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<I>Cancer Irroratus</I>

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<I>Cancer Irroratus</I> BULLETIN OF MARINE SCIENCE, 36(3): 454-466,1985 RHYTHMICITY UNDER CONSTANT CONDITIONS IN THE ROCK CRAB, CANCER IRRORATUS Steve Rebach ABSTRACT The existence of activity rhythms in migratory populations of the rock crab, Cancer ir- roratus, which spend most of the year in deep water, was investigated. Mature individuals from the Mid-Atlantic Bight were tested under laboratory conditions of constant light (LL) and constant dark (DD) after an initial ambient photoperiod was presented. Activity levels were simultaneously monitored for 20 crabs maintained in separate compartments, using an infrared beam-break system. Activity was analyzed using an Enright periodogram. Activity under LL was at a relatively low level and with no rhythmic periodicity exhibited. In DD, activity was concentrated at approximately 25-h intervals, approximating a tidal period. Under ambient (natural photoperiod) conditions, a 24-h rhythm was present with activity greatest during the scotophase and with peaks at dawn and dusk. Possible advantages to the presence of a tidal rhythmicity in a deep water species are discussed and include temporal partitioning of the environment, a timekeeping mechanism for the initiation of migration, and an evolutionary or ecological remnant from shallow water populations. The common rock crab, Cancer irroratus, is found from Labrador to South Carolina (Haefner, 1976) and is most abundant from Maine to North Carolina. Rock crabs inhabit cold water (50-15°C, Haefner, 1976) and in the mid-Atlantic region they live at depths of 10-700 m (depending on age and season), with highest densities between 40 and 60 m (15-30 km offshore) (Musick and McEachran, 1972; Haefner, 1976). Rock crabs in this region also undergo a yearly mass migration into inshore and lower Chesapeake Bay areas (depths of 1-20 m) in October/November and remain until March/April (Shotton, 1973; Haefner and van Engel, 1975). Since some rock crabs enter Chesapeake Bay, they can tolerate some degree of fresh water, but they do not survive in salinities lower than 200/00 (Haefner and van Engel, 1975). Molting occurs in this region from December to February (Winget et al., 1974), with females undergoing their yearly molt in December (Terretta, 1973) in preparation for mating. Molting and mating are confined to the shallower areas of the continental shelf. After mating occurs, the female returns to offshore waters. The males remain along the shelf, or move to even shallower areas, molting in January and continuing to feed in shallow water until spring (Terretta, 1973). A review of this migration can be found in Rebach (1983) and a synopsis of biological data on Cancer irroratus in Bigford (1979). Green crabs, Carcinus maenas, exhibit two rhythms, circa tidal and circadian, in which peaks of activity occur at high tide, with the greatest activity during a nighttime high tide. Over time, the smaller daytime peak increases as it approaches midnight and the larger nighttime peak decreases as it approaches dawn. Under constant conditions, the tidal rhythm persists for about a week. Crabs kept in the laboratory for a month showed a circadian rhythm with no tidal component (Naylor, 1958; 1962). Sand fiddler crabs (Uca pugilator) exhibit an activity pattern with directionality dependent on the stage ofthe tide (Hermkind, 1968; 1972). These directionalities can be exhibited in the laboratory, far removed from the beach. Brown et al. (1956) and Bennett et al. (1957) observed a spontaneous tidal locomotor rhythm in Uca pugnax. Statistical manipulation of the data uncovered 454 REBACH: RHYTHMICITY IN ROCK CRABS 455 an underlying, lower amplitude, diurnal rhythm (although some controversy still exists on this point-for a review of this aspect see DeCoursey, 1983). Other physiological systems in this genus exhibit circadian and tidal periodicities. Color change (melanophore activity) (Brown and Webb, 1948) and oxygen consumption (Brown et al., 1954) are but two well known examples. Evidence indicates a relationship between solar and lunar rhythms in Uca (Barnwell, 1966; Brown et al., 1953; Webb and Brown, 1965). Reviews of movements and rhythmicity in Uca and in Carcinus can be found in Palmer (1974), Creutzberg (1975), Rebach (1983) and Webb (1983), and of Uca in Hermkind (1972). Many workers have found both solar and tidal com- ponents in other intertidal species of crabs. For a review of these rhythms and the relationships between solar and tidal periodicities see Palmer (1974; 1976), Aschoff (1981), DeCoursey (1983) and Webb (1983). The rock crab, Cancer irroratus, spends 5 to 7 months each year offshore in deep waters, where photoperiodic cues are not readily available, and the remainder of its time in shallow coastal waters where it comes under the influence of both solar and tidal periodicities. Furthermore, populations ofthis same species appear to be non-migratory in New England (Turner, 1953; Saila and Pratt, 1973), where they spend the entire year in shallow water. In this study, animals from mid- Atlantic populations were tested for rhythmicity (as evidenced by locomotor activity). Terrestrial animals (including semi-terrestrial Crustacea) utilize circa- dian entraining agents such as photoperiod and, to a lesser extent, temperature cycles. Intertidal crustaceans usually do not respond strongly to light-dark cycles and entrain to artificial tides, simulated wave action, and cycles of temperature and/or salinity changes associated with tide (DeCoursey, 1983). This population was chosen because of its seasonal change of habitat (from deep to shallow waters) and the accompanying difference in available cues to which it is exposed. METHODS AND MATERIALS Rock crabs were captured at different times of year by bottom dredge from the New York Bight to the Mid-Atlantic Bight at depths that ranged between 20 and 70 m. The mean depth of capture was 23 m. Most samples produced an approximate I: I ratio of males to females. Some samples, especially those taken during the migratory period, resulted in ratios as high as 16: I male: female. The mean (±I SO) point-to-point width was 59.6 ± 14.4 mm. The mean weight (±l SO) was 39.9 ± 29.1 g, and the overall sex ratio was 1.8: I m:f. Gonadal inspection of males and the presence of eggs on females from the coastal areas near Virginia suggested that southern populations may mature at about 30 mm carapace width (Shotton, 1973). The size at sexual maturity is greater in the northern part of their range (60 mm, Scarratt and Lowe, 1972). AIl animals tested were sexually mature (confirmed by post-mortems). After capture, crabs were transported to the laboratory within 1-3 h in chilled, light-tight containers. They were maintained in combined housing and testing facilities consisting of duplicate 1,265-liter recirculating, temperature-controlled units, each capable of separately testing 10 crabs (each in an individual compartment) (Rebach, 1977). The two systems were maintained at the same conditions of temperature, salinity, and photoperiod. All activity testing series were conducted simultaneously on 20 crabs. Collection of data from individuals maintained in separate compartments, but under identical conditions (since each set of 10 shared a common water supply), permitted treatment as replicates. Many earlier studies used "actographs" of various designs for measuring locomotor activity. Most of these involved tipping containers, pie plates or buckets to make an electrical contact. These containers were usually very small and locomotor activity was severely restricted. The compartments (26 x 48 cm each) used in this study did not restrict movements and allowed recordings of natural activity patterns. Activity was measured by infrared beam-break using seven photoresistors (Clairex CL703L-peak sensitivity at 735 nm) wired in series and arranged in an "H" pattern under each of the 20 testing chambers. Four photoresistors were located at the comers of the compartment and three were placed across the center. An integrated circuit a,nplification system was slightly modified from Rebach (1977) to interface with a Hewlett-Packard HP-9815A desk calculator via an HP-98133A BCD interface. 456 BULLETIN OF MARINE SCIENCE, VOL. 36, NO.3, 1985 The calculator produced an hourly printout oftotal activity for each of the 20 test animals. An Esterline- Angus A620X 20-pen continuous event recorder was used as a backup, Between tests, each testing unit was cleaned, water quality analyzed, and 10% of the water replaced with freshly mixed artificial seawater. The crabs were fed daily on a randomized schedule with Crustacean Ration (Rebach, 1981). By using a combination ofrelays and elapsed time meters (Kessler-Ellis Products- ETMVS 15,13) connected to a dimmer switch and a reversible motor, a 20-min dawn was introduced before the photophase ("day") and a 20-min dusk before scotophase ("night") in photoperiod tests, Kavanau (1962) criticized the use of abrupt experimental transitions and demonstrated that LD cycles involving simulated twilight periods permitted a greater degree of experimental control. The times of onset and offset of dawn, day, dusk and night were recorded by the HP-98 I SA. At lights on, light intensity in the tanks was 250-300 lux (approximate light intensity at 10 m), depending on water clarity and turbulence. Pilot experiments conducted over a period of 9 months with a single group of animals held under ambient photoperiod indicated that when rhythmicity was exhibited, the peak became obvious after 1 week, reached its maximum definition during the second week and only began to decrease in amplitude after 6 months of testing. The length of the period (periodicity) was fairly constant. It was, therefore, decided to terminate activity tests as soon as possible after the rhythm was clearly defined (14 days). Some species do not continue to maintain their activity levels for such extended periods. Bregazzi and Naylor (1972) found that activity decreased after 2 weeks of testing in Ta/ilrus sa/talor.
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