Relationship between seasonal growth,photosynthetic production and apex mortality of Caulerpa paspaloides (Chlorophyceae)

Seasonal changes in rates of rhizome elongation, frond initiation, photosynthesis, respiration, starch content and mortality of rhizome apices were measured for a population of Caulerpa paspaloides (Bory) Greville located off Key Largo, Florida, USA. Maximal growth rates occur during spring and coincide with high photosynthetic production and low mortality of rhizome apices. A secondary period of rapid growth occurs during fall; however, rates are less than during spring. Minimal growth rates occur during winter and summer. Reduced growth during winter coincides with (1) low photosynthetic rates, (2) low photosynthesis: respiration ratios, and (3) reduced frond biomass, indicating that photosynthetic production is limiting. Reserve carbohydrate (starch) is apparently utilized to maintain vegetative growth during the winter. Increased mortality of rhizome apices is responsible for most of the reduction in growth during summer and fall.     

Relationship between body size, provisioning rate, longevity and reproductive success in females of the solitary bee Osmia cornuta

Body size has often been related to reproductive success in bees and wasps. The objective of this 3-year study was to analyze the relationship between nesting female body size, provisioning rate and longevity and their effect on several traits related to parental investment and reproductive success in the solitary bee Osmia cornuta. Body size was not correlated to longevity, and it was only correlated to provisioning rate in the third year (with poor weather conditions during nesting). Variation in fecundity, offspring size and offspring mortality was not well explained by nesting female body size in any of the 3 years. However, in the third year, small females biased their investment toward males, the sex requiring smaller pollen–nectar provisions. Large females were more successful usurpers of other females' nests, but fecundity of usurpers was no higher than fecundity of nonusurpers. Large females were more likely to establish at the release site, probably in relation to size-dependent vigor at emergence. A review of the literature on parental investment in solitary aculeate Hymenoptera showed a stronger relationship between body size and reproductive success in wasps than in bees. In O. cornuta, fecundity was strongly related to longevity and provisioning rate in all 3 years. Offspring size was associated with provisioning rate in 1 year, when females with higher provisioning rates tended to produce larger sons and daughters. Both longevity and provisioning rate appeared to be strongly conditioned by stochastic events.     

Experimental evidence for effective and altruistic colony defence against natural predators by soldiers of the gall-forming aphid Pemphigus spyrothecae (Hemiptera : Pemphigidae)

Summary The thick-legged first instar soldiers of the gall-forming aphid Pemphigus spyrothecae Pass. are able to protect the aphids in the gall from being eaten by a range of insect predators. In artificial galls, the soldier aphids were able to kill first instar ladybirds Adalia bipunctata (L.) (Coleoptera: Coccinellidae), early third instar hoverfly larvae Eupeodes (Metasyrphus) corollae (Fab.) (Diptera: Syrphidae), and first-third instar Anthocoris nemoralis (Fab.) (Hemiptera: Cimicidae). Almost all the aphids that attacked the predators were themselves killed. The soldiers were also able to kill predators introduced into natural galls. Experiments were devised in which individuals of Anthocoris minki, which is the most important insect predator of the gall generations of Pemphigus at the study site, were free to enter and leave the gall: the soldiers were effective both in preventing the predator's access to the gall and in killing those predators that did manage to get in. In galls with experimentally manipulated numbers of soldiers and non-soldiers, it was clearly shown that it is the soldiers alone that kill the predators (Anthocoris minki and 1st instar Adalia bipunctata) (see Table 3). Even though many soldiers may die during these encounters, the selective advantage of killing the predators is high, since observations show that individual A. minki can pass through more than one instar inside a gall and kill all the aphids therein. The aphids were not observed to attack conspecific aphids from other galls or the cohabiting aphid Chaitophorus leucomelas Koch. The primary role of the soldier caste is therefore the defence of the aphid colony against predators.     

Photosynthesis,photorespiration, and dark respiration in eight species of algae

The rates of photosynthesis and dark respiration for 7 marine algae and 1 fresh-water alga were measured and compared. The dinoflagellates Glenodinium sp. and zooxanthellae have high dark respiration rates relative to photosynthetic rates, which may decrease their net growth rates. Photorespiration in the 8 algal species was studied by examining the effects of the concentration of oxygen on the rates of photosynthesis, on the incorporation of ¹⁴CO₂ into the photorespiratory pathway intermediates glycine and serine, and on the postillumination burst of carbon dioxide production and oxygen consumption. A combination of these results indicates that all the algae tested can photorespire, but that Glenodinium sp., Thalassiosira pseudonana, and zooxanthellae either have a photorespiratory pathway different from that proposed for freshwater algae (Tolbert, 1974), or an additional pathway for glycolate metabolism.     

An annual carbon budget for the kelp Laminaria longicruris

Oxygen evolution and uptake by whole thalli of the large marine alga Laminaria longicruris de la Pylaie were measured for 24 h, once every 2 weeks for a year, using large chambers to incubate the plants on the sea bed. Diel rates of photosynthesis and respiration were calculated from these measurements and continuous light measurements were used to extrapolate the data between observation dates. The resulting estimates were combined with measurements of growth and carbon content to give an annual carbon budget for a typical mature plant. Annual net assimilation was 6.8 mgC per cm² of frond surface (71 cal cm^-2). Approximately 45% of this appeared in the production of new frond tissue, and a further 12% was accounted for by storage of carbon in mature frond tissue. About 8% was needed for stipe growth, and the remaining 35% was assumed to be lost as dissolved organic carbon. Diel net photosynthetic rates reached a maximum in June and July and were negative only in November, indicating an ability to produce a photosynthetic surplus throughout winter. In early winter the plants drew on stored reserves to supplement photosynthesis in providing carbon for growth, but from January onwards photosynthesis provided more than enough carbon for growth.     

Solitary nesting and reproductive success in the paper wasp Polistes aurifer

Female Polistes paper wasps are capable of independent nesting, yet many populations demonstrate a mixture of solitary and cooperative nest foundation. Previous studies of Polistes have found survival and/or productivity advantages of cooperative nest foundation compared to solitary nesting, and reproductive skew models have been designed to predict the dynamics of such flexible cooperation. In this paper, we examine the success of different nesting strategies in a previously unstudied population of Polistes aurifer in southern California. The colony cycle of this population is less synchronous than that of other temperate species, and the frequency of solitary nesting averages 86.2%. Our data suggest that this low rate of cooperative nest founding is adaptive, as demonstrated by the lack of survival or productivity advantages for cooperative foundress associations. Due to foundress turnover and nest foundation later in the season, many nests produce only one set of offspring. This results in a loss of the eusocial nature of some nests in the population. Data from a small sample of multifoundress nests show significant positive reproductive skew, despite concession model predictions that skew should be low in populations with low ecological constraints on independent nesting. This lack of support for the concessions skew model reflects a diminished incentive for cooperation.Communicated by L. Keller     

Daily photosynthesis,respiration, and carbon budgets in a tropical marine jellyfish (Mastigias sp.)

From measured diel photosynthesis and respiration rates, using oxygen electrodes, estimates of carbon flux between symbiotic algae (zooxanthellae) and host animal are presented for the marine scyphomedusan Mastigias sp. from a marine lake in Palau, Western Caroline Islands, during February and March 1982. The carbon budgets calculated for these lake medusae indicate that carbon fixed photosynthetically by zooxanthellae and made available to the host may satisfy up to 100% of the host's daily metabolic carbon demand (CZAR). The stable carbon isotope (¹³C) signature of the mesogleal carbon of lake Mastigias sp. was close to that of the zooxanthellae, supporting the interpretation that while these medusae may feed holozoically, some of their carbon comes from their symbionts. The diel photosynthesis, respiration, and preliminary estimates of carbon budgets of three individuals of another ecotype of Mastigias sp. collected from nearby oceanic lagoons are also given. Photosynthesis of lagoon medusae was generally greater than that for lake medusae of similar size, and lagoon medusae were phototrophic with respect to carbon, with commensurately greater CZAR values. Carbon translocated from the symbiotic algae also may contribute to the growth requirements of both lake and lagoon medusae. From carbon flux data, the lake jellyfish were estimated to contribute about 16% to the total primary productivity of their marine lake habitat.     

Host plant quality and local adaptation determine the distribution of a gall-forming herbivore

Herein we report results of transplant experiments that link variation in host plant quality to herbivore fitness at the local scale (among adjacent plants) with the process of local (demic) adaptation at the landscape scale to explain the observed distribution of the specialist gall former Belonocnema treatae (Hymenoptera: Cynipidae) within populations of its host plant, Quercus fusiformis. Field surveys show that leaf gall densities vary by orders of magnitude among adjacent trees and that high-gall-density trees are both rare (< 5%) and patchily distributed. B. treatae from each of five high-gall-density trees were reared on (1) the four nearest low-gall-density trees, (2) the four alternative high-gall-density trees, and (3) their natal trees (control). Each treatment (source X rearing site) was replicated three times. Nine components of performance that sequentially contribute to fitness were evaluated with over 21000 galls censused across the 25 experimental trees. When reared on their natal trees and compared with low-gall-density neighbors, transplanted gall formers had higher gall initiation success (P < 0.05), produced more (P < 0.001) and larger galls (P < 0.001), and produced a higher proportion of galls that exceeded the threshold size for natural enemy avoidance (P < 0.05). Comparison of gall-former performance on natal vs. alternative high-gall-density trees demonstrated significant (P < 0.001) differences in six performance measures with five differing in the direction predicted by the hypothesis of local adaptation. Overall, these linked experiments document direct and indirect effects of host plant variation on gall-former performance and demonstrate convincingly that (1) high-gall-density trees equate to high-quality trees that are surrounded by trees of relatively lower quality to the herbivore and (2) gall-former populations have become locally adapted to individual trees.     

Sensitivity of mesquite shrubland CO2 exchange to precipitation in contrasting landscape settings

In semiarid ecosystems, physiography (landscape setting) may interact with woody-plant and soil microbe communities to constrain seasonal exchanges of material and energy at the ecosystem scale. In an upland and riparian shrubland, we examined the seasonally dynamic linkage between ecosystem CO2 exchange, woody-plant water status and photosynthesis, and soil respiration responses to summer rainfall. At each site, we compared tower-based measurements of net ecosystem CO2 exchange (NEE) with ecophysiological measurements among velvet mesquite (Prosopis velutina Woot.) in three size classes and soil respiration in sub-canopy and inter-canopy micro-sites. Monsoonal rainfall influenced a greater shift in the magnitude of ecosystem CO2 assimilation in the upland shrubland than in the riparian shrubland. Mesquite water status and photosynthetic gas exchange were closely linked to the onset of the North American monsoon in the upland shrubland. In contrast, the presence of shallow alluvial groundwater in the riparian shrubland caused larger size classes of mesquite to be physiologically insensitive to monsoonal rains. In both shrublands, soil respiration was greatest beneath mesquite canopies and was coupled to shallow soil moisture abundance. Physiography, through its constraint on the physiological sensitivity of deeply rooted woody plants, may interact with plant-mediated rates of soil respiration to affect the sensitivity of semiarid-ecosystem carbon exchange in response to episodic rainfall.     

A comparative analysis of the photobiology of zooxanthellae and zoochlorellae symbiotic with the temperate clonal anemone <Emphasis Type="Italic">Anthopleura elegantissima</Emphasis> (Brandt). III. Seasonal effects of natural light and temperature on photosynthesis and respiration

The sea anemone Anthopleura elegantissima hosts two phylogenetically different symbiotic microalgae, a dinoflagellate Symbiodinium (zooxanthellae, ZX) and a chlorophyte (zoochlorellae, ZC). The photosynthetic productivity (P), respiration (R), and contribution of algal carbon translocated to the host (CZAR) in response to a year’s seasonal ambient changes of natural light and temperature are documented for both ZX- and ZC-bearing anemones. Light and temperature both affect photosynthesis, respiration, and CZAR, as well as various algal parameters; while there are evident seasonal differences, for the most part the relative effects on P, R, and CZAR by the two environmental variables cannot be determined. Net photosynthesis (P_n) of both ZX and ZC was significantly higher during spring and summer. During these seasons, the P_n of ZX was always greater than that of ZC. Regardless of algal symbiont, anemone respiration (R) was significantly higher during the spring and summer. The annual net carbon fixation rate of anemones with ZX and ZC was 325 and 276 mg C anemone⁻¹ year⁻¹, respectively, which translates to annual net community productivity rates of 92 and 60 g C m⁻¹ year⁻¹ for anemones with ZX or ZC, respectively. CZAR did not show a clear relationship with season; however the CZAR for ZX was always significantly greater than for ZC. Lower ZX growth rates, coupled with higher photosynthetic rates and higher CZAR estimates, compared to ZC, suggest that if A. elegantissima is simply carbon limited, ZX-bearing anemones should be the dominant symbiont in the field. However ZC-bearing anemones persist in low light and reduced temperature microhabitats, therefore more than the translocation of carbon from ZC must be involved. Given that global climate change will increase water temperatures, the potential for latitudinal range shifts of both ZC and ZX (S. californium and muscatinei) might be used as biological indicators of thermal shifts in the littoral zone of the Pacific Northwest.     

Host-plant genotypic diversity mediates the distribution of an ecosystem engineer

Ecosystem engineers affect ecological communities by physically modifying the environment. Understanding the factors determining the distribution of engineers offers a powerful predictive tool for community ecology. In this study, we examine whether the goldenrod bunch gall midge (Rhopalomyia solidaginis) functions as an ecosystem engineer in an old-field ecosystem by altering the composition of arthropod species associated with a dominant host plant, Solidago altissima. We also examine the suite of factors that could affect the distribution and abundance of this ecosystem engineer. The presence of bunch galls increased species richness and altered the structure of associated arthropod communities. The best predictors of gall abundance were host-plant genotype and plot-level genotypic diversity. We found positive, nonadditive effects of genotypic diversity on gall abundance. Our results indicate that incorporating a genetic component in studies of ecosystem engineers can help predict their distribution and abundance, and ultimately their effects on biodiversity.     

Impact of an invasive oak gall wasp on a native butterfly: a test of plant-mediated competition

Phytophagous insects commonly interact through shared host plants. These interactions, however, do not occur in accordance with traditional paradigms of competition, and competition in phytophagous insects is still being defined. It remains unclear, for example, if particular guilds of insects are superior competitors or important players in structuring insect communities. Gall-forming insects are likely candidates for such superior competitors because of their ability to manipulate host plants, but their role as competitors is understudied. We investigate the effect of invasive populations of an oak gall wasp, Neuroterus saltatorius, on a native specialist butterfly, Erynnis propertius, as mediated by their shared host plant, Quercus garryana. This gall wasp occurs at high densities in its introduced range, where we stocked enclosures with caterpillars on trees that varied in gall wasp density. Biomass production of butterflies was lower in enclosures on high-density than on low-density trees because overwintering caterpillars were smaller, and fewer of them eclosed into adults the following spring. To see if the gall wasp induced changes in foliar quality, we measured host plant quality before and after gall induction on 30 trees each at two sites. We found a positive relationship between gall wasp density and the percentage change in foliar C:N, a negative relationship between gall wasp density and the percentage change in foliar water at one site, and no relationship between the percentage change in protein-binding capacity (i.e., phenolics) and gall-wasp density. Additionally, there was a negative relationship between foliar quality and butterfly performance. Our results provide evidence for a plant-mediated impact of an invasive oak gall wasp on a native butterfly and suggest that gall wasps could act as superior competitors, especially when they occur at high densities.     

Role of echinoderms in benthic remineralization in the Chukchi Sea

The role of large, epibenthic organisms in carbon cycling at high latitudes is difficult to assess using standard ship-board collection techniques. We used a remotely operated vehicle equipped with video imaging to examine the distribution and abundance of epibenthic organisms in the northeast Chukchi Sea during June 1998. At each of 11 sites, we collected between 25 and 50 images from a minimum of 20 min of video. We observed 15 different epibenthic taxa, with the echinoderms (Ophiura sarsia, O. maculata, Ophiopholis aculeata, Stegophiura nodosa, and Echinarachinus parma) overwhelmingly dominating the epibenthos. Echinoderm density was highly variable, ranging from 0.2 to 256.6 individuals m^-2 (median=16.3), and echinoderm biomass varied between <0.5 and 4,988 mg C m^-2 (median=737). The highest biomass of ophiuroids recorded (3,388 mg C m^-2) is 30% higher than the highest previously reported from an Arctic shelf. Using a relationship between biomass and respiration developed for deep-sea organisms living at cold temperatures, we estimated respiration rates from <0.1 to 15.0 mg C m^-2 day^-1 (median=1.9). Respiration rates measured on board were several orders of magnitude higher than those obtained from the predictive equation. Further work is needed to assess echinoderm respiration rates accurately under in situ conditions. Even with calculated minimal values for respiration rates, a comparison of epifaunal and infaunal respiration at four stations revealed that echinoderm respiration accounted for as much as 25% of total respiration. High epifaunal respiration rates and biomass values are likely supported by high concentrations of particulate organic carbon carried by Bering Sea water flowing through the eastern Chukchi Sea. Our observations support observations from the Eurasian Arctic that echinoderms dominate the epibenthos of Arctic shelves and that the role of these organisms in carbon remineralization must be considered if we are to generate accurate models of carbon cycling in the Arctic.     

Habitat type determines herbivory controls over CO2 fluxes in a warmer Arctic

High-latitude ecosystems store large amounts of carbon (C); however, the C storage of these ecosystems is under threat from both climate warming and increased levels of herbivory. In this study we examined the combined role of herbivores and climate warming as drivers of CO2 fluxes in two typical high-latitude habitats (mesic heath and wet meadow). We hypothesized that both herbivory and climate warming would reduce the C sink strength of Arctic tundra through their combined effects on plant biomass and gross ecosystem photosynthesis and on decomposition rates and the abiotic environment. To test this hypothesis we employed experimental warming (via International Tundra Experiment [ITEX] chambers) and grazing (via captive Barnacle Geese) in a three-year factorial field experiment. Ecosystem CO2 fluxes (net ecosystem exchange of CO2, ecosystem respiration, and gross ecosystem photosynthesis) were measured in all treatments at varying intensity over the three growing seasons to capture the impact of the treatments on a range of temporal scales (diurnal, seasonal, and interannual). Grazing and warming treatments had markedly different effects on CO2 fluxes in the two tundra habitats. Grazing caused a strong reduction in CO2 assimilation in the wet meadow, while warming reduced CO2 efflux from the mesic heath. Treatment effects on net ecosystem exchange largely derived from the modification of gross ecosystem photosynthesis rather than ecosystem respiration. In this study we have demonstrated that on the habitat scale, grazing by geese is a strong driver of net ecosystem exchange of CO2, with the potential to reduce the CO2 sink strength of Arctic ecosystems. Our results highlight that the large reduction in plant biomass due to goose grazing in the Arctic noted in several studies can alter the C balance of wet tundra ecosystems. We conclude that herbivory will modulate direct climate warming responses of Arctic tundra with implications for the ecosystem C balance; however, the magnitude and direction of the response will be habitat-specific.     

Different habitats, different habits? Response to foraging information in the parasitic wasp Venturia canescens

The parasitic wasp, Venturia canescens (Gravenhorst) (Hymenoptera: Ichneumonidae), has two reproductive modes, namely, thelytoky or arrhenotoky, and occurs in habitats with highly variable or relatively stable host abundances, respectively. Since information processing is costly, we expected that information indicating resource availability would be mainly used by the thelytokous wasps and less so by the arrhenotokous type. This idea was explored by two different approaches. In a study on patch-time allocation, we used females from ten populations and measured patch-residence times of individuals that visited multiple patches at different encounter rates. In a more detailed approach, thelytokous and arrhenotokous females from a single location were observed continuously while foraging, and all behaviors were recorded. Wasps of both reproductive modes (i.e., both habitat types) used information for the assessment of habitat quality. However, the way that the information was used differed between them. Whereas thelytokous females used foraging information to maximize their efficiency at high patch-encounter rates, arrhenotokous females merely reduced the number of offspring produced without changing patch times. The behavior of the arrhenotokous females should result in a spreading of offspring across the habitat and, thus, reduced sib-mating. The foraging strategy of these wasps might therefore be an adaptation to reduce costs associated with inbreeding.     

The mechanism of calcification and its relation to photosynthesis and respiration in the scleractinian coral <Emphasis Type="Italic"> Galaxea fascicularis</Emphasis>

The mechanism of calcification and its relation to photosynthesis and respiration were studied with Ca²⁺, pH and O₂ microsensors using the scleractinian coral Galaxea fascicularis. Gross photosynthesis (Pg), net photosynthesis (Pn) and dark respiration (DR) were measured on the surface of the coral. Light respiration (LR) was calculated from the difference between Pg and Pn. Pg was about seven times higher than Pn; thus, respiration consumes most of the O₂ produced by the algal symbiont's photosynthesis. The respiration rate in light was ca. 12 times higher than in the dark. The coupled Pg and LR caused an intense internal carbon and O₂ cycling. The resultant product of this cycle is metabolic energy (ATP). The measured ATP content was about 35% higher in light-incubated colonies than in dark-incubated ones. Direct measurements of Ca²⁺ and pH were made on the outer surface of the polyp, inside its coelenteron and under the calicoblastic layer. The effects on Ca²⁺ and pH dynamics of switching on and off the light were followed in these three compartments. Ca²⁺ concentrations decreased in light on the surface of the polyp and in the coelenteron. They increased when the light was switched off. The opposite effect was observed under the calicoblastic layer. In light, the level of Ca²⁺ was lower on the polyp surface than in the surrounding seawater, and even lower inside the coelenteron. The concentration of calcium under the calicoblastic layer was about 0.6 mM higher than in the surrounding seawater. Thus Ca²⁺ can diffuse from seawater to the coelenteron, but metabolic energy is needed for its transport across the calicoblastic layer to the skeleton. The pH under the calicoblastic layer was more alkaline compared with the polyp surface and inside the coelenteron. This rise in pH increased the supersaturation of aragonite from 3.2 in the dark to 25 in the light, and brought about more rapid precipitation of CaCO₃. When ruthenium red was added, Ca²⁺ and pH dynamics were inhibited under the calicoblastic layer. Ruthenium red is a specific inhibitor of Ca-ATPase. The results indicated that Ca-ATPase transports Ca²⁺ across the calicoblastic layer to the skeleton in exchange for H⁺. Addition of dichlorophenyldimethylurea completely inhibited photosynthesis. The calcium dynamics under the calicoblastic layer continued; however, the process was less regular. Initial rates were maintained. We conclude that light and not energy generation triggers calcium uptake; however, energy is also needed.     

The flux of particulate carbon in an estuary

The major sources and sinks of suspended particulate carbon are identified for northern Chesapeake Bay, Maryland, USA. The area of the bay under consideration was divided into two sections. The northernmost section (upper bay), from the head of the bay to the vicinity of Baltimore, was characterized by a high input of particulate carbon from upland drainage. The section from Baltimore to the mouth of the Potomac River (middle bay) was dominated by an internal supply of particulate carbon from primary production. The northernmost section was characterized by major particulate carbon losses to the bottom and to benthic respiration, with slightly less than half (43%) of the particulate carbon respired in the water column or converted to dissolved organic matter. In contrast, respiration in the water column or conversion to dissolved organic matter was responsible for the loss of about 65% of the particulate carbon in the middle section of the bay, while losses to the bottom and to benthic respiration totaled only 12%. Even though the supply of particulate carbon to the upper bay was about 1.5 times the supply to the middle bay, the loss of carbon due to biological activity (biological efficiency) in each area was similar, and amounted to about 70% of the supply of particulate carbon.The research reported here was partially supported by the University of Maryland and by the U. S. Bureau of Sport Fisheries and Wildlife under Contract No. 14-16-0005-2096. Contribution No. 471 of the Natural Resources Institute, University of Maryland, USA.     

Nutrient cycling between the water column and a marine sediment. I. Organic carbon

The carbon flow through the sediments at a station located in 18.3 m of water off the Scripps Institution of Oceanography, San Diego, California (USA) was determined. The parameters studied [and their mean rates of input (+) or output (-) to the benthos] were macro-detritus (+0.028 gC m^-2day^-1), fallout of particulate debris (+3.3 gC m^-2day^-1), benthic net photosynthesis during the day (-0.06 gC m^-2 daylight period^-1), burial (0 gC m^-2day^-1), benthic respiration at night (-0.28 gC m^-2 night period^-1), and resuspension (-3.0 gC m^-2day^-1). Resuspension of sediment at this station was found to have a controlling effect on the sediment organic carbon content. Benthic photosynthesis was able to provide 79% of the organic carbon required by the benthos for respiration during the daylight hours. A carbon-flow diagram linking together all of the above measurements is presented.     

Freezing rate and duration determine the physiological response of intertidal fucoids to freezing

Differential thermal analysis (DTA) was used to measure the freezing temperature of nine species of red brown intertidal macroalgae from the coast of Maine, USA in 1991. Using slow and rapid cooling rates approximating those found in the field for Ascophyllum nodosum (L.) Le Jol. we found that, for a given rate, the freezing points of all species were similar: -7.06 to -8.02°C for slow cooling (ca. 0.25°C min^-1) and -3.42 to -4.56°C for rapid cooling (ca. 5.0°C min^-1). In the low shore species, Fucus evanescens C. Ag., photosynthesis was inhibited to a greater extent when plants were frozen or thawed rapidly than after slow freezing or thawing. However, in the upper shore species, F. spiralis (L.), photosynthesis recovered rapidly and completely regardless of freezing rate. Rapidly frozen F. evanescens also experienced greater loss of plasmalemmal integrity, evidenced by a greater loss of cellular contents on re-immersion, than those frozen slowly. Light-limited photosynthesis following freezing was more severely inhibited than light-saturated photosynthesis. Respiration was generally enhanced immediately after freezing, but then declined to rates below those of unfrozen controls within 2 h following re-immersion, with control rates of respiration being achieved after a 24 h recovery period. Our data suggest that the physiological consequences of winter emersion at sub-zero temperatures may vary widely between individual plants of freezing-susceptible species, due to the wide variations in freezing rate associated with microhabitat effects.