Thermal tolerance,evaporative water loss,air-water oxygen consumption and zonation of intertidal prosobranchs: a new synthesis

Duration of emergence increases with tidal height on rocky shores therefore, emergence adaptations in intertidal species such as littorine and other prosobranch gastropods have been considered correlated with zonation patterns; temperature tolerance, desiccation resistance and aerial respiration rate all commonly assumed to increase progressively with increasing zonation level. Such direct correlations are rarely observed in nature. Maximal aerial gas exchange occurs in mid-shore, not high shore species. Temperature tolerance and desiccation resistance do not increase directly with shore height. Thus, hypotheses regarding physiological correlates of zonation require revaluation. A new hypothesis is presented that the high tide mark presents a single major physiological barrier on rocky shores. Above it, snails experience prolonged emergence and extensive desiccation; below it, predictable submergence and rehydration with each tidal cycle. Thus, desiccation stress is minimal below the high tide mark and maximal above it. Therefore, species restricted below high tide (the eulittoral zone) should display markedly different adaptive strategies to emergence than those above it (the eulittoral fringe). A review of the literature indicated that adaptations in eulittoral species are dominated by those allowing maintenance of activity and foraging in air including: evaporative cooling; low thermal tolerance; elevated aerial O₂ uptake rates; and high capacity for radiant heat absorption. Such adaptations exacerbate evaporative water loss. In contrast, species restricted to the eulittoral fringe display adaptive strategies that minimize desiccation and prolong survival of emergence including: foot withdrawal, preventing heat conduction from the substratum; aestivation in air; elevated thermal tolerance reducing necessity for evaporative cooling; position maintenance by cementation to the substratum and increased capacity for heat dissipation. In order to test of this hypothesis the upper thermal limits, tissue and substratum temperatures on emergence in direct sunlight and evaporative water loss and tissue temperatures on emergence in 40 °C were evaluated for specimens of six species of eulittoral and eulittoral fringe gastropods from a granite shore on Princess Royal Harbour near Albany, Western Australia. The results were consistant with adaptation to the proposed desiccation barrier at high tide. The eulittoral species, Austrocochlea constricta, Austrocochlea concamerata, Nerita atramentosa and Lepsiella vinosa, displayed adaptations dominated by maintenance of activity and foraging during emergence while the eulittoral fringe littorine species, Bembicium vittatum and Nodilittorina unifasciata displayed adaptations dominated by minization of activity and evaporative water loss during emergence. The evolution of adaptations allowing tolerance of prolonged desiccation have allowed littorine species to dominate high intertidal rocky shore gastropod faunas throughout the world's oceans.     

Tolerance of forced air emergence by a fish with a broad vertical distribution, the rockpool blenny, Hypsoblennius gilberti (Blenniidae)

Many intertidal fishes, particularly among the Blenniidae and Cottidae, possess amphibious adaptations, including the ability to breathe in air and to avoid desiccation in terrestrial conditions. These traits are absent in subtidal species of blennies and cottids. Hypsoblennius gilberti, the rockpool blenny, is found in shallow rockpools in the mid to high intertidal areas of Southern California, and deeper to 18 m in the subtidal zone. This broad vertical distribution could indicate that this blenny is adapted for tidal air emergence, although H. gilberti has not been observed out of water in its natural habitat. H. gilberti does not emerge voluntarily from hypoxic sea water in the laboratory, but it easily withstands 3 h out of water. The aerial respiratory exchange ratio (CO2 released compared to O2 consumed) is 0.70, similar to that of amphibious intertidal fishes in air, indicating sufficient release of metabolically produced CO2 while emerged. There is no increase in aquatic respiration following emergence. However, unlike other amphibious fishes that maintain aerial oxygen consumption at a level similar to aquatic oxygen consumption, H. gilberti has an aerial oxygen consumption rate one-third that in water. H. gilberti can recover rapidly from terrestrial water loss, and shows no change in evaporative water loss rates at 93% and 77% relative humidities. The amphibious capabilities in H. gilberti, even if rarely used, permit survival in air during tidal emergence. These findings suggest that H. gilberti may demonstrate an intermediate condition between the amphibious species of intertidal fishes that regularly emerge from water, and the subtidal fishes that do not survive air emergence and are completely restricted to an aquatic habitat.     

红树植物胎生现象研究进展

植物胎生是指有性繁殖产生的后代在母体上直接萌发的现象, 在红树植物中最为常见。红树植物生长在热带亚热带海岸潮间带, 耐受高盐、高温、淹水缺氧和海浪冲击等复杂环境。胎生被认为是红树植物对这种特殊生境的重要适应方式。该文从形态发育、生理生化、分子水平、生态适应4个层次讨论红树植物胎生现象对复杂生境的适应性, 并指出现有研究存在的不足, 对将来的研究方向进行了展望。与非胎生胚胎发育相比, 红树植物胎生是一个遗传的程序, 在进化过程中形成了一些特殊的结构。植物激素对胎生发育起关键的调控作用, 繁殖体发育过程中, 其盐离子的种类与浓度的动态变化则是对海岸潮间带生境的重要适应特征。这种胎生繁殖体依靠在母体上完善的一系列功能性特征能更有效地适应落地后的滩涂环境。然而, 红树植物胎生发育过程的分子机理及调控机制还有待研究。理解胎生这一特殊适应性现象的本质及其进化过程将为红树林保护繁育、适应气候变化提供理论依据。     

Factors delimiting the boundary between vertically contiguous mussel beds ofSeptifer virgatus (Wiegmann) andHormomya mutabilis (Gould)

On moderately wave-exposed rocky shores in middle Japan, the upper interidal mytilid,Septifer virgatus, and lower intertidal mytilid,Hormomya mutabilis, occur together, forming vertically contiguous mussed beds. Factors limiting the lower distribution limit ofSeptifer and the upper limit ofHormomya were investigated by collections of natural mussel clumps and single- and mixed-species transplantation experiments. Newly settled juvenileSeptifer (<5 mm shell length) were significantly fewer in the natural and artificialHormomya clumps than in theSeptifer clumps. Both natural and artificialHormomya clumps accumulated a much greater amount of sediment than did theSeptifer clumps.Hormomya clumps inhibited the recruitment ofSeptifer, presumably through accumulation of sediment, which resulted in setting the lower limit of theSeptifer zone. Survivorship of small (5–10 mm) and large (>15 mm)Hormomya was much lower inHormomya clumps transplanted upwards into theSeptifer zone than it was inHormomya clumps in theHormomya zone. In mixed-species clumps in theSeptifer zone, however, survivorship of smallHormomya was not significantly different from that in mixed-species clumps in theHormomya zone.Septifer had a positive effect on the survival of smallHormomya and increased the upper limit ofHormomya, presumably by providing shelter and thus protecting them from desiccation. The upper limit ofHormomya zone was thus considered to be set by desiccation exceeding the physiological tolerance of the species.     

COMPONENTS OF PHOTOSYNTHESIS IN THE INTERTIDAL SACCATE ALGA HALOSACCION AMERICANUM (RHODOPHYTA,PALMARIALES)1

Rates of photosynthesis for the intertidal saccate alga Halosaccion americanum Lee were determined under submersed and emersed conditions. By fitting the data to a hyperbolic tangent function, P _max was 4.08 mmol CO₂. m^?2. h^?1 and I_k was 116.4 μE. m^?2. s^?1. under submersed conditions. Under emersed conditions, P _max was 1.89 mmol CO₂. m^?2. h^?1 and I_k was 22.9 μE. m^?2. s^?1. Dark fixation represented 3.7% of P_max in submersed thalli, whereas it equalled 33.3% of P_max in emersed thalli. Photosynthetic uptake from the thallus cavity represented a significant source of carbon, achieving 68.8% of that from the atmosphere and 29.4% of that from seawater. Retained seawater also greatly reduced drying under emersed conditions. Experimental thalli lost 70.4% of their water after 120 min under desiccating conditions, whereas control thalli lost only 6.3%. Emersed photosynthetic rates were enhanced by desiccation, At times, rates for desiccated thalli were two times those of fully-hydrated ones. Only after water loss exceeded 47% did photosynthetic rates fall below fully-hydrated rates. Utilizing data from this study a model was constructed to determine total photosynthetic production of H. americanum over a single daylight period. These caluclations demonstrate that photosynthetic contributions from emersed photosynthesis and retained seawater are significant. Because production from all sources is almost equal, total photosynthesis over a single day does not change greatly regardless of the time spent in air or in water.     

Effects of desiccation and CO2 concentrations on emersed photosynthesis in Porphyra haitanensis (Bangiales,Rhodophyta), a species farmed in China

The effects on photosynthesis of CO₂ and desiccation in Porphyra haitanensis were investigated to establish the effects of increased atmospheric CO₂ on this alga during emersion at low tides. With enhanced desiccation, net photosynthesis, dark respiration, photosynthetic efficiency, apparent carboxylating efficiency and light saturation point decreased, while the light compensation point and CO₂ compensation point increased. Emersed net photosynthesis was not saturated by the present atmospheric CO₂ level (about 350?ml?m^?3), and doubling the CO₂ concentration (700?ml?m^?3) increased photosynthesis by between 31% and 89% at moderate levels of desiccation. The relative enhancement of emersed net photosynthesis at 700?ml?m^?3 CO₂ was greater at higher temperatures and higher levels of desiccation. The photosynthetic production of Porphyra haitanensis may benefit from increasing atmospheric CO₂ concentration during emersion.     

SURVIVAL OF FUCOID EMBRYOS IN THE INTERTIDAL ZONE DEPENDS UPON DEVELOPMENTAL STAGE AND MICROHABITAT1

Embryos of the fucoid alga Pelvetia fastigiata (J. Ag.) DeToni were outplanted into the intertidal zone to assess survival during the physical stress brought about by emersion during a single low tide. Survival varied among microhabitats. Under the adult Pelvetia canopy, survival of 6-h-, 24-h-, 48-h-, and 1-wk-old embryos was nearly 100%. Almost all embryos of all ages died in exposed habitats on bare rock or within habitats where the Pelvetia canopy was removed experimentally. However, within red algal turfs, where most juvenile Pelvetia occur, survival was unusually age specific: 24- to 48-h-old embryos survived poorly compared to younger (6 h old) or older embryos (1 wk old). Survival patterns reflected microhabitat temperatures during the experiments. The fate of young post-settlement stages must be studied at these fine temporal and spatial scales to understand the organization of intertidal communities.     

THE IMPACT OF FROND CROWDING ON FROND BLEACHING IN THE CLONAL INTERTIDAL ALGA MAZZAELLA CORNUCOPIAE (RHODOPHYTA, GIGARTINACEAE) FROM BRITISH COLUMBIA, CANADA

The importance that frond crowding represents for the survival of fronds of the clonal intertidal alga Mazzaella cornucopiae (Postels et Ruprecht) Hommersand (Rhodophyta, Gigartinaceae) was investigated in Barkley Sound, British Columbia, Canada. Frond density is high for this species, up to 20 fronds·cm^?2 in the most crowded stands. Frond crowding imposes a cost in the form of reduced net photosynthetic rates when fronds are fully hydrated as a result of reduced irradiance compared with experimental (not found naturally) low-density stands. However, the interaction between desiccation and irradiance alters this relationship between net photosynthetic rates and frond density. During a typical daytime low tide in spring, irradiance is 10–30 μmol·m^?2·s^?1 below the canopy of fronds, and frond desiccation (relative to total water content) can reach 43% at the end of the low tide. In contrast to natural stands, fronds from experimentally thinned stands are subjected to irradiances up to 2000 μmol·m^?2·s^?1 because of the spatial separation among fronds and can desiccate up to 81% at the end of the same low tide. Laboratory experiments showed that negative net photosynthetic rates occur between 40% and 80% desiccation at an irradiance of 515 μmol·m^?2·s^?1, and the literature suggests that strong bleaching could occur as a result. At 20 μmol·m^?2·s^?1 of irradiance and desiccation levels up to 40%, simulating understory conditions of natural stands, net photosynthetic rates are never negative. Experimental thinning of stands of M. cornucopiae done during spring effectively resulted in a stronger extent of frond bleaching compared with natural stands. Therefore, the cost of reduced net photosynthetic rates at high frond densities when fronds are fully hydrated is counterbalanced by the protective effects of frond crowding against extensive bleaching, essential for survival at the intertidal zone. Future research will have to demonstrate the possible relationship between the frequency and duration of negative net photosynthetic rates and the extent of frond bleaching.