Respiration in Dodecaceria
page 2
B. L. Gasior
ABSTRACT
Respiratory rates of the common California intertidal
Polychaete Dodecaceria fewkesi were measured under different
experimental conditions using Warburg constant- volume
respirometry. At a constant temperature of 19°C, mean respir-
atory rates ranged from 69.08 to 648.61 1 09 / gram worm/
hour (wet weight), depending upon the environmental conditions.
No significant difference was found between the rates in and
out of their natural tubes in water, whereas in air the res-
piratory rate was higher out of their tubes. Respiratory rates
in water were significantly higher than those in air under
all conditions. Headless worms had a much higher respiratory
rate than whole worms.
Respiration in Dodecaceria
page 3
B. L. Gasior
INTRODUCTION
Dodecaceria fewkesi was first described from California
by Fewkes (1889) under the name Sabella pacifica. Moore
(1909, 1923) later reclassified it with regards to specie and
genera, and Berkeley and Berkeley (1954) recently elaborated
on Moore's description.
This specie is distinguished from others in the family
Lirratulidae by the presence of setae resembling acicular
spines distally spoon shaped, construction of its own calcar-
eous tube masses, and its dark green to black color in life
(Blake, 1975). Dodecaceria fewkesi lives in asexually repro-
ducing clones. It has not yet been experimentally established
uhether autotomery precedes regeneration or schizametamery
(Dehorne, 1933) occurs.
I chose Dodecaceria fewkesi for these experiments because
I was particularly interested in respiratory rates under dif-
ferent conditions and stresses experienced by an intertidal
tube dweller. There appears to be atleast one previous inves-
tigation of the influence of the natural tubal environment
on respiratory metabolism of Polychaetes (Fox, 1938), and a
few authors (Hyman, 1932 and Courtney,1958) have studied the
effect of life in glass tubes.
Respiration in Dodecaceria
page 4
B.L.Gasior
ME THODS
Worms were collected on April 25, 1976, from rocks at
the zero to negative one foot tide level along the boat works
beach of Cannery Row, Monterey Bay, California. A chisel and
hammer were used to break chunks of the calcareous tube masses
from the rocks. The worms were brought to the laboratory and
kept in aquaria of running sea water at an average temperature
of 12°C at Hopkins Marine Station.
In preparation for these experiments I chiselled off
smaller chunks of tubes from the colony a minimum of twelve
hours before each run. Uhen I needed worms out of their natural
tubes I placed the tubes with worms in 100% Mocl, solution
for approximately 45 minutes before carefully chopping up the
tube chunks into even smaller pieces, and extracting the worms
with a pair of fine forceps. I tried to be as gentle as possible,
but it is virtually impossible to cut off a portion of the
colony without harming at least a few of the inhabitants.
Animals in each vessel were weighed after extraction from tubes,
and before experiments run out of tubes. The animals were placed
on a KIMWIPE for ten minutes before weighing; the weight of
worms varied from 0.03 to 2.37 grams/ vessel. After extraction
from tubes I counted the number alive, dead, whole, and head¬
less, and weighed the tubular mass. On handling, these worms
secrete a pigment from the skin, which has been characterized
by R.P. Dales (1963).
page 5
Respiration in Dodecaceria
B. L. Gasior
Ihe duration of all experiments ranged from ten to twelve
hours between 0600 and 2400; readings were taken every hour.
Ihe vessels were immersed in a constant temperature water bath
at 19°C, and shaken at a rate of 50 per minute. Instant ocean
was the aqueous medium used throughout the experiments, and
the gas phase was air. Respiratory rates were calculated as
Al gram wet weighthour.
RE SULT
All results of these experiments are summarized in lable
I. These values for respiratory rate will be compared below
for the different experimental conditions used. However, one
must first consider the possible contribution of the respir-
atory rates of non-worm tubular material to the values given
in Experiments I and IV. To examine this question I determined
the respiratory rate of the tubular mass in the Warburg vessels
in 17 experiments. The resulting rates varied inconsistently,
and in some experiments live worms were found in the tubes.
According to Berkeley and Berkeley (1954), even one single
segment of Dodecaceria may have regenerative powers. Since
in breaking up the tubes, a single segment or more of worm
was often times left behind, I attributed the small respira-
tory rates obtained to this fact. No correction was therefore
applied for respiration of the tubes.
Respiration in Dodecaceria
page 6
B. L. Gasior
I. COMPARISON OF RESPIRATORY RATES IN AND OUT OF TUBES
OUT DF TUSES
IN TUBES
P VALUE
IN WATER
119.80
9.50
128.39
.o01
IN AIR
69.08
89.84
In water there was no significant difference between
respiratory rates in and out of their natural tubes. In air,
however, the respiratory rate of worms out of tubes was higher
than that of worms in tubes.
Perhaps the respiratory rates of worms in and out of tubes
in water were not significantly different because there is a
layer of water surrounding the animal in its natural tube,
and placing it in water out of its tube was not dramatically
stressful. I observed that worms out of their tubes in water
maintained their natural U-folded position, and did not aggre-
gate as they did in air.
The significant difference in rates in and out of their
tubes in air may be attributed to the left-over water in their
tubes as opposed to their semi-dry integument, which was most
likely a very stressful situation.
Respiration of Dodecaceria
page 7
B. L. Gasior
II. COMPARISON OF RESPIRATORY RATES IN AIR AND WATER
IN AIR
IN UATER
P VALUE
2.005
69.08
IN TUBES
119.80
OUT OF TUBES
89.84
2.005
128.39
Respiratory rates of worms in water were significantly
higher than those in air, both in and out of tubes, The be¬
haviour of Dodecaceria has not been studied in its natural
tube, however, perhaps D. fewkesi has the same capabilities
as stated for Arenicola and Nereis by Wells (1949). He contends
that both Arenicola and Nereis can, when the water level falls
below that of the burrow openings, resort to aerial respira-
tion by trapping bubbles in the water which remains in the
burrow. This ability could be important for intertidal species
experiencing long exposure, both for obtaining oxygen to main-
tain the standard metabolic rate, and in preventing CO, build-
up, but there are no experimental data. De-tubed worms placed
in air, assumingly, are under great stress because they are
not receiving a constant fresh supply of sea water. The slight
drying of their integument may be enough to initiate decreased
respiration. A decrease in activity may also explain the de-
creased respiration, but it is yet unknown whether activity
is substantially decreased when these worms are exposed at
low tide.
The comparison of respiratory rates of whole and headless
worms in air and water will be found in the following section.
Respiration in Dodecaceria
page 8
B. L. Gasior
III. COMPARISON OF RESPIRATORY RATES OF WHOLE UORMS WITH
HEADLESS UORMS
WHOLE
HEADLESS
P VALUE
96.29
IN AIR
.002
137.02
123.94
o01
IN UATER
648.61
P VALUE
.001
.001
A. IN UATER
It is interesting to note that the respiratory rate of
whole worms in water is not significantly different from that
of worms out of their tubes in water. However, headless worms
in water had a respiratory rate 5 times as high as whole worms
under the same conditions. I feel the remarkable significant
difference noted in the rates under these conditions can be
attributed, in large, to regenerative properties of this specie.
The standard metabolism during growth is known to be appreciably
higher than during adulthood (Hemmingsen, 1950 ).
Much research has been done on the role of a crown or
anterior branchial filaments in respiration.(Zoond, 1931 and
Wells, 1949a, 1952 and Courtney, 1958 ). Wells concluded that
the localisation of oxygen uptake depends more on behaviour
than anatomy (1952).
Respiration in Dodecaceria
page 9
B. L. Gasior
B. IN AIR
The significantly lower respiratory rate found for whole
worms as opposed to headless worms in air may be due to the
innate fact that whole worms are not regenerating. Being
exposed in air is a stressful enough situation to lead to a
decreased respiratory rate.
C. UHOLE UORMS
In comparing the respiratory rates of whole worms in air
and water one sees that they are in accord with those out of
tubes in air and water. The lower respiratory rate in air can
most likely be accounted for by a decrease in activity.
D. HEADLESS UORMS
The respiratory rate of headless worms in water is seen
to be slightly more than 5 times their respiratory rate in
aif. I believe they are maximizing efficiency of metabolism
while in water. There are most likely some essential elements
in water for regeneration, respiration, and other metabolic
functions that are missing in air. Perhaps their activity is
always highest while immersed in water, in compensation for
the time spent exposed during low tides. It would be to any
intertidal animal's advantage to function in this manner.
page 10
Respiration in Dodecaceria
B. L. Gasior
DISCUSSION
Bounhiol found respiratory rates of Cirratulids small
compared with values obtained for active errant forms of
Nereis and Nephtys, and for Arenicola, Amphitrite, and Sabella
pavonina. He attributed this low metabolic rate to the small
amount of muscular activity, but his results must be interpreted
with caution because of the inadequacy of methods for quanti¬
tative work in 1903. Courtney (1958) stated that the intertidal
Cirratulid Polychaetes Audouinia tentaculata, Lirratulus cir-
ratus, and Dodecaceria concharnum do not irrigate their burrows.
Both food and oxygen are obtained from the surface of the habitat
by means of branchial and tentacular filaments.
Many conflicting data have been compiled on respirometry
of various other Polychastous Annelids. Ewer & Fox (1940) con¬
tend that 0 consumption of Sabella varies with 0, tension,
while Hyman (1932) holds that the fall in 0, consumption with
time in Nereis virens is largely conditioned by some other
factor(s) than the fall in 0, content of its environment.
The worms showed a relatively constant respiratory rate over
the duration of each of my experiments. This trend is in accor¬
dance with the findings of Dolk & Paauw (1929). They found that
earthwormscare efficient breathers, respiring at a constant
rate down to 1/8 the 0, concentration of air.
Fox (1938) showed that O, consumption of S. spallanzanii
declined progressively after the removal of the worm from its
Respiration in Dodecaceria
page 11
B. L. Gasior
hatural tube. The decline of S. pavonina was shown to be
negligible during the first 5 hours of de-tubing. Hyman
(1932) , with respect to Nereis virens, showed that O, uptake
was higher in worms unenclosed than worms enclosed in glass
tubes. My results are not in accord with these findings for
a significantly lower value was not found for worms out of
their tubes.
CONCLUSTON
It seems as though the intertidal specie of Dodecaceria fewkesi
can adjust its respiratory rates to meet different stresses.
My experimental results seem more consistent with those of
the earthworm (Dolk & Paauw, 1929). Much more work needs to
be done on the respiratory rates of invertebrates, for there
seems to be no uniformity in the results relating 09 consump-
tion to 0, tension or habitat. As Hemmingsen suggests (1950)
and Courtney (1958) states, "...it is clear that among the
Polychaetes there is considerable variation in the absolute
values of the respiratory rates, owing to diversity of way
of life...'
Respiration in Dodecaceria
page 12
B. L. Gasior
ACKNOULEDGEME
I wish to thank the faculty and staff at Hopkins Marine
Station, and especially thank Dr. Fred Fuhrman for his
invaluable aid and loyalty as an adviser.
Respiration in Dodecaceria
page 13
B. L. Gasior
REFERENCES
Berkeley, E. and C. 1954. Notes on the Life History of the
Polychaete Dodecaceria fewkesi (nom. n.). Fish. Res.
Bd. Canada, 11: 920 - 334.
Blake, J. A. 1975. Phylum Annelida: Class Polychaeta. pp.151-
243. Chapter in Light's Manual. Intertidal Invertebrates
of the Central California Coast. Smith, R. I. and Carlton,
J. T. (eds). Univ. Calif. Press.
Bounhiol, J. 1903. Recherches biologiques experimentales sur
la respiration des annelides polychetes. Ann. Sci. nat.,
Paris. ser 8, 16: 1 - 131.
Lourtney, W. A. M. 1958. Certain aspects of the Biology of the
Lirratulid Polychaetes. (phD thesis) Univ. of London.
Dales,R. P. 1963. Pigments in the skins of the Polychaetes
Arenicola, Abarenicola, Dodecaceria, and Halla.
Lomp. Biochem. Physiol., 8 : 99 - 108.
Dehorne, A. 1933. La schizometamerie et les segments tetra-
gemmes de Dodecaceria
aulleryi, N. Sp.
Bull. biol. 67: 298 .
520.
Dolk,
H. E.,and Paauw, F. van der. 1929. Die Leistungen des
Hamoglobins beim Regenwurm. Zeitsch. f. vergleich. Physiol.,
10, S. 324.
Ewer
D. W. and Fox, H. M. 1940. On the function of chlorocruorin,
Proc. Roy. Soc., London. B 129 : 137 - 153.
Feukes, J. N. 1889. New invertebrates from the coast of Cali-
fornia. Bull. Essex Inst., Boston. 21: 99 - 146.
H. M. 1938. On the blood circulation and metabolism of
Fox,
Sabellids. Proc. roy. Soc., London. B, 125 : 554 - 69.
Hemmingsen, A. M. 1950. The relation of standard (basal) energy
metabolism to total fresh weight of living organisms.
Rep. Steno Hosp., Copenh., 4 : 7 - 58.
tension to 0 consumption
Hyman, L. H. 1932. Relation of 0.
in Nereis virens. J. Exptl. Zool. 61 : 209.
Moore, J. P. 1909. Polycheatous annelids from Monterey Bay
and San Diego, California. Proc. Acad. Nat. Sci.
Philadelphia. 61 : 235 - 294.
--------1923. The Polychaetous annelids dredged by the
U.S.S. " Albatross " off the Coast of Southern Ca. in
1904. 4. Spionidae to Sabellariidae. Ibid., 75: 179 - 259.
Respiration in Dodecacer
page 14
B. L. Gasior
Wells, G. P. 1949. Respiratory movements of Arenicola marin:
L. : intermittent irrigation of the tube and intermittent
aerial respiration. J. Mar. Biol. Ass. U. K. 28: 447 - 64.
Zoond, A. 1931. Studies in the Localisation of Respiratory
Exchange in invertebrates. J. Exptl. Biol. 8: 258 - 62.
1/15
mm
Ve
FIGURE I















t






.

S





ket


V


in




.


a
DODECACERIA
FEWKESI
650
500
500
400
300
5 200
100
——

9












-

IN AIR
5


IN WATE

FIGURE II
Figure I : Anterior portion of an Intertidal Dodecaceria
fewkesi as drawn from Light's Manual
Figure II : Mean Respiratory Rates of Intertidal Dodecaceria
fewkesi under different experimental conditions
TABLE I
EXPERIMENTAL
CONDITIONS
I. worms in tubes in air
II.
worms out of tubes in
air
III.
whole worms out o
tubes in air
worms in tubes in water
IV.
whole worms out of tubes
in water
VI.
worms out of tubes
in water
VII. headless worms out of
tubes in air
headless worms out of
VIII.
tubes in water
N*
11
13
13
14
MEAN RESPIRATORY RATE**
Al 0/m worm/hr
69.08 + 14.74
89.84 + 20.13
96.29 + 16.80
119.80 + 37.92
123.94 + 11.90
128.39 + 40.17
137.02 + 16.71
648.61 + 148.94
Of Intertidal Dodecaceria
Table I : Mean Respiratory Rates
under different Experimental Conditions
* N = number of determinations
Legend:
** value  standard deviation