Camp. Biochem. Physiol., 1973, Vol. 44A, pp. 1185 to 1197. Perganon Press.

Printed in Great Britain

THE EFFECT OF BODY SIZE AND TEMPERATURE

ON THE RESPIRATION OF POLINICES DUPLICATUS

JUDITH D. HUEBNER*

Department of Zoology, University of Massachusetts, Amherst, Massachusetts 01002

(Received 11 July 1972)

Abstract-1. Although variable, respiration is proportional to the 0.536 power

of wet weight for Polinices duplicatus at all temperatures and seasons tested.
2. QrO is inversely proportional to size between 15 and 30°C and directly
porportional to size between 5 and 10°C.
3. Level of metabolism is directly proportional to temperature.
4. Crawling activity is 35 per cent less between 5 and 10°C than between 15
and 30°C.
5. There is no evidence of seasonal acclimation of respiration in P. duplicatus.
6. Possible adaptive significance and physiological mechanisms for patterns
of respiration found, and the possibility of a metabolic “switch” are discussed.

INTRODUCTION
THE TEMPERATE intertidal zone is among the most rigorous habitats with extensive
and rapid changes in temperature (Johnson, 1965 ; Green & Hobson, 1970) both
daily and seasonally, alternations of exposure and inundation, and rapid depletion
of oxygen with depth in the sediment (Brafield, 1965 ; Jansson, 1967). For these
reasons, only animals capable of physiological compensation for such environ-
mental changes live in this habitat. Polities duplicatus is a carnivorous proso-
branch from the intertidal zone along the east coast of the United States as far
north as Cape Anne, Massachusetts. Respiration of individuals from a northern
population was studied in order to (1) investigate respiratory responses of an
intertidal predator to changes in temperature and season and (2) collect sufficient
respiratory data for the construction of energy budgets for the species which will
be the subject of another communication.
In addition to temperature and season, animal size influences respiration in
animals. The expression M = aWb (M = oxygen consumption, W = body weight
and a and b are constants) defines the relationship between metabolism and body
size. A logarithmic plot of M and W results in a straight line whose slope b is the
coefficient of respiration. Although Brody (1945) and Zeuthen (1953) found a
respiration coefficient (b) of 0.67 applicable to homeotherms, suggesting that
respiration is dependent on the surface : volume relationship, Hemmingsen (1960)

* Present address: c/o Institute of Parasitology, Macdonald College 800, Quebec,

Canada.

1185
1186 JUDITH D. HIJEBNER

concluded that 0.75 + 0.015 was the usual value for poikilotherms. Overall, respi-
ration coefficients appear variable, even among closely related species (Barnes &
Barnes, 1969) but most fall between 0.6 and 1.0.
Metabolism increases with temperature as well as with body size (Krogh, 1941;
Bullock, 1955). In addition, temperature often alters the relationship between
weight and respiration (b-values) resulting in size-dependence of Qlo measured
over the same temperature range. (Q1s is the change in respiration for a 10°C
change in temperature.) As temperature increases, Qlo for various invertebrates
decreases (Akerlund, 1969), increases (Rao & Bullock, 1954) or remains unchanged
(Pamatmat, 1969), although an inverse relationship between Qr,, and temperature
is most common (Precht, 1958).
The relationships between respiration and body size and temperature are also
influenced by an animal’s activity at the time of testing (Newell & Northcroft,
1967; Newell, 1969) and by acclimatization to seasonal changes in field tempera-
tures. Simultaneous measurements of activity and respiration of invertebrates are,
however, often imprecise and diflicult to obtain. Moreover, variations in the physio-
logical state of the animals (e.g. reproductive, nutritional) at different times of the
year are often neglected in investigations of seasonal temperature acclimatization.

MATERIALS AND METHODS

Collection and mahenance of maa% in the laboratory
Snails were collected at approximately monthly intervals throughout the yesfr (1970-71)
near Indian Trail Landing, B-table Harbor, Maaoachueetta. The animals were kept
without food for 2-14 days in 3-4 1. of aerated artificial sea water in 7-1. plastic aquaria.
Aquaria were kept in the dark or in dim light at the desired holding temperature. Table 1
lista holding and test temperatures, which were normally those appropriate for the time of

TABLE l--SUMMARY OF HOLDING AND EXPERIMENTAL TEMPER.4~ USED IN RESPIRATION

WPBRIMENTS

Collection Holding temperature (“C) Test temperature (“C)

date (?r l*O”C) ( & 0*.5”C)

28 June 1970 20.0 20.0

11 July 1970 20.0 25.0, 30.0
24 July 1970 20.0 20.0
8 Aug. 1970 20.0 25.0, 30.0
28 Sept. 1970 15.0 15.0, 20.0
7 Nov. 1970 5.0 5.0, 10.0
9 Dec. 1970 5.0 5.0, 10.0
21 Feb. 1971 5.0 5.0, 10.0
8 Mar. 1971 5.0 15.0, 20.0
22 Mar. 1971 5.0 5.0, 10.0
24 Apr. 1971 15.0 15.0, 20.0
22 May 1971 15.0 15.0, 20.0
6 June 1971 20.0 5.0, 10.0
25 June 1971 20.0 20.0, 25.0, 30.0
 RESPIRATION OF POLINICES DUPLICATUS 1187

collection (based on Green & Hobson, 1970, and records of the Woods Hole Oceanographic
Institution). Additional determinations in March and June 1971, were performed at
temperatures approximating those of other seasons.

Apparatus and measurements

The bottom of a 2.8-l. Fembach flask was covered with a thin layer of clean gravel and
filled with filtered, aerated artificial sea water at the desired temperature. After a single
snail was placed inside, the flask was closed with a rubber stopper through which a Clark
type polarographic oxygen electrode (YSI SlA, Yellow Springs Instrument Co., Yellow
Springs, Ohio) was inserted. The flask was kept in a constant temperature bath except
when measurements of oxygen concentration were being made. During the readings, the
water was circulated gently by a magnetic flea. Measurements were made each hour (except
for the two experiments done at off-season temperatures for which measurements were
made only at the start and finish of the run) for 5 hr, or until (as happened at high tempera-
tures) the oxygen concentration fell below 1.5 ppm. At the beginning and end of each
series of experiments, the electrode was calibrated in air; dissolved oxygen was also measured
by the Winkler method at these times (American Public Health Association, 1965). Read-
ings from control flasks containing only artificial sea water and gravel showed no oxygen
consumption, indicating that only the following small corrections for changes in electrode
calibration during the experiments were needed: -0.25 ppm at 5°C; -0.20 ppm at 10°C;
-0.05 ppm at 15 and 20°C; 0.10 ppm at 30°C.
Ten snails from each collection were used at each temperature. When respiration of
snails from one collection was determined at two or three temperatures, temperatures were
alternated daily until the series of experiments was completed thereby assuring that animals
tested at various temperatures were exposed to laboratory conditions for similar lengths of
time. If the experimental temperature differed from the holding temperature, the snail
was allowed 1 hr at the test temperature in a separate container before measurements were
begun. In all experiments the snail’s activity was observed and recorded. A snail was
considered active if it was crawling at the time of the reading, inactive if it was motionless or
withdrawn into its shell and was not classified if it was on its back with its foot partially or
entirely expanded.
At the end of each experiment, snails were measured, and fresh weights were obtained
after removal of the shell and operculum. Dry weights were recorded after 72 hr at 60°C
and ash weights after 24 hr at 600-65O”C. Ash-free dry weights were calculated by differ-
ence.
Manual calculations were done with a Wang programmable calculator (Wang, Model
380). Regression analyses and analyses of covariance were done with a CDC-3600 computer
using UCLA Biomed series programs BMDOlR and BMD03R. Comparisons of slopes
and elevations of regression equations were performed according to Snedecor & Co&ran
(1967). When a common slope was applied to equations, new intercepts were derived from
the means of the data.

RESULTS
Following the convention set by Zeuthen (1953), oxygen uptake/hr per animal
will be referred to as metabolism or respiration and oxygen uptake/hr per g as
metabolic or respiration rate or weight-specific metabolism. Since the correlation
(I) between oxygen uptake and weight was consistently higher with shell-less wet
weight than with ash-free dry weight, all calculations are based on wet weights.
This should be taken into account when comparisons are made with the results of
others, since regression coefficients change when different measures of weight are
1188 JUDITH D. HLBNER

used. To facilitate comparisons, calculations were made for a “standard” animal

of 5 g wet weight. This choice was based upon the mean weight of all snails
tested (5.25 g).
Figure 1 s ummarizes the relation between respiration and fresh weight of
P. duplicatw at different temperatures and seasons. There are no significant differ-
ences among b-values for the temperatures and seasons tested (analvsis of
covariance, >-test, 5 per cent level), so all data (Table 2) were pooled to

FIG. 1. Respiration of P. duplicatus at temperatures from 5 to 30°C using common

respiration coefficient of 0.536. Figure Sa--5°C. b-10°C. ~-15°C. d-20°C.
-25°C. f-30°C. Dates of collection are designated as follows: A = Feb. 1971,
B = Mar. 1971, C = Apr. 1971, D = May 1971, E = June 1970, E’ = June 1971,
F = July 1970, G = Aug. 1970, H = Sept. 1970, J = Nov. 1970, K = Dec. 1970,
* = All tests at the designated temperature.

obtain a common respiration coefficient of b = 0.536. Since slopes for all March
tests (S-2O”C), 5°C in June and December, and 30°C in June 1971 did not differ
statistically from either zero or the common slope, the common respiration co-
efficient was applied to them as well.
Respiration increased directly with temperature to a maximum at 25”C, as
illustrated for a standard (5 g) animal (Fig. 2) ; t = 0439 when the 30°C tests are
excluded. The Y-intercept (u-value), another indication of metabolism, was also
positively correlated with temperature; Y= 0.96 when means of u-values for the
same temperature, adjusted to b = 0.536 are used (Fig. 3). The reduced activity
of the snails at low temperatures may be partly responsible for the relationship:
 RESPIRATION OF POLINICES DUPLICATUS 1189

IO00 - rF-G E-F

l%O-
/
E’
600- H
4
&
\ D/
0" 400r
C/
zi

200-

I 1 I I I
OL I
5 IO 15 20 25 30

Tempemture, C

FIG. 2. The respiration of a standard (5 g) P. duplicates at various temperatures

and seasons using the common respiration coefficient of 0536. Symbols for
collection dates as in Fig. 1.

26-

2.5-

2,4-

f 2.3 -

> 2.2-
b
P 2.1-
8
a PO-
1.9-

I.8
0 5 IO 15 20 25 30
Temperature, C

FIG. 3. Respiration level (u-value) of P. duplicatus for temperatures of S-25°C

using common respiration coefficient of 0536; a = 1*72+0*03 (“C).

snails were active at 88 per cent of the readings made at 15-30°C but were active
at only 52 per cent of the readings at 5 and 10°C. Animals were inactive at 3 and
34 per cent of the readings at 15-30 and 5-lO”C, respectively.
Since analysis of covariance indicated that a-values of the following samples
were not significantly different at the 5 per cent level, they were grouped: all 5°C
tests, all 10°C tests, all winter tests at 5 and lO”C, all 20°C tests except March, all
25°C tests, all 30°C tests (see Table 2).
1190 JUDITHD. HUZBNBR

TABLE 2-TERMS OF REGRESSION EQUATIONS: log Y = U+b lOg x, DESCRIBING THE RELA-
TIONSHIP BETWEEN OXYGEN CONSUMPTION ( y) AND WET WEIGHT (x) OF P. du~licuiur

Date and
temperature (adj:ed to
PC) N a-value b f SE Y b = 0.536)

5 10 2.01 0.329 f 0.218 0.49 1.91

J””
k
5
5
10
10
1.89
1.97
0.409 f0.116
0.322 f 0.462
0.70
0.26
1.82
1.84
A, 5 11 1.99 0.391 f 0.103 0.80 1.92
B, 5 10 1.72 0.695 f 0440 0.58 1.81
10 10 1.97 0.604 f 0.138 0.86 2.01
J””
I;,
10
10
10
10
2.10
1.74
0.514 f 0.083
0.995 f 0.168
0.91
0.91
2.09
1.98
A, 10 10 2.06 0.361 f 0.103 0.64 1.96
B, 10 10 2.01 0.432 f 0.533 0.31 1.95
I-5 15 10 2.28 0.735 f 0.121 0.92 2.42
B, 15 8 2.15 0.326 f 0.331 0.35 2,03
c 15 10 1.96 0.887 f 0.236 0.82 2.20
D, 15 10 2.35 0468 f 0.073 0.92 2.31
E, 20 10 2.11 0.865 f 0.220 0.83 2.35
E’, 20 10 2.34 0.663 f 0.190 0.80 2.42
F, 20 10 2.44 0.621 f OW8 0.98 2.50
H, 20 10 2.41 0.722 f 0.067 0.97 2.53
B, 20 9 2.27 0.278 f 0.258 0.38 2.14
c, 20 10 2.41 0.414 f 0.065 0.93 2.35
D, 20 10 2.53 0.348 f 0.079 0.86 2.44
E’s 25 10 2.48 0.615 f 0173 0.80 2.53
F, 25 10 2.37 0.778 f 0.096 0.95 2.53
G, 25 10 2.69 0.479 f 0.079 0.92 2.66
E’, 30 10 2.65 0.312 f0.153 0.61 2.48
F, 30 10 2.54 0~56Of0~112 0.88 2.60
G, 30 10 2.70 0406 f 0.114 0.80 2.66
All 5 49 1.94 0400 f 0.087 0.56 1.86
All 10 50 2.00 0.527 it 0.078 0.70 2.00
C, D 15 20 2.17 0.541 kO.116 0.58 2.25
All 20 60 2.44 o-531 f 0.050 0.81 2.43
All (exc;$ W
30 2.53 0.594 f 0.072 0.84 2.57
All 30 30 2.66 0.373 f 0.066 0.74 2.56
All E’ 30 2.36 0480 f 0.093 0.72 2.48

Collection dates are designated an follows: A = Feb. 1971, B = Mar. 1971,

C = Apr. 1971, D = May 1971, E = June 1970, E’ = June 1971, F = July 1970,
G = Aug. 1970, H = Sept. 1970, J = Nov. 1970, K = Dec. 1970.

QIO reflects respiratory response to changes in temperature and varies with

both size and temperature for P. duplziutus. Between 15 and 3O”C, Qlo is inversely
proportional to animal size, whereas between 5 and 10°C (except in March), it is
directly proportional to size (Fig. 4). This relationship of Qlo to size is eliminated
 IWSPIRATION OF POLINICES DUPLICATUS 1191

when all equations are adjusted to b = 0.536. Q10 is quite variable at any given
temperature, with no clear relationship to temperature (Fig. 5). Although there is
no broad temperature range over which QIO remains constant for P. duplicatus,
values rarely exceed 2.0 except at low temperatures ( < 1OT). I If Q,, is considered

IOOk (b) c)

5.0 -

I.0 -

Wet wt of PohMes, g

FIG. 4. Relationship between Q10 and wet weight of P. duplicaturat various

temperatures and seasons based on original regression equations. Figure 4(a)-
5-10°C. (b)--1%20°C. (c)-Upper three lines 20-25”C, lower three lines 25-30X!.
Symbols for collection dates as in Fig. 1.

4.01

*J
3.0 -

20-

I .o- PG

OE’

06’
I I I I I
5-10 IO-15 B-20 20-25 25-x,

Temwmtvre, C

FIG. 5. Q,, values for a standard (5 g) P. duplicatusat various seasons and

temperatures based on equations using wmmon respiration coefficient of 0.536.
Circles used for separate collection dates, crosses ( x ) for PI,, derived from all tests
at the given temperature. Symbols for collection dates as Fig. 1.
1192 JUDITHD. HUEBNER

over the classical temperature interval of lO”C, there is a loss of information (e.g.
Ql0 is lower for 20-30°C than for 20-25°C because respiration shows no further
increase above 25°C).
Both Qi,, and metabolism vary with season as well as temperature and animal
size (Figs. 1, 2, 4 and 5). Thus, in the fall when holding and field temperatures
(Green & Hobson, 1970) were lower than in summer, respiration at 20°C was
higher than in summer. When temperatures were still lower (e.g. in March),
however, respiration at 20°C decreased considerably. Though there is a general
decrease in Qis throughout the winter and summer and an increase through the
spring, trends are not very strong. When animals were tested at temperatures
inappropriate for the collection time, the relationship between size and respiration
tended to break down (e.g. 5°C in June, 15 and 20°C in March). In part this is
due to large variability and general immobility of the snails under these conditions.
In addition, respiration appeared to be depressed when animals were tested at
unusually high temperatures such as 15 and 20°C in March and 30°C in June.
Q1,,, activity level, and to a lesser extent, level of metabolism (u-value) (Palo-
heimo & Dickie, 1966) all exhibit a discontinuity between 10 and 15°C. Thus,
Q usually increases with animal size at 5 and lO”C, but decreases with size at
teiperatures 2 15°C (Fig. 4). Activity (as defined in Materials and Methods)
decreases by about 35 per cent between 15 and lO”C, but is fairly constant at
higher and lower temperatures. In addition, the relationship between u-values
and temperature changes somewhat between 10 and 15°C.

DISCUSSION
P. duplicatus conform to the expected pattern of increasing respiration and
decreasing weight-specific respiration with increasing animal size (Fig. 1; Krogh,
1941; Brody, 1945 ; Rao & Bullock, 1954; Bullock, 1955). Studies of many inverte-
brates are consistent with Hemmin gsen’s respiration coefficient of 0.75 (e.g.
Roberts, 1957a; Berg & Ockelmann, 1959; Davies, 1966; Pamatmat, 1969;
Hughes, 1970), but others present quite different values (e.g. Dehnel, 1960;
Newell & Northcroft, 1965; Davies & Walkey, 1966; Akerlund, 1969; Rising &
Armitage, 1969). For P. duplicatus a common respiration coefficient of 0.536,
conforming neither to the range suggested by Hemmingsen (1960) nor Bertalanfiy
(1951, 1957), applies to snails from all collections regardless of season and tempera-
ture. Davies & Walkey (1966) proposed that such nonconformist respiration
coefficients be regarded as “deviation(s) from the phylogenetic b-value of 0*75”,
but available data appear to support neither a phylogenetic nor an ecological con-
sistency in respiration coefficients, e.g. the b-value is 0.536 for P. duplicatus, O-52
for the cestode Schistocephalus solidus (Davies & Walkey, 1966) and 0.95 for the
prosobranch mollusc Theodoxus fluwiatilus (Berg & Ockelmann, 1959). There is
currently no explanation for the various b-values found among poikilotherms,
which suggests that a variety of environmental factors and inherent physiological
mechanisms (Dehnel & McCaughran, 1964) are responsible for determining the
respiration: weight relationships of different species.
 RESPIRATION OF POLINKES DUPLICATUS 1193

Newell & Northcroft (1967) and Newell (1969) among others have divided
metabolism into active and standard phases. The former varies in the predicted
manner, directly with temperature, whereas the latter is considered relatively
temperature-independent (McFarland & Pickens, 1965 ; Newell, 1966; Halcrow &
Boyd, 1967; Sandison, 1967; Newell & Pye, 1970a, b, 1971). This is not true for
all animals, however, as shown recently for the dipteran Calliphora erythrocephala
(Tribe & Bowler, 1968), the polychaete Diopatra cuprea (Magnum & Sassaman,
1969), the crab Cmcinus and the mollusc Pat&a vulgata (Davies & Tribe, 1969).
Whether there are two distinguishable phases of metabolism for P. duplicatus could
not be determined in this study, but depressed metabolism at 5 and 10°C may be
related to reduced activity at these temperatures. Halcrow & Boyd (1967) state
that the influence of temperature on invertebrate metabolism is an expression of
change in locomotor activity; the spontaneous activity on which “routine” meta-
bolism is based (Beamish & Mookherjii, 1964) is temperature-dependent (Halcrow
& Boyd, 1967).
Low respiration and respiration coefficients which do not differ significantly
from zero, as well as cessation of feeding (Hanks, 1953), suggest that for P. duplicatus
5°C is near the lower limit of activity and possibly of aerobic respiration. Lewis
(1971) obtained similar results for tropical gastropods at 20°C. There is also a
levelling off, or decrease in respiration at excessively high temperatures (e.g. 30°C
in summer and 20°C in winter) in P. duplicatus and other species (Read, 1962;
Newell & Pye, 1970a).
The Q10 of invertebrate poikilotherms may vary inversely (Akerlund, 1969;
Barnes & Barnes, 1969), directly (Rao & Bullock, 1954; Pickens, 1965; Mason,
1971) or not at all (Roberts, 1957b; Pamatmat, 1969) with animal size. In P.
dupzicatus it appears to vary inversely with size between 15 and 3O”C, but
(except in February and March) directly with size between 5 and 10°C (Fig. 4)
suggesting that large snails are more sensitive to temperature changes when
cold, or in winter, and small snails are more affected when warm, or in
summer.
Many temperate zone poikilotherms (Bullock, 1955 ; Precht, 1958; Beamish,
1964) exhibit an inverse relationship between Qlo and temperature. Other species
may exhibit a peak Q10 value at a particular temperature with lower values at both
higher and lower temperatures (Read, 1962). Although there is no clear pattern
for P. duplicates, QIO values are relatively low, between 1.0 and 2-O for most
temperatures, and substantially exceed 2.0 only between 5 and 10°C in November
and December (Fig. 5). Such responses to changes in temperature may be bene-
ficial to eurythermal species permitting individuals to preserve a fairly constant
metabolism when subjected to rapid fluctuations in temperature such as may occur
during a tidal cycle. There may be an especially advantageous energy saving for
large animals at high temperatures, reducing somewhat the amount of food they
need for maintenance metabolism. The physiological mechanisms responsible for
increased Q10 values at low temperatures may enable snails to respond to brief
warm spells with increased activity and possibly even feeding. Teal (1957)
1194 JUDITH D.HUEBNER

suggested that high Qr,, values may be important in the conservation of resources
by poikilotherms in winter.
Respiration coefhcients, Qis, level of metabolism and activity level of P.
duplicutus are all independent of season, although all but respiration coefficients
are influenced by temperature. While many species show season-dependent
changes in these parameters (Rao & Bullock, 1954; Berg et al., 1958; Newell & Pye,
1970a; Burky, 1971), others, like P. d&pZicutus,do not (&gal, 1956; Barnes ef al.,
1963; McFarland & Pickens, 1965 ; Davies, 1967). Newell & Pye (1970a, b)
showed that seasonal changes in temperatures of maximal metabolism and low
Qi,, values for both active and standard metabolism of intact Litiotina and M’tiruF
and cell-free homogenates were directly dependent upon acclimation temperatures.
In contrast to the subtidal D. cupreu (Mangum & Sassaman, 1969), in Littoka
and ZUytiIus this acclimation process occurs only after several days (Newell & Pye,
1970b), ‘which would make these intertidal molluscs relatively independent of
rapid daily fluctuations in temperature. Lack of metabolic changes in P. dupZicatu.s
after several days’ acclimation at holding temperatures in the laboratory suggests
similar compensatory mechanisms in this species. It is possible, however, that
gradual acclimation of snails to unseasonable temperatures was responsible for the
high variability during experiments run at such temperatures.
The consistent presence of a discontinuity in QiO, activity level, and to a lesser
extent, level of metabolism between 10 and 15°C suggests the possibihty of a
“metabolic switch” that changes the animal’s respiratory patterns to adapt it to
changes in temperature. It appears that temperature is the critical factor triggering
the “switch” since animals tested at the same temperature at different seasons
responded in the same way.
Recent studies (e.g. Hochachka & Somero, 1968; Somero, 1969b; Baldwin,
1971; Somero & Hochachka, 1971) suggest that alterations in isozyme proportions
(as well as changes in animal activity) may be responsible for general respiratory
responses to temperature, as well as phenomena such as seen between 10 and 15°C
and suggested for temperatures below 5°C in P. duplicatus. Temperature-
dependent interconversions of isoxymes (Somero, 1969a), favoring the isozyme
with the minimal K,,, at the acclimation temperature, could compensate for direct
thermal influences on the rates of metabolic processes.

Acknowledgements--I wish to thank Drs. D. Craig Edwards, Donald Fairbeim and John
L. Roberts for their criticisms of the manuscript and helpful discussions throughout.

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Key Word Indecc-Metabolism ; oxygen consumption ; respiration ; temperature ; size ;

QIO; Polinices duplicatus.