the interval of 14-20 "C. The hatching rate
(l/De) decreased even more abruptly. The Ql0 value for 1/D, was 13.6 within this
temperature interval. The survival and average lifespan of an individual was much
lower at 14" than at 20 "C. These figures show that the temperature of 14" is below
the limits of the optimal temperature range for Ph. roseola.
The highest survival of reproductive females and the maximum lifespan (48 days)
of an individual were observed at 20 "C. The growth of the females did not cease
during the whole reproduction period and reproduction proceeded normally. The sharp-
est increase in the growth and reproduction rates is also found in the interval of
14-20". This temperature (20 "C) is the optimum for lifespan, total increase in body
weight and the number of eggs laid during the whole lifespan of Ph. roseola.
At 26" growth and development accelerate, but the rates of these processes increase
more slowly than in the interval of 14-20 "C. The somatic growth rate remains practi-
cally the same, as a result of the early onset of puberty of the females and their
very high reproduction rate-the maximum Fday value. It is important that somatic
growth of the females continues after reproduction is over. A considerable shortening
of the prereproductive period (D,) and doubling time (T,) at 20-26 "C takes place
against the background of reduced female fecundity ( Fr) .
A t 32" the growth and reproduction acquire some peculiarities. The growth rate
(C,) of the juvenile individuals continues to increase, and the puberty and egg de-
velopment periods become shorter. However, these "satisfactory" quantitative
characteristics of growth and development are accompanied by distinct signs of the
inhibited oppressed state of the rotifer: early cessation of somatic growth, high inor-
tality among breeding females, abnormalities in the egg laying process and a reduction
in lifespan to 5.5 days. Thus, for the majority of the processes investigated, 32 "C
is above the optimum temperature limits. Pirst and foremost, this temperature has
an depressive effect on the reproduction rate and lifespan of the individual. At the
same time, somatic growth of the juvenile rotifers proceeds at a very high rate.
A t 35 "C both the reproduction and growth of Ph. roseola are inhibited. The lifespan
is very short, and the rotifers grow mainly during the first two days after hatching;
the Qlo value for juvenile individuals is 0.44 within the interval of 32-35 "C. Reproduc-
tion is affected worst: hatching is retarded and eggs are formed only in a suiall
number of females. At 35" none of the females laid eggs.
Consequently, 35 "C is the critical temperature for Ph. roseola vital activity and it
is close to the tolerance limits in the high temperature zone. The hatching of young
rotifers, their growth during the first days after hatching, and the puberty of some
individuals can be considered as a form of adaptation in case the temperature de-
creases. Ph. roseola populations can exist for only a few days at 35 "C, as no reproduc-
tion takes place, although a certain amount of somatic production is formed and
young rotifers hatch from the previously laid eggs.
Ph. roseola has rather a broad range of tolerance. The temperature interval of
20-26 "C should be considered the optimum for growth and reproduction and the
temperatures of 14 and 32 "C as being close to the minimum and maximum tempera-
ture tolerance limits.
524 L. I. LEBEDEVA and T. N. GERASIMOVA
7. Summary
1. Pidoclaha roseola survives within a broad teiiiperatmre interval, which was
9-35 "C in our experiments. Solriatic growth takes place between 14 and 35 "C and
reproduction between 14 and 32 "C.
2. The growth of Philodina roseola in length and weight is finite. The teniperature
interval of 20 to 26 "C is the optiniuni for soniatic growth, the lengths of Philodinu
increasing froin 226 to 588 p i and their weights from 1.23 - 10-4 to 35.29. 10-6 ing
(2.6 and 28.7 times, respectively).
3. The growth in length of Philodinn. roseola can he described (with a certain approxi-
mation) by the Bertalanffy equation. The constant k increases from 0.135 to 0.700 as
the temperature rises froni 14 to 35 "C.
4. Pkilodina roseola has two periods of life : the first is a very short period of juvenile
rotifer soniatic growth, and the second one is the growth of adult fernales. The effect
of temperature on the growth rate is evident in the first period but is less clear during
the second.
5. The rnaxiniitni number of eggs laid by a speciinen during its lifespan was recorded
at 20 "C. The potential reproductive perforniance of the rotifers is realized only
partially under the conditions of high and low temperatures. The effect of low teinpe-
ratures (14 "C) manifests itself in a reduction of the reproduction rate by all the para-
meters. When the temperature rises froin 20 to 35 "C, Philodina roseola reproduction
increases on the one hand owing to the acceleration of developnient, but, on the other,
it is reduced because fewer eggs are laid during the reproductive period.
6. Temperature variations beyond the optiiiiuni interval (20-26 "C) affect egg
laying, causing the overfilling of females with eggs and the death of females. Thus,
breeding is most vulnerable: it is disturbed at 14 "C and 32 "C and ceases conipletely
at 35 "C.
7. The Qt,, value for the growth and development rates of Ph. roseoln. is 2.4-3.6
(within a broad range of temperature: 14-32 "C), i.e. i t is similar to the Qlo values of
other poikilothennic animals. The Ql 0 value for the six-degree teniperature intervals
reaches its niaxiniuni in the interval 14-20 "C.
S. Acknowledgements
We consider i t our pleasant duty to t~hitnk Dr. V. B. VASILYEV for his help in mathematical
treatment of our material.
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Philodina roseola under Various Temperature Conditions 525
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Dr. LUDMILA LEBEDEVA, Senior scientist
Department of General Ecology and Hydrobiology
Biological Faculty
Moscow State University
USSR - 117234 Moscow, Leninskie Gory
TATIANA GERASIMOVA, J unior scientist
Laboratory of Aquatic Ecosystems
Water Problems Institute of USSR Academy of Sciences
USSR - 103064 Moscow
Sadovo-Chernogryazskaya 1313
Manuscript accepted: August 15th, 1984