A REVIEW OF LARVAL DEVELOPMENT IN THE INTERTIDAL LIMPET GENUS SIPHONARIA (GASTROPODA: PULMONATA)

A compilation of distributional and life-history data relatingto mode of larval development is presented for 26 species ofSiphonana, a genus of intertidal pulmonates. Most species depositgelatinous benthic egg masses with two species releasing pelagicegg masses. Fifteen species hatch as planktonic-developing larvae,nine hatch as direct-developing juveniles, and in a furthertwo larvae hatch with both the swimming velar apparatus (associatedwith plank-tonic development) and a crawling foot (associatedwith direct development). Data on mode of larval developmentare interpreted with respect to some adaptive models. Despiteimportant exceptions, there is support for adaptive models basedupon egg capsule size (direct developers hatch from larger eggcapsules) and intertidal distribution (direct developers generallyoccur higher on the shore than planktonic developers). Worldwide,planktonic developers are more widespread than direct-developingspecies, and individual planktonic species have a greater meanlatitudinal range. The evidence for adaptive models relatinglatitudinal distribution to developmental mode is equivocal.There appears to be no clear relationship between body sizeand developmental mode in the genus, although the smallest specieshas direct development and the largest has planktonic development.In most siphonariid subgenera, developmental mode appears tobe constant, but two subgenera contain a mixture of developmentaltypes (Received 1 November 1993; accepted 15 April 1994)     

NOTES ON THE TAXONOMY, SPAWN AND LARVAL DEVELOPMENT OF SOUTH AFRICAN SPECIES OF THE INTERTIDAL LIMPET SIPHONARIA (GASTROPODA: PULMONATA)

The taxonomy of South African Siphonaria is reviewed. We concludethat nine species are valid. These are: S. anneae Tomlin, 1944;S. capensis Quoy & Gaimard, 1833; S. compressa Allanson,1959; S. concinna Sowerby, 1824; S. dayi Allanson, 1959; S.nigerrima Smith, 1903; S. oculus Krauss, 1848; S. serrata (Fischer,1807) and S. tenuicostulata Smith, 1903. Of these, Siphonarianigerrima Smith, 1903, has been incorrectly synonymized withSiphonaria carbo Hanley, 1858, which is not present on SouthAfrican shores. S. aspera Krauss, 1848 is reduced to a juniorsynonym of S. Serrata (Fischer, 1807). Shell characteristicsand mode of larval development are described for these nineSouth African species. All species deposit benthic egg massesand development may be either planktonic with swimming veligerlarvae (S. capensis, S. concinna and S. oculus) or direct, withcrawling larvae emerging from the eggs (S. anneae, S. compressa,S. dayi, S. nigerrima, S. serrata and S. tenuicostulata). (Received 20 November 1992; accepted 24 January 1994)     

TEMPORAL VARIATION IN FORAGING BEHAVIOUR OF PATELLA GRANULARIS (PATELLOGASTROPODA) AND SIPHONARIA CONCINNA (BASOMMATOPHORA) ON A SOUTH AFRICAN SHORE

Foraging activity of two mid- to low- shore species of limpet,Patella granulans (Prosobranchia) and Siphonaria concinna (Pulmonata)from an exposed shore on the Eastern Cape coast of South Africawas monitored. In both species, activity was compared duringspring and neap tides and, in P. granularis between summer andwinter. Rhythms of activity of the two species were similar,with foraging excursions being mainly associated with nocturnallow tide times, although some P. granularis foraged during daytimelow tides. It is suggested that foraging excursions in P. granularisare triggered by wave action. Both species foraged further onspring tides than on neap tides and this is suggested to bea result of the limited time limpets have to forage. P. granulariswas also found to forage further during summer when comparedto winter and the possibility that seasonal micro-algal productivityinfluences foraging distances in limpets is discussed. The foraging activity of both species could be divided into3 distinct phases, a relatively rapid outward phase, a muchslower foraging phase and a rapid homeward phase. Whether ornot these limpets graze throughout an excursion is not known.S. concinna was found to home to a fixed scar, although duringthe experiment some scar-swapping occurred. P. granularis didnot home to a fixed scar but possessed a ‘home range’(approx. 5 cm²) to which it returned after each excursion. Patella granularis was found to move randomly during foraging,whilst S. concinna foraged in a non-random direction -whichtook individuals upshore. No tidal-influence is thought to bepresent in this case and the possibility of a learning componentin the foraging behaviour of certain limpet species in relationto the return to optimal feeding patches is discussed. (Received 18 May 1996; accepted 2 September 1996)     

THE STRUCTURE OF THE SECONDARY GILLS OF SIPHONARIA CAPENSIS (GASTROPODA: PULMONATA)

The secondary gill of the pulmonate limpet Siphonaria capensisis located in the dorsal portion of the mantle cavity. Eachlamella of the gill is triangular in shape and bears ciliarytufts which have a density of about 400/mm². The free ends ofsome of the cilia are enlarged biconcave discs. Each gill lamellais covered by a single layer of cuboidal (4x4 µm) epitheliumon each side, separated by a central haemocoelic space. Ciliatedand mucus secreting cells are interspersed amongst the epithelialcells. The haemocoelic space is spanned at intervals by trabeculaecontaining longitudinal and transverse muscle fibres. (Received 29 August 1986;     

高原鼢鼠挖掘取食活动的能量代价及其最佳挖掘取食行为

高原鼢鼠(Myospalax baileyi)是青藏高原典型的地下鼠,它主要挖掘取食杂类草之地下根茎,在挖掘过程中形成取食洞道。本文根据此特点,建立了挖掘取食过程中能量代价(E_(?))与取食道结构参数(土丘间距S、侧道长L、取食道半径b及洞道深度D)之间以及取食获能E_(fg)与S和b之间的两个函数关系。分别以挖掘效率极大和取食效率极大为优化目标,导出同样的结果,即相同的最佳土丘间距S_(opt)。通过实验观测,计算出在矮嵩草(Kobresiahumilis)草甸生境条件下,最佳土丘间距S_(opt)=127.4厘米,与野外实测土丘间距(126.30±47.85厘米)无显著差异(t=0.312,P>0.50),表明高原鼢鼠的挖掘及取食行为符合最佳化原则。     

CIRCADIAN ACTIVITY RHYTHMS AND SENSITIVITY TO NOISE IN THE MONGOLIAN GERBIL (MERIONES UNGUICULATUS)

Since consistent data on endogenous circadian rhythms of Mongolian gerbils are not available, the main aim of our study was to identify suitable conditions to receive stable and reproducible free-running rhythms of activity under different light intensities. Another objective was to determine the role of social cues as an exogenous zeitgeber in the absence of a light-dark (LD) cycle. We performed two long-term sets of experiments with adult male gerbils kept in climatic chambers under various photoperiods of at least 30 days each. In all cases, the time of lights on in the chambers differed from the daily starting hour of work in the animal house. Always, two animals per chamber were kept separately in cages with a running wheel while their activity was monitored continuously. During the first set, only three of eight animals developed intra- and interindividual variable free-running rhythms. The activity patterns seemed to be influenced by human activities outside, indicating high sensitivity to external factors. Subsequently, we damped the chambers and the room and restricted access to the room. In the following noise-reduced set, all gerbils developed comparable free-running rhythms of activity. We determined the mean of the free-running period τ, the activity-rest relationship α/θ and the amount of running wheel activity per day: τ = 23.7h ± 0.08h under low light (5 lux) and 25.5h ± 0.19h under high light intensities (450 lux); α/θ = 0.53 ± 0.08 under 5 lux and 0.34 ± 0.04 under 450 lux. The amount of daily activity was 12 times as high under 5 lux as under 450 lux. There was no indication that the two animals in one chamber socially synchronized each other. In conclusion, the pronounced rhythm changes in accordance with Aschoff's theory support the view that gerbils are mainly nocturnal animals. (Chronobiology International, 17(2), 137–145, 2000)     

FORAGING ACTIVITY OF HELCION PRUINOSUS (PATELLOGASTROPODA) ON A SOUTH AFRICAN BOULDER SHORE

Foraging activity of the mid-shore limpet Helcion pruinosuswas monitored on an exposed boulder beach in the Eastern Capeof South Africa. Numbers of limpets active in 10 fixed quadratswere monitored and compared in different seasons and during springand neap tides. Foraging excursions were mainly associated with nocturnallow tides, limpets only emerging from beneath rocks when theywere completely uncovered by the receding tide, and they hadall retreated to beneath rocks before being splashed by theadvancing tide. The number of animals active was greater onfull moon spring tides. Significantly more limpets were activeon an autumn spring tide than on similar tides in other seasons. ^* To whom correspondence should be addressed. (Received 27 March 2000; accepted 5 July 2000)     

ACTIVITY RHYTHMS OF WILD AND LABORATORY GOLDEN HAMSTERS (MESOCRICETUS AURATUS) UNDER ENTRAINED AND FREE-RUNNING CONDITIONS

The golden hamster (Mesocricetus auratus) is one of the most frequently used laboratory animals, particularly in chronobiological studies. One reason is its very robust and predictable rhythms, although the question arises whether this is an inbreeding effect or rather is typical for the species. We compared the daily (circadian) activity rhythms of wild and laboratory golden hamsters. The laboratory hamsters were derived from our own outbred stock (Zoh:GOHA). The wild hamsters included animals captured in Syria and their descendants (F1). Experiments were performed under entrained (light: dark [LD] 14h:10h) and under free-running (constant darkness, DD) conditions. Locomotor activity was recorded using passive infrared detectors. Under entrained conditions, the animals had access to a running wheel for a certain time to induce additional activity. After 3 weeks in constant darkness, a light pulse (15 min, 100 lux) was applied at circadian time 14 (CT14). Both laboratory and wild hamsters showed well-pronounced and very similar activity rhythms. Under entrained conditions, all hamsters manifested about 80% of their total 24h activity during the dark portion of the LD cycle. The robustness of the daily rhythms was also similar. However, interindividual variability was higher in wild hamsters for both measures. All animals used the running wheels almost exclusively during the dark portion of the LD cycle, although the wild hamsters were three times more active. The period length, measured in constant darkness, was significantly shorter in wild (23.93h ± 0.10h) than in laboratory hamsters (24.06 ± 0.07h). The light-induced phase changes were not different (about 1.5h). In summary, these results indicate that the laboratory hamster is not much different from the wild type. (Chronobiology International, 18(6), 921–932, 2001)     

ACTIVITY RHYTHMS OF WILD AND LABORATORY GOLDEN HAMSTERS (MESOCRICETUS AURATUS) UNDER ENTRAINED AND FREE-RUNNING CONDITIONS

The golden hamster (Mesocricetus auratus) is one of the most frequently used laboratory animals, particularly in chronobiological studies. One reason is its very robust and predictable rhythms, although the question arises whether this is an inbreeding effect or rather is typical for the species. We compared the daily (circadian) activity rhythms of wild and laboratory golden hamsters. The laboratory hamsters were derived from our own outbred stock (Zoh:GOHA). The wild hamsters included animals captured in Syria and their descendants (F1). Experiments were performed under entrained (light: dark [LD] 14h:10h) and under free-running (constant darkness, DD) conditions. Locomotor activity was recorded using passive infrared detectors. Under entrained conditions, the animals had access to a running wheel for a certain time to induce additional activity. After 3 weeks in constant darkness, a light pulse (15 min, 100 lux) was applied at circadian time 14 (CT14). Both laboratory and wild hamsters showed well-pronounced and very similar activity rhythms. Under entrained conditions, all hamsters manifested about 80% of their total 24h activity during the dark portion of the LD cycle. The robustness of the daily rhythms was also similar. However, interindividual variability was higher in wild hamsters for both measures. All animals used the running wheels almost exclusively during the dark portion of the LD cycle, although the wild hamsters were three times more active. The period length, measured in constant darkness, was significantly shorter in wild (23.93h ± 0.10h) than in laboratory hamsters (24.06 ± 0.07h). The light-induced phase changes were not different (about 1.5h). In summary, these results indicate that the laboratory hamster is not much different from the wild type. (Chronobiology International, 18(6), 921-932, 2001)     

PRESENCE OF CIRCADIAN RHYTHMS IN THE LOCOMOTOR ACTIVITY OF A CAVE-DWELLING MILLIPEDE GLYPHIULUS CAVERNICOLUS SULU (CAMBALIDAE,SPIROSTREPTIDA)

The locomotor activity of the millipede Glyphiulus cavernicolus (Spirostreptida), which occupies the deeper recesses of a cave, was monitored in light-dark (LD) cycles (12h light and 12h darkness), constant darkness (DD), and constant light (LL) conditions. These millipedes live inside the cave and are apparently never exposed to any periodic factors of the environment such as light-dark, temperature, and humidity cycles. The activity of a considerable fraction of these millipedes was found to show circadian rhythm, which entrained to a 12:12 LD cycle with maximum activity during the dark phase of the LD cycle. Under constant darkness (DD), 56.5% of the millipedes (n = 23) showed circadian rhythms, with average free-running period of 25.7h ± 3.3h (mean ± SD, range 22.3h to 35.0h). The remaining 43.5% of the millipedes, however, did not show any clear-cut rhythm. Under DD conditions following an exposure to LD cycles, 66.7% (n = 9) showed faint circadian rhythm, with average free-running period of 24.0h ± 0.8h (mean ± SD, range 22.9h to 25.2h). Under constant light (LL) conditions, only 2 millipedes of 11 showed free-running rhythms, with average period length of 33.3h ± 1.3h. The results suggest that these cave-dwelling millipedes still possess the capacity to measure time and respond to light and dark situations. (Chronobiology International, 17(6), 757–765, 2000)     

EFFECTS OF LIGHT AND MELATONIN TREATMENT ON BODY TEMPERATURE AND MELATONIN SECRETION DAILY RHYTHMS IN A DIURNAL RODENT,THE FAT SAND RAT

Many mammals display predictable daily rhythmicity in both neuroendocrine function and behavior. The basic rest-activity cycles are usually consistent for a given species and vary from night-active (nocturnal), those mostly active at dawn and dusk (i.e., crepuscular), and to day-active (diurnal) species. A number of daily rhythms are oppositely phased with respect to the light/dark (LD) cycle in diurnal compared with nocturnal mammals, whereas others are equally phased with respect to the LD cycle, regardless of diurnality/nocturnality. Pineal produced melatonin (MLT) perfectly matches this phase-locked feature in that its production and secretion always occurs during the night in both diurnal and nocturnal mammals. As most rodents studied to date in the field of chronobiology are nocturnal, the aim in this study was to evaluate the effect of light manipulations and different photoperiods on a diurnal rodent, the fat sand rat, Psammomys obesus. The authors studied its daily rhythms of body temperature (T_b) and 6-sulphatoxymelatonin (6-SMT) under various photoperiodic regimes and light manipulations (acute and chronic exposures) while maintaining a constant ambient temperature of 30°C?±?1°C. The following protocols were used: (A) Control (CON) conditions 12L:12D; (A1) exposure to one light interference (LI) of CON-acclimated individuals for 30?min, 5?h after lights-off; (A2) short photoperiod (SP) acclimation (8L:16D) for 3 wks; (A3) 3 wks of SP acclimation with chronic LI of 15?min, three times a night at 4-h intervals; (A4) chronic exposure to constant dim blue light (470nm, 30 lux) for 24?h for 3 wks (LL). (B) The response to exogenous MLT administration, provided in drinking water, was measured under the following protocols: (B1) After chronic exposure to SP with LI, MLT was provided once, starting 1?h before the end of photophase; (B2) after a continuous exposure to dim blue light, MLT was provided at 15:00?h for 2?h for 2 wks; (B3) to CON animals, MLT was given intraperitoneally (i.p.) at 14:00?h. The results demonstrate that under CON acclimation, Psammomys obesus has robust T_b and 6-SMT daily rhythms in which the acrophase (peak time) of T_b is during the photophase, whereas that of 6-SMT is during scotophase. LI resulted in an elevation of T_b and a reduction of 6-SMT levels. A significant difference in the response was noted between acute and chronic exposure to LI, particularly in 6-SMT levels, which were lower than CON after LI and higher after chronic LI, implying an acclimation process. Constant exposure to blue light abolished T_b and 6-SMT rhythms in all the animals. MLT administration resumed the T_b daily rhythm in these animals, and had a recovery effect on the chronic LI-exposed animals, resulting in a T_b decrease. Altogether, the authors show in this study the different modifications of T_b rhythms and MLT levels in response to environmental light manipulations. These series of experiments may serve as a basis for establishing P. obesus as an animal model for further studies in chronobiology. (Author correspondence: hagitsc@research.haifa.ac.il)     

A COMPARATIVE STUDY OF THE MORPHOLOGY OF THE PULMONATE SNAIL PSEUDOTACHEA LITTURATA (PFEIFFER) AND OTHER SPECIES OF PSEUDOTACHEA, IBERUS AND CEPAEA

A population of Pseudotachea litturata (Pfeiffer, 1851) fromTarifa (Càdiz, Spain) has been studied. The morphologicalresults are compared with those from P. splendida, Iberus gualtierianus,I. alonensis, I. marmoralus, I. guiraoanus and four speciesof the genus Cepaea using, as an exploratory method, the Wagnerparsimony procedure and 18 characters of the shell, genitalsystem and karyotype have been analysed. According to this methodit seems that the taxonomical position of P. litturata in thegenus Pseudotachea is confirmed, and agrees with the phylogeneticalrelationships in this group of species. The genus Cepaea seemsto be well established, although two species groups can be distinguished:C. nemoralis—C. hortensis and C. syluatica—C. vindobonensis.These differ mainly in chromosome number, diverticulum lengthand degree of shell polymorphism. Although the present resultsdo not allow us to clarify the current taxonomical problemswithin the genus Iberus, the species studied seem to belongto a natural group (Received 15 September 1987; accepted 1 January 1988)     

DIURNAL FLUCTUATION OF NITRATE REDUCTASE ACTIVITY IN THE MARINE RED ALGA GRACILARIA TENUISTIPITATA (RHODOPHYTA)1

The biochemical characteristics and diurnal changes in activity of the enzyme nitrate reductase (NR; EC 1.6.6.1) from the marine red alga Gracilaria tenuistipitata var. liui Zhang et Xia are described. Different assay conditions were tested to determine the stability of NR. The crude extract of G. tenuistipitata has a NR specific activity of 10.2 U.mg⁻¹, which is higher than the NR activities found for other algae, plants, and fungi. This NR is highly active at a slightly alkaline pH and is stable over a wide range of temperature, with an optimal activity at 20° C. The apical portions of the thallus contain 64.9 ± 6.6% of the total NR specific activity. The apparent Michaelis-Menten (K_m) constant found for KNO₃ was 197 μM, and it was 95 μM for NADH. The NR from G. tenuistipitata can be included in the NADH-specific group, because no activity was found when NADPH was used as an electron donor. In extracts of algae grown under either continuously dim light or a light-dark cycle, the activity of NR exhibits a daily rhythm, peaking at the middle of the light phase, when activity is 30-fold higher than during the night phase.     

FAECAL PRODUCTION AND AN ESTIMATE OF FOOD INTAKE IN THE WILD OF THE ANTARCTIC LIMPET NACELLA CONCINNA (STREBEL)

The Antarctic limpet Nacella concinna is common in shallow watersaround the Antarctic Peninsula, and has a mean density of 125individuals/m² at Signy Island (Picken, 1980). In the australsummer 1986/87 ammonia excretion and faecal egestion were measuredas part of a programme to develop a detailed individual energybudget for this species. Ammonia production was measured in 102 individuals ranging from3 to 720 mg dry weight. For a standard limpet of 200 mg dryweight, ammonia excretion was 0.13 µg-at/hr. Comparedwith previous measures of oxygen uptake these data suggest anO:N atomic ratio of between 15 and 25. The relationship betweenammonia excretion and dry weight could be expressed by a powercurve with a weight exponent of 0.82 (SE 0.042). Faecal production was measured in limpets freshly sampled fromthe field, and placed in dean seawater for 4 days. During thistime faecal production decreased (since the limpets were notfeeding), although only data from the first 24 hours were used.Again the relationship between faecal egestion and dry weightcould be expressed by a power curve, although this time theweight exponent was 0.94 (SE 0.101). The ash content of thefaecal strings increased significantly with the size of thelimpet, possibly because larger limpets were ingesting a greaterproportion of substrate during feeding. If this is so, thenthis would also explain the weight exponent dose to 1.0, ratherthan the value of about 0.8 to be expected from metabolic measures(since larger limpets would be passing relatively more faecalmaterial). If assimilation efficiency is known, then food intake in thewild may be estimated from a measurement of faecal egestionand faecal organic content (Clarke et at., 1988). Applying typicalvalues of assimilation efficiency for limpets (40 to 60%) suggestsa daily food intake in Nacella of 2 to 3% body weight per day.This is a quite typical figure for a grazing limpet, and suggeststhere is no resource limitation for Nacella at Signy in summer.     

VARIATIONS IN INFESTATIONS OF SIPHONARIA CAPENSIS AND S. CONCINNA(PULMONATA) BY LARVAL TREMATODES IN SOUTHERN AFRICA

The prevalence of digenean trematode parasites in Siphonariacapensis (Pulmonata) was examined in populations from 24 sites,and in S. concinna from 10 sites along the coast of southernAfrica from 1989 to 1991. Sporocysts containing cercariae werefound in the digestive gland of both species. To date, two typesof cercaria have been isolated from S. capensis and one typefrom S. concinna, these types possibly representing three distinctspecies of trematode. In approximately 30% of parasitized S.capensis and 10% of parasitized S. concinna the digestive glandwas completely destroyed. In both species of Siphonaria therewas considerable temporal and spatial variation in parasiteprevalence. At some locations up to 40% of S. capensis wereinfected whereas at others, no parasites were found. Prevalenceof trema-todes in S. concinna rarely exceeded 12%. At severalsites the prevalence of infestation within the limpets increasedwith increasing host size. (Received 8 June 1992; accepted 6 July 1992)