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Effect of Starvation on the First Larvae of <I>Homarus Americanus</I>

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Effect of Starvation on the First Larvae of <I>Homarus Americanus</I> BULLETIN OF MARINE SCIENCE, 61(1): 73–80, 1997 EFFECT OF STARVATION ON THE FIRST LARVAE OF HOMARUS AMERICANUS (DECAPODA, NEPHROPIDAE) AND PHYLLOSOMAS OF JASUS VERREAUXI AND J. EDWARDSII (DECAPODA, PALINURIDAE) Fernando A. Abrunhosa and Jiro Kittaka ABSTRACT Food is one of the important factors controlling decapod larval culture, however, little is known about the effect of the starvation regimen on the physiological condition of the lar- vae. In the present study, the influence of starvation upon survival rate and the intermolt period was observed in the first instar of the American lobster, Homarus americanus, the first instars of phyllosomas of the red rock lobster, Jasus edwardsii, and the green rock lobster, J. verreauxi. Larvae were reared in receptacles of 150 ml capacity filled with sea water and submitted to two feeding regimens: larvae were submitted to an initial period of starvation and larvae were submitted to an initial period of feeding. Larvae of H. americanus were cultured individually at 17 - 18°C, while phyllosomas were cultured at five larvae per receptacle at temperatures of 19 - 22°C and 16 - 17° C for J. edwardsii and J. verreauxi, respectively. No larvae succeeded in molting if completely starved or if they were fed after a prolonged starvation period. However, the species showed a period of tolerance before food was introduced. The average interval between the first day of feeding and the first day of molting was relatively constant within each species: about 4, 10 and 12 d for H. americanus, J. edwardsii and J. verreauxi, respectively. The starvation tolerance period (50% survival) averaged about 5, 4 and 8 d for these species, respectively. The interval between the ending of starvation and the initial molting period were roughly equivalent although it was shorter in H. americanus than in the Jasus species. The beginning of molting in each species was delayed in accordance with prolonged days of starvation. The feeding period that allowed at least 50% of the larvae to molt to the 2nd instar was 1, 5 and 7 d for these species, respec- tively. Molting in each species began after a rather constant intermolt period (5, 12, 13 d, respectively), regardless of the length of the initial feeding period. These results indicate that the first instar phyllosoma of these Jasus spp. are less tolerant of starvation and require a longer feeding period to molt compared to Homarus larvae. Decapod larvae usually hatch as zoea and begin to take plant or animal food. Food supply is considered critical for the successful culturing of such larvae. Food should be provided before the beginning of the feeding stage and the timing of the first feeding has to be syn- chronized just after hatching for Panulirus japonicus (Mikami and Takashima, 1993) and for Homarus americanus (Aiken and Waddy, 1995) or after metamorphosis from nauplius to first zoea stage of penaeid shrimp (Hudinaga, 1942; Shigueno, 1975) and naupliosoma to phyllosoma instar (e.g., genus Jasus spp. Kittaka, 1988; Kittaka et al., 1988). Recently, concepts of the “point-of-no-return” (PNR), which is the maximum period (days) of starvation prior to the first feeding which will allow the instar to molt to the next instar, and “point-of-reserve-saturation” (PRS), which is the minimum feeding time (depending on the species and, or, its rearing conditions, hours or days) during which nutrients are reserved to allow the instar to molt into the next instar, (differing for each larvae instar in a complete development series). The concepts have been introduced into larval culture tech- nology (Anger and Darwirs, 1981; Mikami et al., 1994). Estimation of the PNR is consid- ered to give us more practical information on allowable initial non-feeding time because our 73 74 BULLETIN OF MARINE SCIENCE, 61(1): 73–80, 1997 common practice was to begin feeding immediately after hatching. PRS will also give us the possible time to stop feeding before each molt because feeding after some time of PRS is considered unnecessary. Such information will make it possible to improve the feeding schedule, because food such as Artemia nauplii and mussel flesh induces water quality deterioration (e.g., for phyllosomas of spiny lobster, Radhakrishnan and Vijayakumaran, 1995; Kittaka, 1995), thus helping to maintain water quality at a more satisfactory level and keep larvae in a healthy condition. Thus, starvation is not only an interesting subject for studies on digestive physiology but for critical measures in larval culture. Nevertheless, related studies are very few. The effect of starvation on larvae survival was examined for several decapod crustaceans (spider crab, Hyas arenaeus (Anger and Darwirs, 1981), king crab Paralithodes camtchaticus (Kurata, 1959), tanner crab Chionescetes opilio (Kon, 1979), red frog crab Ranina ranina (Minagawa, 1989). Recently, Mikami and Takashima (1993) and Mikami et al. (1994) studied effect of starvation on phyllosomas of the Japanese spiny lobster P. japonicus and a scyllarid Thenus sp. Anger et al. (1985) studied change in hepatopancreas of starved larvae of American lobster H. americanus in relation to the molting cycle. Phyllosomas are very difficult to culture (Kittaka, 1994). Therefore, knowledge of spe- cific PNR and PRS levels may help to improve method of culture phyllosomas of different species. The present paper reports on starvation effect on two different types of larvae: the first instar of phyllosomas of New Zealand red rock lobster Jasus edwardsii (H. Milne Edwards, 1875), the green rock lobster of eastern Australia and northern New Zealand, J. verreauxi (H. Milne Edwards, 1851), and larvae of the American clawed lobster, H. americanus (H. Milne Edwards, 1837). MATERIALS AND METHODS The first larval instars of H. americanus, J. verreauxi and J. edwardsii were submitted to the follow- ing two contrasting feeding experiments: (1) an initial period of starvation and (2) an initial period of feeding. The larvae were collected after hatching and transferred to a 10-liter aquarium. The apparently healthy larvae were pipetted and transferred into transparent glass containers of 150 ml capacity filled with 100 ml of filtered and sterilized seawater. Each day, larvae in the two experiments were trans- ferred into newly prepared containers, and in accordance with the feeding regimen, they were fed with newly hatched Artemia nauplii at the density of 4 ind. ml-1. The containers were immersed partially into a water-bath with the temperature adjusted to 18 ± 1° C for H. americanus and 19° C (ranging from 19 to 22° C) for J. edwardsii. Phyllosomas of J. verreauxi were exposed to ambient temperature conditions in the laboratory, approximately 16 - 17° C. The photoperiod has an effect on larval development and survival of H. americanus for which the use of near normal light-dark day-lengths (LD 10:14 to LD 14:10) are recommended (Aiken and Waddy for H. americanus, 1995). For phyllosomas, the effect of light and/or day length under labora- tory conditions was not tested experimentally, but they showed a repetitive rotatory movement when exposed to light reflected in the culture receptacles. This behavior ceased in complete darkness. Thus, the larvae of H. americanus were exposed to laboratory day/night photoperiods while the phyllosomas were reared in darkness. The PNR50 and PRS50 were estimated using the correlation between the number of larvae molting (at both treatments) and the initial feeding period or starvation period taken at each monitored group (Figs. 1,3). Experiments with H. americanus and J. edwardsii were carried out at Sanriku while those with J. verreauxi were done at Nemuro, Hokkaido, Japan. ABRUNHOSA AND KITTAKA: STARVATION IN CRUSTACEAN LARVAE 75 LA RVA E OF H. AMERICANUS. In February of 1994 ovigerous females of the American lobster were placed in a tank with running seawater and constant aeration (salinity 35‰, temperature 14 - 18° C). They were fed mussels, Mytilus edulis, every 2 d. After hatching, the larvae were organized into several groups (6 - 10 individual/group) in accordance with the starvation treatment. The larvae were placed individually in the receptacles because cannibalism in H. americanus is related to their feeding behavior (Atema and Cobb, 1980). Experiment 1 - Larvae of H. americanus submitted to an initial period of starvation. Eight groups were formed with 10 larvae/group (10 replicates), in which they were starved for 0 (i.e., they were not starved) and 1, 2, 3, 4, 6, 8 and 10 d after hatching, except those groups starved 6, 8 and 10 d held 7, 6 and 6 larvae/group, respectively. After the starvation period larvae were fed Artemia nauplii. Experiment 2 - Larvae of H. americanus submitted to an initial period of feeding. Five groups were formed with 10 larvae/group (10 replicates), in which larvae were fed Artemia nauplii for 0 ( i.e., they were not fed) and 1, 2, 3 and 4 d, etc., before being starved, except those groups fed for 0 and 4 d held 50 and 8 individuals/group (i.e., 50 and 8 replicates, respectively). PHYLLOSOMAS OF J. VERREAUXI AND J. EDWARDSII. After mating and oviposition in the laboratory, egg- bearing females were placed in a tank with running seawater and aeration (salinity 35‰, temperature 15 - 18° C). They were fed mussels every 2 d. Larvae of J. edwardsii that hatched out in March of 1994, and those of J. verreauxi in August of 1995 were used for the experiments. Unlike H. americanus, Jasus phyllosomas are not cannibalistic (Kittaka, 1994). Thus, these larvae were placed in transparent glass containers communally at 5 phyllosomas per container (4 and 5 replicates per group for J. edwardsii and J. verreauxi, respectively). Experiment 1- Phyllosomas submitted to an initial period of starvation. For J. edwardsii, ten groups were formed with 4 replicates (5 larvae/receptacle) in which larvae were starved for 0, 1, 2, ..., 10 d after hatching.
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