Sensitivity and acclimation to UV radiation of zoospores from five species of Laminariales from the Arctic

Spores of five Laminariales from Arctic Spitsbergen were exposed in the laboratory to photosynthetically active radiation (PAR; 400–700 nm), PAR+UVA radiation (UVAR; 320–400 nm) and PAR+UVAR+UVB radiation (UVBR; 280–320 nm). Subsequently, germination was monitored over periods of 3, 6 and 9 days. The investigated species were the upper sublittoral Saccorhiza dermatodea, the upper to mid-sublittoral Alaria esculenta and Laminaria digitata, the mid-sublittoral L. saccharina and the lower sublittoral L. solidungula. The germination capacity decreased sharply after 16 h exposure to PAR+UVAR+UVBR in all species. However, S. dermatodea was able to recover from the damaging effects of UVBR. There was also a small increase in percentage germination of A. esculenta 6–9 days after the treatment. No recovery was evident in the other species. After 8 h exposure to PAR+UVA+UVB, L. digitata recovered completely, and L. saccharina and L. solidungula, partially. The only species susceptible to PAR+UVAR was L. solidungula. One prominent cytological feature of UVR-exposed spores was the enlargement of phenolic vesicles (physodes) (particularly seen in S. dermatodea and A. esculenta), which may have a protective function against UVR. Pilot experiments under natural irradiance conditions indicate that the PAR component of solar radiation exerts an additional stress. Overall the data show that zoospores of the species from the upper sublittoral are less sensitive to UVR or have the capacity to recover from UV stress in contrast to species from deeper waters, probably due to their UV protective and repair capabilities.Communicated by O. Kinne, Oldendorf/Luhe     

Standing crop and leaf area index of the sublittoral Laminaria species near Helgoland

Standing crop values of Laminaria species sampled between 0 to 2 m depth in the sublittoral of Helgoland, a small rocky island in the southern North Sea, are similar to those reported by other workers from Norwegian or Scottish coasts. Below 2 m water depth, however, standing crop values are found to be reduced to a far greater extent with increasing depth than in the Norwegian or Scottish areas studied. Maximum standing crop values are not found in Laminaria digitata and L. saccharina growing just below the low water mark of spring tides, but in the upper L. hyperborea forest at 2 m depth. Population density and leaf area index (LAI) decrease with increasing depth (but not the average weight of individuals constituting the L. hyperborea forest). It is suggested that the depth at which LAI becomes 1 represents an important ecological threshold, since growth rates of L. hyperborea specimens appear to decrease at greater depths but not to increase above the threshold. Self-shading might equalize, to a certain extent, the quantity of underwater irradiance available to an individual phylloid at all depths at which a close vegetation of Laminaria species occurs.     

CO2 fixation and translocation in benthic marine algae. III. Rates and ecological significance of translocation in Laminaria hyperborea and L. saccharina

Translocation of ¹⁴C-labelled assimilates in Laminaria hyperborea and L. saccharina is strictly basipetal within the thallus. The lowermost 20 cm of the frond accumulates 71% (L. hyperborea) or 63% (L. saccharina) of the basipetallymoving translocate after a period of 96 h from initial feeding with NaH¹⁴CO₃. Whereas translocation was found to occur at all seasons in L. saccharina, which grows throughout the year, no translocation of newly synthesized photosynthate occurred in L. hyperborea in October, when no frond enlargement takes place, nor in January, when the new fond occurs as a tiny outgrowth. In L. hyperborea, organic material, assimilated by the old frond during the period of rapid growth, is exported at the rate of 0.5 mg C · dm^-2 · h^-1. In L. saccharina, 14 to 18 cm frond length are necessary for an elongation rate of the growing zone, situated in lowermost 10 cm of the frond, of 1 cm per week. The significance of translocation in the two Laminaria species is demonstrated by comparison of export rates with growth rates measured in situ and in the laboratory.Dedicated to Professor Dr. Otto Stocker on his 85th birthday.     

Influence of ultraviolet-radiation on chlorophyll fluorescence and growth in different life-history stages of three species of Laminaria (phaeophyta)

Zoospores, gametophytes, young sporophytes and discs cut from mature sporophytes of Laminaria digitata, L. hyperborea and L. saccharina were exposed in the laboratory to UV-radiation, with a spectral composition and irradiance similar to natural sunlight, for periods ranging from 15 min to 8 d, and were then returned to white light. Germination of zoospores and the growth of gametophytes were reduced after exposures to UV longer than 1 h, whereas UV had little effect on the growth of young or mature sporophytes unless exposure continued for more than 48 h. The variable fluorescence (F _v:F_m) of all stages was strongly reduced immediately after short exposures to UV, but recovered almost completely within 24 h. However, exposure of gametophytes to UV for >4 h resulted in little or no recovery of F _v:F_m, whereas >16 h of UV were required to produce this result in young sporophytes, and >48 h in mature sporophytes. Thus, sensitivity to UV-radiation decreased from gametophytes to sporophytes, and with increasing age of sporophytes, but, in gametophytes, growth appeared to be a more sensitive indicator of UV-damage than F _v:F_m after 24 h recovery. The responses to UV of the zoospores and gametophytes of all three species were similar, but both growth and fluorescence measurements suggested that the sporophytes of L. saccharina were more sensitive to UV than those of the other two species.     

Optimal growth and maximal survival temperatures of Atlantic Laminaria species (Phaeophyta) in culture

The effects of temperature on growth rate of rapidly-growing cultured macrosporophytes of 9 isolates of Atlantic Laminaria comprising 4 species have been investigated. No significant population variation was observed within species despite wide variations in temperature between the original collecting sites. L. saccharina showed a broad temperature optimum in the 10°–15°C range, whereas L. longicruris had a sharp optimum at 10°C. L. digitata and L. hyperborea grew more slowly, with only slightly sub-optimal growth over a wide temperature range, but with peaks at 10°C (L. digitata) and 15°C (L. hyperborea). The maximum survival temperatures of individual male and female vegetatively-growing gametophytes were ascertained for these species plus the Arctic L. solidungula, and were as follows: L. saccharina and L. longicruris, 23°C; L. digitata (male), 23°C; L. digitata (female), 22°C; L. hyperborea, 21°C; L. solidungula, 18°C. The lack of within-species differences demonstrates that the success of the genus in areas with different temperature regimes is brought about by phenotypic plasticity of individuals rather than the selection of temperature races or ecotypes.     

Effects of ultraviolet radiation on different life cycle stages of the south Pacific kelps, Lessonia nigrescens and Lessonia trabeculata (Laminariales, Phaeophyceae)

The effects of exposure to ultraviolet radiation (UVR), 280–400 nm, in different life histories and development stages of the kelps, Lessonia nigrescens and L. trabeculata, collected in the south-east Pacific coast (30°S) were evaluated in the laboratory. Germination and viability (motile zoospores, settled spores), diameter of the primary cell of the gametophytes, percentage of female gametophytes, fertility and sporophytes production were measured after exposure to three radiation treatments (PAR; PAR + UVA; PAR + UVA + UVB). The effects of UVR in young sporophytes (diploid stage) were evaluated as changes in maximal quantum yield of chlorophyll fluorescence of photosystem II (PSII) (F _v/F _m). A significant decrease in all variables was observed for the treatment that included UVB (PAR + UVA + UVB) after 2 and 4 h of exposure, in relation to the control. The motile spores were more sensitive to UVR exposure compared to settled spores and gametophytes, suggesting that along with an increase in ontogenetic development; there is an increase in the tolerance to UVR. In addition, it was observed that early stages of the intertidal L. nigrescens were more tolerant to UVR compared to the subtidal L. trabeculata. These results allow initially to infer that UVR may be regarded as an important environmental factor influencing the upper limit of distribution of these species, mainly through its detrimental effects on the early stages of the life cycle.     

Transversal profiles of carbon assimilation in the fronds of three Laminaria species

The brown macroalgae Laminaria digitata (Huds.) Lamour., L. hyperborea (Gunn.) Foslie, and L. saccharina (L.) Lamour. (Laminariales, Phaeophyceae) have been investigated for carbon assimilation and assimilate biosynthesis in different tissues (meristoderm, cortex, medulla) of stipes and blades. Carboxylation via ribulose-1,5-bisphosphate carboxylase (EC 4.1.1.39) and phosphoenolpyruvate carboxykinase (EC 4.1.32) follows a transversal gradient from the outer to the inner tissues with maximum activity of enzymatic CO₂ fixation in the meristoderm. Such transversal profiles are also recorded for photosynthetic and dark carbon assimilation as well as for kinetics of assimilate ¹⁴C-labelling in isolated tissues. Differential chemical constitution of the blade and stipe tissues examined and results from assimilate biosynthesis are discussed with regard to a transversal translocation of photosynthates from meristodermal to cortical and medullar cells.     

UV-radiation versus grazing pressure: long-term floating of kelp rafts (<Emphasis Type="Italic">Macrocystis pyrifera</Emphasis>) is facilitated by efficient photoacclimation but undermined by grazing losses

Large quantities of floating macroalgae are traveling in coastal waters of the SE Pacific and in other temperate climate zones. While afloat, these algae are potentially exposed to full solar radiation, including UVA and UVB, which can have profound effects on their physiological and growth performance. Latitudinal variations in UV-radiation (UVR) are hypothesized to affect floating algae differently with higher impacts at low latitudes than at high latitudes. In addition, UVR together with grazing might accelerate the demise of floating kelps. This hypothesis was tested with outdoor laboratory experiments in which sporophytes of the giant kelp Macrocystis pyrifera (L.) C. Agardh were exposed to a combination of different UVR regimes (PAR only, PAR + UV) and grazing at three sites along the Chilean coast (20°S, 30°S, and 40°S). A latitudinal trend in irradiance was detected with increasing values from 40°S to 20°S. Surprisingly, floating M. pyrifera responded with a high acclimation potential within this latitudinal UVR gradient. At 20°S, floating kelps were slightly sensitive to UVR, which was reflected in reduced blade growth. At 30°S, physiological responses were hardly affected by the prevailing irradiance but sporophyte growth and thus persistence mainly depended on the presence or absence of amphipod grazers. At high latitudes, grazing had only minor impacts on algal biomass and blade growth, and kelps thrived well under all tested environmental conditions. Overall, our results reveal that floating M. pyrifera was only slightly affected by UVR and that sporophytes can efficiently acclimate over a latitudinal UVR gradient that spans from 20°S to 40°S. Given this high acclimation potential, we suggest that these (and possibly other) positively buoyant algae are important dispersal agents over a wide range of temperate latitude conditions.     

Growth of amputated and dark-exposed individuals of the brown alga Laminaria hyperborea

The seasonal growth of the brown alga Laminaria hyperborea (Gunn.) Foslie, which forms dense forests in the sublittoral zone of Helgoland, a rocky island in the Southern North Sea, was investigated by transplanting specimens of medium size onto PVC or wooden plates fixed to sub-tidal grwoth stations. The 2-year study revealed that, during the season of fast growth (January to June), young plants produce each year a new frond, larger in size than in the preceding year. This observation can be interpreted, in part, by assuming that the growth of the young frond is accomplished not only by its own assimilatory surplus, but also by reserve materials of the old frond assimilated during the preceding season of slow growth. This assumption is supported by experimental evidence: (1) Even in complete darkness Laminaria hyperborea is capable of producing a small new frond during the season of fast growth; (2) New fronds with stipe, but without old frond, grow (during the same period) considerably faster than isolated new fronds separated from the possible sources of reserve materials. Maximum growth occurs in normal plants which possess their old frond until April or May.An abbreviated version of this paper has been presented at the Sixth International Seaweed Symposium, held at Santiago de Compostela, Spain, September 9 to 13, 1968.     

Effects of ambient levels of solar ultraviolet radiation on zooxanthellae and photosynthesis of the reef coral Montipora verrucosa

Paired flat plates of the hermatypic coral Montipora verrucosa from Kaneohe Bay, Oahu, Hawaii, were acclimated to photosynthetically active radiation (PAR) only and to full sunlight (PAR+UV) for several weeks in the summer of 1990. After the acclimation period, photosynthesis, both in PAR-only and PAR+UV as well as dark respiration were measured. Levels of the UV-absorbing compounds, S320, density of zooxanthellae, and chlorophyll a concentration were determined. Corals acclimated in PAR+UV had higher levels of the UV-protective compounds and lower areal zooxanthellae densities than corals acclimated in PAR-only. Chlorophyll a per unit volume of coral host and per algal cell did not differ between corals from the two acclimation treatments. Corals acclimated to PAR+UV displayed higher photosynthesis in full sunlight than corals acclimated to PAR-only, but when photosynthesis was measured in the light regime to which the corals had been acclimated, there were no differences in photosynthesis. Dark respiration was the same for corals from the two acclimation treatments regardless of the light quality immediately preceding the dark period.Contribution No. 902 HIMB     

Ecotypic differentiation in the kelpLaminaria saccharina: Phase-specific adaptation in a complex life cycle

Comparison of cultured sporophytes and gametophytes in common-garden experiments confirmed the existence of ecotypic differences in light-related traits among populations ofLaminaria saccharina (L.) Lamour. Cultured sporophytes from the turbid habitat in Long Island Sound, New York, USA, grew faster under both limiting and saturating daily irradiances than sporophytes from shallow and deep habitats along the Atlantic coast of Maine. Rapid growth of turbid plants was attributable to several factors, including high photosynthetic capacity and efficiency [due to differences in photosynthetic unit (PSU) number and size], low respiration rates, and high surface area:weight ratios. In contrast to sporophytes, microscopic gametophytes from the three kelp populations grew at similar rates under limiting and saturating daily irradiances. Biomass-specific photosynthesis vs irradiance (PI) parameters were similar for gametophytes from the shallow, deep, and turbid sites, despite population differences in chlorophyll-specific PI parameters and PSU characteristics. However, turbid gametophytes produced microscopic sporophytes more rapidly than gametophytes from the shallow and deep sites, apparently due to a lower blue-light requirement for gametogenesis. Ecotypic differences in sporophytes and gametophytes ofL. saccharina from shallow, deep, and turbid habitats can be understood as phase-specific adaptations.     

The section Digitatae of the genus Laminaria (Phaeophyta) in the northern and southern Atlantic: crossing experiments and temperature responses

Hybridization experiments between seven north and south Atlantic Laminaria species were carried out. Morphologically normal F₁ sporophytes developed from the following crosses among south Atlantic species: L. pallida x L. schinzii, L. pallida x L. abyssalis and L. schinzii x L. abyssalis. Normal F₁ sporophytes also resulted from the crosses L. digitata (north Atlantic) x L. pallida (south Atlantic) and L. digitata (north Atlantic) x L. abyssalis (south Atlantic). Hybrids between north Atlantic L. ochroleuca and south Atlantic L. pallida, L. schinzii and L. abyssalis and between north Atlantic L. digitata and south Atlantic L. schinzii initially developed as normal sporophytes but became deformed later on and further development was retarded. No hybrids resulted from attempted crosses between northeastern Atlantic L. saccharina and L. abyssalis from Brazil. Temperature tolerance, relative growth rates and temperature demands for gametogenesis revealed the existence of a warm temperate group within the digitate Laminaria species consisting of L. ochroleuca, L. pallida, L. schinzii and L. abyssalis. Hybridization experiments and temperature responses suggest that north Atlantic L. digitata and L. ochroleuca are still similar to south Atlantic Laminaria species, confirming the speculation that a transequatorial migration of a warm-temperate L. ochroleuca-like ancestor may have taken place.     

Ecotypic differentiation in light-related traits of the kelp Laminaria saccharina

Light-related traits were compared for Laminaria saccharina Lamour. collected from three habitats in Maine and New York, USA, with different ambient light regimes. Light-level, expressed as a proportion of surface irradiance (I₀), ranged from 0.04 to 0.32 I₀ in the shallow habitat, but rarely exceeded 0.04 I₀ in the deep and turbid habitats. Juvenile sporophytes collected from each habitat in April, 1985, were grown at four acclimation light-levels (0.065, 0.12, 0.26, and 0.54 I₀) in a common-garden, laboratory experiment. Photosynthesis vs irradiance (PI) parameters, light-harvesting characteristics, and rates of carbon-assimilation and growth were determined for each group of plants. The results indicated that ecotypic differentiation had occurred among the three kelp populations. Photosynthetic capacity (P_max) and photosynthetic efficiency () were generally highest for plants from the turbid habitat, lowest for deep plants, and intermediate for shallow plants. These differences were largely attributable to variations in light-harvesting characteristics. The nature and magnitude of photoacclimation responses also differed among populations. Population differences in photosynthetic parameters resulted in different rates of C-assimilation and growth by plants from shallow, deep, and turbid habitats. Predictions of in situ growth rates indicated that the severity of light-limitation and, therefore, the adaptive significance of efficient light-utilization vary among the three populations. It is concluded that ecotypic differentiation in light-related traits is important to the broad environmetal range of L. saccharina.     

Influence of local environmental conditions on the seasonal acclimation process and the daily integrated production rates of Laminaria digitata (Phaeophyta) in the English Channel

Two populations of Laminaria digitata (Hudson) Lamouroux (Ann Mus Hist Nat Paris 20:21–47, 1813) were examined for their seasonal photosynthetic acclimation to clear and turbid-light environments along the French coast of the English Channel. Photosynthesis–irradiance curves, pigment concentrations and the daily in situ integrated oxygen production rates were measured in both populations. Despite the great differences in light attenuation between the sites, the two populations achieved similar oxygen production rates in the field, in relation to high maximal photosynthetic rates, total pigment concentrations and antenna (fucoxanthin + chlorophyll c)/chl a pigment ratios in sporophytes from the turbid environment. Environmental conditions (i.e. light, temperature and nitrogen availability) changed throughout the year in both sites. While the seasonal acclimation trends were evident in the clear-light environment, the strategy in the turbid-light environment differed, tending to maximize light capture throughout the year. This study highlights the diversity of the response of a single species to contrasted light environments.     

Effects of ultraviolet radiation and water motion on the reef coral Porites compressa Dana: a flume experiment

The effects of water flow and ultraviolet radiation (UVR, 280–400 nm) on the reef coral Porites compressa Dana were explored in a manipulative flume experiment. The aim of this study was to determine whether this coral responds to changes in the UVR environment by adjusting the tissue concentration of UV-absorbing compounds (mycosporine-like amino acids, MAAs), and to see whether such an acclimation is affected by water flow. Also, calcification rate and chlorophyll-a concentration were measured during the experiment to estimate the potential costs (in the form of slowed growth and/or reduced photosynthetic capacity) to the coral–alga symbiosis of being exposed to UVR and producing MAAs. Branches of P. compressa from a single male colony were exposed to high or low flow (15 cm s⁻¹ and 3 cm s⁻¹, respectively) and ambient or no UVR in an outdoor, continuous-flow seawater system. Chlorophyll-a and MAA concentrations were determined after zero, 3 and 6 weeks of exposure to the experimental conditions. Increase in buoyant weight during the two 3-week periods was used to calculate calcification rate. The presence of UVR had a significant positive effect on total MAA concentration in the P. compressa colonies; however, there were significant interactions present. In colonies exposed to UVR, MAA concentration increased and then decreased to initial levels in high water flow, and increased steadily in low water flow. In colonies receiving no UVR, MAA concentration decreased steadily, declining 23% in 6 weeks. The absence of UVR did not result in higher chlorophyll-a concentrations, but the calcification rate was slightly affected by UVR. This study supports the putative photoprotective role of MAAs in P. compressa, and suggests that the costs of mitigating the effects of ambient UVR are detectable, but they are very small. Received: 29 February 2000 / Accepted: 20 September 2000     

Temperature acclimation of respiration and photosynthesis in the brown algaLaminaria saccharina

Sporophytes of the brown algaLaminaria saccharina (L.) Lamour grown at 15°C contained significantly more chlorophylla (chla) than did similar plants grown at 5°C. The increase in chla in 15°C plants was due to increased numbers of photosystem II reaction centes, and possibly to increased photosynthetic unit size, compared with 5°C plants. These changes were associated with increased values (photosynthetic efficiencies) in 15°C-grownL. saccharina relative to 5°C-grown plants. The changes in together with reduced respiration rates allowed 15°C-grownL. saccharina to achieve net photosynthesis and light-saturated photosynthesis at a lower photon fluence rate (PFR) than 5°C plants when both groups were assayed at the same temperature (15°C). The photon fluence rates necessary to reach the compensation point and achieve light-saturated photosynthesis (I _c andI _k , respectively) increased with increasing incubation temperature inL. saccharina grown at both 5 and 15°C. However, acclimation responses to growth temperature compensated for the short-term effect of temperature onI _c andI _k . Consequently, plants grown at 5 and 15°C were able to achieve similar rates of light-limited photosynthesis, and similarI _c andI _k values at their respective growth temperatures. These responses are undoubtedly important for perennial seaweeds such asL. saccharina, which frequently grow in light-limited habitats and experience pronounced seasonal changes in water temperature.Please address all correspondence and requests for reprints to I.R. Davison     

Modeling kelp forest distribution and biomass along temperate rocky coastlines

We address the global deficit of data describing kelp forest distribution, relative covers and biomass by testing the ability of species distribution models to predict these attributes at locations where data are currently limited. We integrated biological ground truth data with high-resolution environmental datasets to develop generalized additive models that accurately predict the structure of Laminaria forests within the Bay of Morlaix (48°42′42″N, 3°55′40″W). Forest distribution and proportional covers were predicted using water depth, light availability, wave exposure and sediment dynamics. The biomass of individual kelp species was modeled by supplementing these same variables with measures of seafloor slope and benthic position. Biomass predictions for Laminaria digitata and Laminaria hyperborea contrast the physiological tolerances of these species to light and wave exposure gradients. As a direct management output, we produced high-resolution maps (25 m² grids) that closely match independent field data and provide vital information for marine spatial planning.     

Ultraviolet radiation effects on the behavior and recruitment of larvae from the reef coral Porites astreoides

We tested the rarely considered hypothesis that the ultraviolet portion (UVR, 280–400 nm) of the light spectrum affects patterns of recruitment in reef-building corals. The premise for this hypothesis rests in the fact that biologically relevant intensities of UVR penetrate to considerable depths (>24 m) in the clear waters surrounding many coral reefs, and that reef organisms allocate substantial resources to prevent and repair UVR damage. The ability of larvae spawned by the brown morph of the Caribbean coral, Porites astreoides, to detect and avoid UVR was assessed in petri dishes where one-half of the dish was shielded from UVR and the other exposed. Observations made every 30 min between 10:30 and 13:30 h showed significantly higher densities of larvae swimming in regions shielded from UVR. To determine how this behavior affects settlement patterns, larvae collected from P. astreoides adults at 18 m depth were released into chambers deployed at 17 m depth where they were given a choice of three different light regions in which to settle: PAR (PAR=400–700 nm), PAR+UVAR (UVAR=320–400 nm), and PAR+UVAR+UVBR (UVBR=280–320 nm). At the end of the experiment, greater numbers of P. astreoides larvae had settled in the region of the tube where UVR was reduced than would be expected if dispersion were random. To our knowledge, this is the first demonstration in any reef-building coral species that planula larvae can detect UVR and that it affects their choice of a settlement site. These results indicate that the capacity to detect and avoid habitats with biologically damaging levels of UVR may be one factor contributing to the successful recruitment of coral larvae.     

“Free space-photosynthesis” in the algae Fucus virsoides and Laminaria saccharina

This paper deals with the role of bicarbonates in the apparent free space (AFS) and their importance for the photosynthetic process. Both the littoral alga Fucus virsoides (Mediterranean Sea) and the sublittoral alga Laminaria saccharina (Baltic Sea) show the same response. Unloading of Clions and subsequent reloading of HCO₃-ions result in a positive photosynthetic rate in distilled water (free of CO₂). Time of exposure of an algal thallus to bicarbonate solutions, as well as the concentration of these solutions, influences AFS-assimilation of the ions.     

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.