Effects of temperature on photosynthesis and respiration in hermatypic corals

Photosynthesis and respiration rates of the reef corals Pocillopora damicornis (Linn.), Montipora verrucosa (Lamarck), Porites compressa Dana and Fungia scutaria Lamarck were measured under controlled temperatures. Results indicate that coral metabolism is closely adapted to ambient temperature conditions. Tropical corals measured at Enewetak, Marshall Islands, showed greater primary production compared to maintenance requirements at elevated temperatures than did subtropical varieties of the same species in Hawaii. Photosynthesis: respiration (P:R) ratios were significantly and negatively related with temperature between 18° and 31°C for all Hawaiian corals, whereas at Enewetak this ratio generally showed a curvilinear relationship for this temperature range. Extrapolations of P:R regressions on temperatures to a value of 2.0 (estimated as a minimum required for long-term functional autotrophy) coincide for Hawaiian specimens with published upper lethal temperatures. Extrapolation of P:R regressions for Enewetak specimens at temperatures above 25°C suggests lethal temperatures for these corals to be 2 to 5 C° higher than for Hawaiian corals, in good agreement with recent experimental findings. Interspecific differences in P:R temperature regressions for Hawaiian corals correlating with upper lethal temperature tolerances are described.Contribution No. 505 of the Hawaii Institute of Marine Biology.     

Synergistic effects of temperature,salinity and light on the hermatypic coral Montipora verrucosa

Temperature tolerance in the reef coral Montipora verrucosa (Lamarck) is affected by salinity and light. Low salinity reduces ability of the coral to survive shortterm exposure to elevated temperature. High natural light intensity aggravates damage sustained by corals at high temperature. In long-term growth experiments, high light intensity caused substantial loss of zooxanthellar pigment, higher mortality rates, reduced carbon fixation and lowered growth rate at both upper and lower sublethal temperatures Effects of light at optimal temperature were less dramatic. Interactions between physical environmental factors appear to be most important near the limits of tolerance for a given factor. Acclimation capability was indicated, and was influenced by both thermal history and pigmentation state of stressed corals.Contribution No. 543 of the Hawaii Institute of Marine Biology.     

Depth and temperature of the blue marlin,Makaira nigricans,observed by acoustic telemetry

Multiplex acoustic transmitters were used to monitor the depth, swimming speeds, body temperature and water temperature preference of six blue marlin, Makaira nigricans (Lacépède), near the Hawaiian islands in July and August 1989. The blue marlin ranged in size from 60 to 220 kg and were tracked for 1 to 5 d. All of the fish moved away from the point of capture and were followed up to 253 km from the island of Hawaii. The blue marlin tracked remained in the top 200 m of the water column, spending half the time in the upper 10 m, and rarely ventured below the thermocline. In the nearsurface waters the temperature was uniformly warm (25 to 27°C). The coldest water temperature, 17°C, was encountered on the deepest descent recorded (209 m). Depth changes occurred rapidly and excursions below 10 m were usually less than 60 min in duration. Muscle temperature was similar to water temperature except for a 2°C elevation in muscle temperature observed at the beginning of tracking one individual. This initial rise in body temperature was associated with the anaerobic muscle activity during capture and is an indication of the physiological stress involved in capture.     

Critical thermal maxima of two species of estuarine fish

Critical thermal maxima (CTM) were determined for winter flounder (Pseudopleuronectes americanus, Walbaum) and scup (Stenotomus chrysops, L.), acclimated to ambient water temperature or constant increments above ambient. The acclimation regimes simulated a natural environment by tracking the temperature fluctuations in Point Judith Pond, a small Rhode Island estuary. An elevation in CTM from 30.2° to 35.6°C was observed for scup as the mean acclimation temperature increased from 14.8° to 22.2°C. CTM for flounder increased from 26.4° to 32.4°C over an acclimation range of 4.2° to 23.0°C. The use of CTM studies for predicting the effects of heated effluents is discussed.     

Skeletal Sr content and density in Porites spp. in relation to environmental factors

The coral genus Porites was investigated to evaluate the use of skeletal strontium content as a recorder of seasonal and annual temperature oscillations. In the Hawaiian archipelago, the mean annual water temperature fluctuates by ±0.5C°, with seasonal temperature ranges of 4 to 8C°; the resolution of the Sr thermometer appears to be ±1.5C°. Of this error term, ±0.7C° is analytical, the remainder is biological. Corals from some locations yield temperatures which are consistently offset from the Sr vs temperature calibration line, suggesting genetic population differences. Analysis of cores collected in 1980 from Hawaiian Porites spp. showed no discernible long-term trends over a 100 yr period. Although absolute temperatures are poorly resolved, subannual oscillations in skeletal Sr values accurately reflect recorded seasonal temperature variations. The most useful application of the Sr thermometer is in deciphering the skeletal density band pattern. Subannual oscillations in Sr-temperature values when correlated with density values showed a consistent pattern. When the sections chosen for x-radiography closely followed the growth axis, an abrupt shift from minimum to maximum skeletal density was evident in September/October each year, followed by a gradual decrease in density. The density pattern, shown by microdensitometry, is independent of latitude or temperature range over the Hawaiian archipelago. The annual density shift coincides with high but declining water temperature and solar insolation. If low-density growth represents optimum calcification conditions, the density shift in Hawaiian Porites spp. reflects a change in conditions from optimal to suboptimal. Analyses of samples from other Indo-Pacific locations confirm the generality of this density pattern and suggest a complex relationship between density and environmental light and temperature.Hawaii Institute of Geophysics Contribution No. 1209; Hawaii Institute of Marine Biology Contribution No. 618     

Influence of thermal history on the response of <Emphasis Type="Italic">Montastraea annularis</Emphasis> to short-term temperature exposure

Acclimation of reef corals to environmental conditions has been related to metabolic response at large geographic scales, but regional relationships have rarely been described. Physiological responses to temperature increases of Montastraea annularis (Ellis and Solander 1786) from an inner lagoon and an outer barrier reef in the Gulf of Honduras, southern Belize, were compared in May 2003. The hypothesis that inferred differences in thermal history would result in contrasting responses to elevated temperature was tested. Ambient seawater temperatures adjacent to corals at 4–5 m depth were measured every 15 min at inner lagoon and outer barrier reef collection sites for 1 year (June 2002–May 2003). Monthly averages and 3-day running averages (warmest period, July–October 2002) of daily maximum seawater temperatures were significantly higher (by ∼0.5°C) at inner lagoon reef compared to outer barrier reef sites. M. annularis photosynthesis (P) and respiration (R) rates were measured in respirometers at six temperatures between 29°C and 35°C approximately every hour, with repeated measurements over 3 h. P and R were significantly lower across most temperature treatments for samples collected from the inner lagoon compared to outer barrier reef. Both inner and outer reef M. annularis displayed an increase in P and R with increasing temperature between 29°C and 32°C, but above 32°C P and R sharply declined. P/R ratio versus temperature showed a significant difference between the elevations of the regression lines suggesting that M. annularis from the outer barrier reefs may have been more physiologically stressed than those from the inner lagoon reefs when exposed to acute temperature changes. These results emphasize that thermal stress must be considered within the context of acclimation temperature, and that short-term exposures may have physiologically important effects on this species.     

Effect of acclimatization to low temperature and reduced light on the response of reef corals to elevated temperature

This study tested the effects of acclimatization on the response of corals to elevated temperature, using juvenile massive Porites spp. and branching P. irregularis from Moorea (W149°50′, S17°30′). During April and May 2006, corals were acclimatized for 15 days to cool (25.7°C) or ambient (27.7°C) temperature, under shaded (352 μmol photons m⁻² s⁻¹) or ambient (554 μmol photons m⁻² s⁻¹) natural light, and then incubated for 7 days at ambient or high temperature (31.1°C), under ambient light (659 μmol photons m⁻² s⁻¹). The response to acclimatization was assessed as biomass, maximum dark-adapted quantum yield of PSII (F _v/F _m), and growth, and the effect of the subsequent treatment was assessed as F _v/F _m and growth. Relative to the controls (i.e., ambient temperature/ambient light), massive Porites spp. responded to acclimatization through increases in biomass under ambient temperature/shade, and low temperature/ambient light, whereas P. irregularis responded through reduced growth under ambient temperature/shade, and low temperature/ambient light. Acclimatization affected the response to thermal stress for massive Porites spp. (but not P. irregularis), with an interaction between the acclimatization and subsequent treatments for growth. This interaction resulted from a lessening of the negative effects of high temperature after acclimatizing to ambient temperature/shade, but an accentuation of the effect after acclimatizing to low temperature/shade. It is possible that changes in biomass for massive Porites spp. are important in modulating the response to high temperature, with the taxonomic variation in this effect potentially resulting from differences in morphology. These results demonstrate that corals can acclimatize during short exposures to downward excursions in temperature and light, which subsequently affects their response to thermal stress. Moreover, even con-generic taxa differ in this capacity, which could affect coral community structure. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effects of elevated temperature on larval settlement and post-settlement survival in scleractinian corals, <Emphasis Type="Italic">Acropora solitaryensis</Emphasis> and <Emphasis Type="Italic">Favites chinensis</Emphasis>

We examined the effects of elevated temperature under different exposure periods on larval settlement and post-settlement survival in scleractinian corals, Acropora solitaryensis and Favites chinensis. In the first experiment with the subtropical coral, A. solitaryensis, the numbers of larvae settling and those dead were examined daily for 5 days at 20, 23 (ambient), 26 and 29°C conditions. Larval settlement of A. solitaryensis was initially greater at higher temperature conditions, but the peak in number of settled larvae shifted from 29 to 26°C by day 5, due to ca. 90% post-settlement mortality at 29°C condition. In order to determine the effects under short-term exposure, larvae of F. chinensis were exposed to 27 (ambient), 31 or 34°C only for one hour in the second experiment. The number of larvae settling for 24 h after the exposure and their survivorship over subsequent week was monitored in the ambient temperature condition. Larvae of F. chinensis exhibited greater settlement at higher temperature treatments and constantly low post-settlement mortalities (< ca. 17%) in all temperature treatments, resulting in the highest number of settled larvae at 34°C treatment. These results suggested two different effects of elevated temperature on the early stages of recruitment process of scleractinian corals; (1) the positive effect on larval settlement and (2) the negative effect on post-settlement survival under prolonged exposure.     

Effects of delayed feeding and temperature on the age of irreversible starvation and on the rates of growth and mortality of Pacific herring larvae

The time periods from exhausion of the yolk to the age of irreversible starvation for Pacific herring Clupea harengus pallasi larvae were 8.5, 7.0 and 6.0 d at 6°, 8° and 10°C, respectively. These periods are within the range perviously measured for Atlantic herring larvae and other temperature zone fish species; they are long compared to the periods for tropical species. The variation in the length of this period is due almost entirely to temperature; the natural logarithm of the time period from fertilization to irreversible starvation is highly correlated (r=0.91) with the mean rearing temperature for 25 species of pelagic marine fish larvae. The rates of growth and mortality, measured for 26 experimental populations of Pacific herring larvae reared at 6°, 8° and 10°C and ten ages of delayed first feeding, decreased and increased, respectively with increasing age of first feeding and increasing temperature. These rates, adjusted for the effects of rearing conditions, were compared with the rates for natural populations of herring larvae. Growth is generally faster in the sea than in experimental enclosures. Two of the eleven estimates of natural mortality rate were high enough to indicate possible catastrophic mass starvation. This is consistent with Hjort's critical period concept of year class formation and it suggests that mass starvation occurs in 18 to 36% of the natural populations of first feeding herring larvae.     

Delayed growth of mussel (Mytilus edulis) and scallop (Pecten maximus) veligers at low temperatures

Veliger larvae of Mytilus edulis (L.) from Menai Straits, North Wales, were maintained for up to 2 mo during 1981 at 5°C and then grown on to metamorphosis at 17°C. Larvae so treated showed similar low mortality and equivalent spat production to control larvae. Growth rate at 17°C was less in treated larvae than in controls, but treated larvae grew a little during the period at low temperature. Larvae of Pecten maximus (L.) from the Irish Sea suffered high mortality at low temperature but larvae surviving 2 wk at 8°C could be grown on to spat at 17°C. The longevity of M. edulis larvae is discussed in relation to the genetic homogeneity of adult populations around the UK.     

Embryogenesis and larval biology of the ahermatypic scleractinian<Emphasis Type="Italic"> Oculina varicosa</Emphasis>

The ivory tree coral Oculina varicosa (Leseur, 1820) is an ahermatypic branching scleractinian that colonizes limestone ledges at depths of 6–100 m along the Atlantic coast of Florida. This paper describes the development of embryos and larvae from shallow-water O. varicosa, collected at 6–8 m depth in July 1999 off Fort Pierce, Florida (27°32.542 N; 79°58.732 W). The effect of temperature on embryogenesis, larval survival, and larval swimming speed were examined in the laboratory. Ontogenetic changes in geotaxis and phototaxis were also investigated. Embryos developed via spiral cleavage from small (100 µm), negatively buoyant eggs. Ciliated larvae developed after 6–9 h at 25°C. Embryogenesis ceased at 10°C, was inhibited at 17°C, and progressed normally at 25°C and 30°C. Larval survival, however, was high across the full range of experimental temperatures (11–31°C), although mortality increased in the warmest treatments (26°C and 31°C). Larval swimming speed was highest at 25°C, and lower at the temperature extremes (5°C and 35°C). An ontogenetic change in geotaxis was observed; newly ciliated larvae swam to the water surface and remained there for approximately 18 h, after which they swam briefly throughout the water column, then became demersal. Early larvae showed no response to light stimulation, but at 14 and 23 days larvae appeared to exhibit negatively phototactic behavior. Although low temperatures inhibited the development of O. varicosa embryos, the larvae survived temperature extremes for extended periods of time. Ontogenetic changes in larval behavior may ensure that competent larvae are close to the benthos to facilitate settlement. Previous experiments on survival, swimming speeds, and observations on behavior of O. varicosa larvae from deep-water adults indicate that there is no difference between larvae of the deep and shallow populations.Communicated by J.P. Grassle, New Brunswick     

Do corals select zooxanthellae by alternative discharge?

Loss of zooxanthellae (dinoflagellate Symbiodinium) from corals will sometimes lead to mass mortality of corals. To detect and quantify Symbiodinium released from corals, we developed a zooxanthellae “trap” and a quantitative PCR (qPCR) system with Symbiodinium clades A–F-specific primer sets. The trap was attached to a branch or the surface of several wild stony corals, and the water samples within the traps, including released Symbiodinium, were subjected to qPCR. All tested corals released clade C Symbiodinium at estimates of ~5,900 cells h⁻¹ cm⁻² of coral surface. Although all tested Pocillopora eydouxi harboured both clades C and D, some of these colonies released only clade C or released a lesser amount of clade D than that in the tissues. Our Symbiodinium quantification system revealed that wild hermatypic corals constantly release Symbiodinium to the environment. Our result suggests that some corals may discharge certain clades of Symbiodinium alternatively.     

Effects of temperature on fertilization and early cleavage of some tropical echinoderms,with emphasis on Echinometra mathaei

Select temperatures, above normal, are shown to reduce success of fertilization and normal early cleavage in the laboratory for the echinoderms Acanthaster planci (L.), Culcita novaeguineae Muller and Troschel, Linckia laevigata (L.), Echinometra mathaei (de Blainville), and Diadema savignyi Michelin. The data indicate that cleavage is more sensitive to increased temperature than is fertilization. Upper tolerance limits for early cleavage in most of the species examined is near 34.0°C. The early developmental stages of A. planci were the most sensitive to elevated temperature, and those of E. mathaei, the least sensitive. Further experiments with E. mathaei showed that unfertilized ova were still viable, dividing normally when fertilized after 2 h exposure at 36.0°C. The ova were significantly less viable after 3 h. Early cleavage stages of E. mathaei were resistant to 36.0°C for exposure times of up to 40 min, but were inhibited beyond this period. It is suggested that the ability of E. mathaei to develop normally at 34.0°C (6C° above ambient temperature) and to withstand limited exposure to 36.0°C may account for the wide distribution of this species in habitats which are often subjected to broad temperature fluctuations, such as reef flats.Contribution No. 46 from the University of Guam Marine Laboratory.     

Elevated temperature impairs onset of symbiosis and reduces survivorship in larvae of the Hawaiian coral, <Emphasis Type="Italic">Fungia scutaria</Emphasis>

Many corals obtain their obligate intracellular dinoflagellate symbionts from the environment as larvae or juveniles. The process of symbiont acquisition remains largely unexplored, especially under stress. This study addressed both the ability of Fungia scutaria (Lamarck 1801) larvae to establish symbiosis with Symbiodinium sp. C1f while exposed to elevated temperature and the survivorship of aposymbiotic and newly symbiotic larvae under these conditions. Larvae were exposed to 27, 29, or 31°C for 1 h prior to infection, throughout a 3-h infection period, and up to 72 h following infection. Exposure to elevated temperatures impaired the ability of coral larvae to establish symbiosis and reduced larval survivorship. At 31°C, the presence of symbionts further reduced larval survivorship. As sea surface temperatures rise, coral larvae exposed to elevated temperatures during symbiosis onset will likely be negatively impacted, which in turn could affect the establishment of future generations of corals.     

Studies on differential fitness of PGI genotypes with regard to temperature in Gammarus insensibilis (Crustacea: Amphipoda)

This paper deals with an experimental study of survival, combined with estimates of biochemical activity, of different genotypes at the PGI (phosphoglucose isomerase) locus in Gammarus insensibilis, in relation to temperature. Samples were collected in the lagoon of Venice during 1987. No mortality occurred at 10°C whereas at 27°C, where mortality reached the value of 50%, heterozygotes exhibited significantly higher survival than homozygotes. Experiments conducted in order to evaluate the PGI biochemical activity of homo- and heterozygote genotypes at three different temperatures (4°, 20° and 37°C) showed in all genotypes an increased activity from 4° to 20°C and a fall of activity from 20° to 37°C. Heterozygotes exhibited higher activity at all temperatures tested. The difference between homo- and heterozygotes became more obvious at 37°C. Our results suggest that in G. insensibilis the PGI locus, as already shown in other organisms, may be subject to selection and that the heterozygous genotypes possess superior fitness. The biochemical bases of the observed differences between genotypes are briefly discussed.     

Temperature ecotypes near the southern boundary of the kelp Laminaria saccharina

Effects of temperature on survival, growth, and photosynthesis were compared for two USA populations of Laminaria saccharina Lamour. One population was located in New York State, near the southern latitudinal boundary of the species in the western North Atlantic. This southern boundary population was exposed to ambient temperatures 20°C for about 6 wk each summer. The second population was located in Maine, toward the center of the latitudinal range of the species, and was rarely exposed to temperatures>17°C. sporophytes from the New York (NY) population exhibited greater tolerance of high temperature than plants from the Maine (ME) site. Juvenile sporophytes from the two sites had similar rates of survivorship and growth at temperatures below 20°C, but showed different responses at 20°C in laboratory experiments. NY plants survived and grew for 6 wk at 20°C. ME plants showed negative growth during wk 2 and 100% mortality during wk 3. NY and ME plants held in situ at the NY site during June to September, 1985, also exhibited differential survivorship when ambient temperatures exceeded 20°C. Results of photosynthesis and dark respiration measurements on NY and ME plants grown at various temperatures suggested that the high-temperature tolerance of NY plants was attributable to their ability to maintain positive daily net C-fixation at 20°C. The high-temperature tolerance of the NY plants appeared to be due to genetic adaptation and is probably crucial to the persistence of the species near its southern boundary.     

Population dynamics and production of the planktonic copepods in a eutrophic inlet of the Inland Sea of Japan. I. Centropages abdominalis

Population dynamics and production of the calanoid copepod Centropages abdominalis were studied from November 1986 to November 1987 in Fukuyama Harbor, in the central part of the Inland Sea of Japan. This species was present in the plankton during a cold-water period from November to June (temperature range: 8.9 to 21.1 °C), with a peak abundance (23 600 ind m^–3) in February. During this period, six generations could be detected, and each generation time agreed well with that predicted from food-satiated laboratory experiments, indicating that the natural population was not food-limited. The population suffered extremely high mortality during the period from egg to naupliar stage (N) II: only 0.02 to 4% of the eggs survived to NII. However, the mortality in stages older than NII was almost negligible. The growth rate of C. abdominalis increased exponentially with increasing temperature. Its biomass and production rate showed marked seasonal variations largely in parallel with numerical abundance. The estimated production between 7 November 1986 and 29 May 1987 was 355 mg C m^–3 or 2.66 g C m^–2, 95% of which occurred during February and March. The daily production rate to biomass ratio increased exponentially with temperature from 0.18 at 8.9°C to 0.37 at 19°C.     

Rearing Pandalus borealis larvae in the laboratory

Larvae of the northern shrimp Pandalus borealis (Krøyer) are pelagic. In the Estuary and Gulf of St. Lawrence, Canada, the early stages are found in the upper 25-m of the water column in spring and early summer and are expected to experience a range of water temperatures from as low as 0°C to as high at 6°C. Little is known of the impact of water temperature on metabolic requirements of northern shrimp larvae. In this study, routine respiration (VO₂), maximum respiration (electron transport system activity, ETSA) and metabolic scope for growth (MS, ETSA–VO₂) of northern shrimp larvae were measured as a function of temperature (3, 5 and 8°C), developmental stage (I–V at 3°C, I–VII at 5°C and 8°C) and growth rate in dry mass. After logarithmic transformation, all three metabolic variables were linearly related to dry mass. The increase in VO₂ with body mass was faster at 5°C than at 3 or 8°C, whereas with ETSA this increase was slower. As a result, MS increased more slowly with dry mass at 5°C than at 3 and 8°C. However, MS did not limit growth in this study, since it explained only 39% of the variability in growth. All three metabolic variables as well as growth varied together as a function of temperature and ontogeny. Q₁₀ of all three metabolic variables ranged from 1.6 and 2.2 for stages I–V larvae, except for VO₂ at stage I (3.9) and stage III (2.9).     

Temperature tolerance of the polychaete worms Diopatra cuprea and Clymenella torquata

The resistance to high temperatures of two species of intertidal tube-dwelling polychaete worms has been tested to show seasonal and geographic variations. The summer 50% survival temperature at Beaufort, North Carolina, was 42.5 °C for Diopatra cuprea (Bosc) and 40.5 °C for Clymenella torquata (Leidy). Winter 50% survival temperatures for both species at Beaufort were approximately 4.0 C° lower. Both species showed a geographic difference in 50% survival temperature of more than 4.0 C° between North Carolina and Massachusetts in summer. D. cuprea from the Mississippi coast showed a lower survival temperature, probably due to combined temperature and salinity effects. Laboratory acclimation of C. torquata from Beaufort at low temperatures, during summer months, produced experimental results similar to those from winter animals. The seasonal differences in temperature tolerance are phenotypic expressions of a physiological response which can be related to environmental temperature patterns.     

A dispersal model for larvae of the deep sea red crab Geryon quinquedens based upon behavioral regulation of vertical migration in the hatching stage

Behavioral responses to gravity, hydrostatic pressure, and thermoclines are described for Stage I zoeae of the deep sea red crab Geryon quinquedens Smith. Survival and rate of development as a function of temperature is presented for all larval stages. Although temperatures between 10° and 25°C have no direct effect upon survival, development time is five times longer at 10°C than at 25°C. Stage I larvae show strong negative response to gravity. Swimming rate increases with an increase in pressure up to 20 atm above ambient at 11°C, but not at 15°C. Swimming rates at 15°C are higher than those measured at 11°C at each pressure tested. Stage I larvae readily penetrate sharp thermoclines. Potential dispersal ranges of G. quinquedens larvae in the Mid-Atlantic Bight are suggested based on larval behavior, development time, and coastal hydrography. A testable recruitment model is proposed for G. quinquedens.Contribution no. 1365 of the Center for Environmental and Estuarine Studies