Environmental characteristics,field water relations,and photosynthetic responses of C4 Hawaiian Euphorbia species from contrasting habitats

Summary Four endemic Hawaiian Euphorbia species range in habitat from open arid coastal strand to shaded mesic forest and in growth-form from small prostrate shrubs to trees. As shown in the present study, these large differences in habitat and growth-form are paralleled by equally large differences in maximal photosynthetic rate (13.7 to 37.1 mol CO₂ m^-2s^-1), dark respiration rate (0.7 to 4.1 mol CO₂ m^-2s^-1), light level for saturation of photosynthesis (0.9 to over 2.0 mmol m^-2s^-1), light compensation point (0.01 to 0.11 mmol m^-2s^-1), leaf conductance to CO₂ (1.7 to 4.9 mm s^-1), and mesophyll conductance to CO₂ (3.7 to 8.5 mm s^-1). A principal consequence of this differentiation is that the capacity for photosynthesis at high light levels is higher in open site species, such as E. celastroides and E. degeneri, and at low light levels is higher in shade species, such as E. forbesii. E. hillebrandii, a species from intermediate semiopen habitats, exhibits an intermediate photosynthetic capacity at both high and low light levels. Despite this remarkable diversity, all four species exhibit the distinguishing physiological features of C₄ photosynthesis.     

Photosynthesis and dark respiration in Antarctic mosses—an initial comparative study

Rates of dark respiration (DR), gross photosynthesis (GPS) and net photosynthesis (NPS) were investigated for 14 species of moss from a maritime Antarctic locality. The rates found were similar to those reported in studies of temperate, alpine and Arctic species, indicating no physiological specialisation to the Antarctic environment. There was no relationship between the habitat occupied by a species (hydric, mesic, xeric) and physiological measures. There was, however, a loose correlation between NPS and a species' ecology or reproductive behaviour in the maritime Antarctic — species with high NPS being either colonists or those that show high and regular investment in sporophyte production.     

RATES OF PHOTOSYNTHESIS AND RESPIRATION OF THE MOSS BRYUM SANDBERGII AS INFLUENCED BY LIGHT INTENSITY AND TEMPERATURE

Using differential respirometry and air enriched to 3% CO₂ (v/v), the rates of photosynthesis and dark respiration of the moss Bryum sandbergii were measured as influenced by temperature and light intensity. The optimal temperature for net (apparent) photosynthesis was between 24 to 30 C; however, the photosynthesis/respiration ratio was about 11 to 27 between 4 to 24 C and dropped to lower values at 34 C., which indicates a wide temperature tolerance for this moss in short-term experiments. The maximum temperature for photosynthesis was about 41 C and the minimum was below –5 C. At 20 C light saturation was approached at 6.2 mw cm^–2 (ca. 700 ft-c) but not completely reached at 12 mw cm^-2; the light compensation point was estimated to be 0.4 mw cm^-2 (ca. 40 ft-c). At 4 C light saturation and the compensation point were at lower levels and apparently solarization occurred at 12 mw cm^-2. Light intensity had little or no apparent effect on dark respiration. However, respiration increased with temperature over various ranges extending from –5 to 39 C with temperature quotients of about 2.5 to 1.2. The significance of these characteristics is discussed with respect to the ecological relationships of the species.     

Response patterns of net photosynthesis to moisture of mosses in xeric habitats

The response patterns of net photosynthesis to moisture level of mosses in xeric habitats were compared with those in mesic habitats, in order to determine whether the former species are better adapted to the xeric condition with regard to carbon gain. Moss species examined wereRhacomitrium lanuginosum andR. barbuloides in xeric open habitats andDicranum japonicum, Hypnum plicatulum, Ptilium crista-castrensis, Pleurozium schreberi andHylocomium splendens in mesic habitats on the coniferous forest floor in the upper subalpine zone of Mt. Fuji. Three additional xerophytic species collected at other localities,Ptychomitrium polyphylloides, Grimmia pilifera andHedwigia ciliata, were also examined. Five species in the xeric habitats showed an optimum range of moisture level for net photosynthesis, 2 to 3g·g⁻¹. On the other hand, species in the forest showed a wider optimum range, 3 to 8g·g⁻¹. Net photosynthetic rate at the moisture level of 0.5g·g⁻¹ was positive in xerophytic mosses, but negative in most forest mosses. Moisture levels where external capillary water disappeared and drop of water potential began was determined by blotting water-saturated shoots with membrane filters. These moisture levels were low in the xerophytic mosses and high in the forest mosses, although there were some exceptions. It was concluded that mosses in xeric habitats are better adapted for the efficient use of water for photosynthesis than those in mesic habitats.     

Phenotypic plasticity of desiccation resistance in Glossina puparia: are there ecotype constraints on acclimation responses?

Phenotypic plasticity allows organisms to cope with environmental variation and may aid in the evolution of novel traits. However, whether phenotypic plasticity is beneficial, or if acclimation responses might be constrained to particular ecotypes is generally poorly explored. Here we test the beneficial acclimation hypothesis (BAH) and its alternatives for desiccation resistance to atmospheric moisture in mesic‐ and xeric‐adapted Glossina species. Highly significant interactions among acclimation and test humidity were detected for water loss rates indicative of significant phenotypic plasticity. Ordered‐factor anova was unable to reject predictions of the ‘drier is better’ acclimation hypothesis in xeric Glossina morsitans and mesic G. austeni. Evidence for the ‘deleterious acclimation hypothesis’ was found for mesic G. palpalis as expected from the moist habitats it typically occupies. By contrast, support for the ‘optimal acclimation hypothesis’ was found in xeric G. pallidipes. Little support for BAH was obtained in the present study, although other hypotheses, which might enhance fitness within the environments these species are typically exposed to, were supported. However, acclimation responses were not necessarily constrained to xeric/mesic ecotypes which might be expected if adaptation to a particular environment arose as a trade‐off between plastic responses and living in a particular habitat. These results highlight the complexity of acclimation responses and suggest an important role for phenotypic plasticity in moderating environmental effects on evolutionary fitness in Glossina.     

Metabolic recovery of continental antarctic cryptogams after winter

The activation of metabolism after the winter period was investigated in several mosses and lichens in continental Antarctica. Thalli that were still in their over-wintering inactive state in early spring were sprayed artificially and the time-dependent activation of photosystem II (PSII), carbon fixation and respiration was determined using gas exchange and chlorophyll a fluorescence techniques. The investigated lichens recovered PSII activity almost completely within the first few minutes and gross photosynthesis was fully reactivated within a few hours. In contrast, photosynthesis took much longer to recover in mosses, which could indicate a general difference between the green-algal symbionts in lichens and moss chloroplasts. Only small and quickly reversible increased rates of respiration were observed for the foliose lichen Umbilicaria aprina from a more xeric habitat. In contrast, species occurring near persistent meltwater, such as the moss Bryum subrotundifolium and the lichen Physcia caesia, had highly increased respiration rates that were maintained for several days after activation. Calculation of the carbon balances indicated that the activation pattern strongly dictated the length of time before a carbon gain was achieved. It appears that the differences in recovery reflect the water relations of the main growth period in summer.     

Reactivation of photosynthesis in the photoinhibited green alga Chlamydomonas reinhardtii: Role of dark respiration and of light

Effect of quality, quantity and minimum duration of light on the process of recovery was investigated in the photoinhibited cells of the green alga Chlamydomonas reinhardtii. Complete and rapid reactivation of photosynthesis took place in diffuse white light of 25 mol m^–2 s^–1. The recovery was partial (< 10%) in the dark. Far red (725 nm), red (660 nm) and blue light (480 nm) in the range of 10 to 75 mol m^–2 s^–1 did not enhance the process of reactivation. Photoinhibited cells incubated in dark for 15 min when exposed for 5 min to diffuse light (25 mol m^–2 s^–1) showed complete reactivation. Even exposure of 15 min dark incubated photoinhibited cells to photoinhibitory light (2500 mol m^–2 s^–1) for 5 s fully regained the photosynthesis. The study indicated a very precise and triggering effect of light in the process of reactivation. The dark respiratory inhibitor KCN and uncouplers FCCP and CCCP increased the susceptibility of C. reinhardtii to photoinhibition and also prevented photoinhibited cells to reactivate fully even after longer period of incubation under suitable reactivating conditions. Of the various possibilities envisaged to assign the role of dark respiration in recovery process, supply of ATP by mitochondrial respiration appeared sound and pertinent.Abbreviations CCCP- carbonyl cyanide m-chlorophenylhydrazone - D1- 32 kDa protein of PS II reaction center - FCCP- carbonyl cyanide p-(trifluoromethoxy)phenylhydrazone - KCN- potassium cyanide - PBQ- phenyl-p-benzoquinone - PFD- photon flux density - SHAM- salicylhydroxamic acid NBRI Research Publication No. 431.     

Foliar δ13C within a temperate deciduous forest: spatial,temporal, and species sources of variation

Summary Foliar ¹³C-abundance (¹³C) was analyzed in the dominant trees of a temperate deciduous forest in east Tennessee (Walker Branch Watershed) to investigate the variation in foliar ¹³C as a function of time (within-year and between years), space (canopy height, watershed topography and habitat) and species (deciduous and coniferous taxa). Various hypotheses were tested by analyzing (i) samples collected from the field during the growing season and (ii) foliar tissues maintained in an archived collection. The ¹³C-value for leaves from the tops of trees was 2 to 3%. more positive than for leaves sampled at lower heights in the canopy. Quercus prinus leaves sampled just prior to autumn leaf fall had significantly more negative ¹³C-values than those sampled during midsummer. On the more xeric ridges, needles of Pinus spp. had more positive ¹³C-values than leaves from deciduous species. Foliar ¹³C-values differed significantly as a function of topography. Deciduous leaves from xeric sites (ridges and slopes) had more positive ¹³C-values than those from mesic (riparian and cove) environments. On the more xeric sites, foliar ¹³C was significantly more positive in 1988 (a dry year) relative to that in 1989 (a year with above-normal precipitation). In contrast, leaf ¹³C in trees from mesic valley bottoms did not differ significantly among years with disparate precipitation. Patterns in foliar ¹³C indicated a higher ratio of net CO₂ assimilation to transpiration (A/E) for trees in more xeric versus mesic habitats, and for trees in xeric habitats during years of drought versus years of normal precipitation. However, A/E (units of mmol CO₂ fixed/mol H₂O transpired) calculated on the basis of ¹³C-values for leaves from the more xeric sites was higher in a wet year (6.6±1.2) versus a dry year (3.4±0.4). This difference was attributed to higher transpiration (and therefore lower A/E) in the year with lower relative humidity and higher average daily temperature. The calculated A/E values for the forest in 1988–89, based on ¹³C, were within ±55% of estimates made over a 17 day period at this site in 1984 using micrometeorological methods.     

Photoinhibition as a control on photosynthesis and production of Sphagnum mosses

The effect of high light intensity on photosynthesis and growth of Sphagnum moss species from Alaskan arctic tundra was studied under field and laboratory conditions. Field experiments consisted of experimental shading of mosses at sites normally exposed to full ambient irradiance, and removal of the vascular plant canopy from above mosses in tundra water track habitats. Moss growth was then monitored in the experimental plots and in adjacent control areas for 50 days from late June to early August 1988. In shaded plots total moss growth was 2–3 times higher than that measured in control plots, while significant reductions in moss growth were found in canopy removal plots. The possibility that photoinhibition of photosynthesis might occur under high-light conditions and affect growth was studied under controlled laboratory conditions with mosses collected from the arctic study site, as well as from a temperate location in the Sierra Nevada, California. After 2 days of high-light treatment (800 mol photons m^–2 s^–1) in a controlled environmental chamber, moss photosynthetic capacity was significantly lowered in both arctic and temperate samples, and did not recover during the 14-day experimental period. The observed decrease in photosynthetic capacity was correlated (r ²=0.735, P<0.001) with a decrease in the ratio of variable to maximum chlorophyll fluorescence (F _v/F _m) in arctic and temperate mosses. This relationship indicates photoinhibition of photosynthesis in both arctic and temperate mosses at even moderately high light intensities. It is suggested that susceptibility to photoinhibition and failure to photoacclimate to higher light intensities in Sphagnum spp. may be related to low tissue nitrogen levels in these exclusively ombrotrophic plants. Photoinhibition of photosynthesis leading to lowered annual carbon gain in Sphagnum mosses may be an important factor affecting CO₂ flux at the ecosystem level, given the abundance of these plants in Alaskan tussock tundra.     

CO2 environment,microclimate and photosynthetic characteristics of the moss Hylocomium splendens in a subarctic habitat

Summary In order to document the natural CO₂ environment of the moss Hylocomium splendens, and ascertain whether or not the moss was adapted to this, and its interactions with other microenvironmental factors, two studies were carried out. Firstly, the seasonal variations of CO₂ concentration, photosynthetically active radiation (PAR), tissue water content and temperature were measured in the natural microenvironment of H. splendens in a subarctic forest during the summer period (July–September). Secondly, the photosynthetic responses of the species to controlled CO₂ concentrations, PAR, temperature, and hydration were measured in the laboratory. CO₂ concentrations around the upper parts of the plant, when PAR was above the compensation point (30 mol m^–2 s^–1), were mostly between 400 and 450 ppm. They occasionally increased up to 1143 ppm for short periods. PAR flux densities below saturating light levels for photosynthesis (100 mol m^–2 s^–1), occurred during 65% (July), 76% (August) and 96% (September) of the hours of the summer period. The temperature optimum of photosynthesis was 20° C: this temperature coincided with PAR above the compensation point during 5%, 6% and 0% of the time in July, August and September, respectively. Optimal hydration of tissues was infrequent. Hence PAR, temperature and water limit CO₂ uptake for most of the growing season. Our data suggest that the higher than normal ambient CO₂ concentration in the immediate environment of the plant counteracts some of the limitations in PAR supply that it experiences in its habitat. This species already experiences concentrations of atmospheric CO₂ predicted to occur over the next 50 years.     

In situ CO2 gas-exchange in fruits of a tropical tree,Durio zibethinus Murray

We examined the in situ CO₂ gas-exchange of fruits of a tropical tree, Durio zibethinus Murray, growing in an experimental field station of the Universiti Pertanian Malaysia. Day and night dark respiration rates were exponentially related to air temperature. The temperature dependent dark respiration rate showed a clockwise loop as time progressed from morning to night, and the rate was higher in the daytime than at night. The gross photosynthetic rate was estimated by summing the rates of daytime dark respiration and net photosynthesis. Photosynthetic CO₂ refixation, which is defined as the ratio of gross photosynthetic rate to dark respiration rate in the daytime, ranged between 15 and 45%. The photosynthetic CO₂ refixation increased rapidly as the temperature increased in the lower range of air temperature T _c (T _c <28.5 °C), while it decreased gradually as the temperature increased in the higher range (T _c 28.5 °C). Light dependence of photosynthetic CO₂ refixation was approximated by a hyperbolic formula, where light saturation was achieved at 100 mol m^–2 s^–1 and the asymptotic CO₂ refixation was determined to be 37.4%. The estimated gross photosynthesis and dark respiration per day were 1.15 and 4.90 g CO₂ fruit^–1, respectively. Thus the CO₂ refixation reduced the respiration loss per day by 23%. The effect of fruit size on night respiration rate satisfied a power function, where the exponent was larger than unity.     

Elevated CO2 and drought alter tissue water relations of birch (Betula populifolia Marsh.) seedlings

The effect of increasing atmospheric CO₂ concentrations on tissue water relations was examined in Betula populifolia, a common pioneer tree species of the northeastern U.S. deciduous forests. Components of tissue water relations were estimated from pressure volume curves of tree seedlings grown in either ambient (350 l l^–1) or elevated CO₂ (700 l l^–1), and both mesic and xeric water regimes. Both CO₂ and water treatment had significant effects on osmotic potential at full hydration, apoplasmic fractions, and tissue elastic moduli. Under xeric conditions and ambient CO₂ concentrations, plants showed a decrease in osmotic potentials of 0.15 MPa and an increase in tissue elastic moduli at full hydration of 1.5 MPa. The decrease in elasticity may enable plants to improve the soil-plant water potential gradient given a small change in water content, while lower osmotic potentials shift the zero turgor loss point to lower water potentials. Under elevated CO₂, plants in xeric conditions had osmotic potentials 0.2 MPa lower than mesic plants and decreased elastic moduli at full hydration. The increase in tissue elasticity at elevated CO₂ enabled the xeric plants to maintain positive turgor pressures at lower water potentials and tissue water contents. Surprisingly, the elevated CO₂ plants under mesic conditions had the most inelastic tissues. We propose that this inelasticity may enable plants to generate a favorable water potential gradient from the soil to the plant despite the low stomatal conductances observed under elevated CO₂ conditions.     

Tests of the contribution of acclimation to geographic variation in water loss rates of the West Indian lizard <Emphasis Type="Italic">Anolis cristatellus</Emphasis>

Phenotypic plasticity can contribute to the process of adaptive radiation by facilitating population persistence in novel environments. West Indian Anolis lizards provide a classic example of an adaptive radiation, in which divergence has occurred along two primary ecological axes: structural microhabitat and climate. Adaptive plasticity in limb morphology is hypothesized to have facilitated divergence along the structural niche axis in Anolis, but very little work has explored plasticity in physiological traits. Here, we experimentally ask whether Puerto Rican Anolis cristatellus from mesic and xeric habitats differ in desiccation rates, and whether these lizards exhibit an acclimation response to changes in relative humidity. We first present microclimatic data collected at lizard perch sites that demonstrate that abiotic conditions experienced by lizards differ between mesic and xeric habitat types. In Experiment 1, we measured desiccation rates of lizards from both habitats maintained under identical laboratory conditions. This experiment demonstrated that desiccation rates differ between populations; xeric lizards lose water more slowly than mesic lizards. In Experiment 2, lizards from each habitat were either maintained under the conditions of Experiment 1, or under extremely low relative humidity. Desiccation rates did not differ between lizards from the same habitat maintained under different treatments and xeric lizards maintained lower desiccation rates than mesic lizards within each treatment. Our results demonstrate that A. cristatellus does not exhibit an acclimation response to abrupt changes of hydric conditions, and suggest that tropical Anolis lizards might be unable to exhibit physiological plasticity in desiccation rates in response to varying climatic conditions.     

Photosynthetic metabolism and cell-wall polysaccharide accumulation inGelidium sesquipedale (Clem.) Born. et Thur. under different light qualities

The influence of different light qualities on the photosynthetic rate, dark respiration, intracellular carbon and nitrogen content, and accumulation of photosynthetic pigments and cell-wall polysaccharides during short-term incubation (5 h) of the red algaGelidium sesquipedale was investigated. The same photon irradiance of 50mol m^–2 s^–2 below the light saturation point of photosynthesis was applied in each case. Blue light stimulated photosynthesis, dark respiration and the accumulation of chlorophyll and biliproteins, phycoerythrin in particular. The accumulation of internal carbon and nitrogen was greater under blue light than under the other light qualities. In contrast, the percentage of cell-wall polysaccharides was higher in red light. The content of cell-wall polysaccharides decreased during the time of incubation in all light treatments except in red light. The action of a non-photosynthetic photoreceptor in the control of cell-wall polysaccharide synthesis is suggested because the accumulation of cell-wall polysaccharides was not correlated with net photosynthesis in contrast to what occurred with carbon, chlorophyll and phycoerythrin accumulation.     

Carbon dioxide exchange characteristics of C4 Hawaiian Euphorbia species native to diverse habitats

Summary The characteristics of the photosynthetic apparatus of 11 Hawaiian Euphorbia species, all of which possess C₄ photosynthesis but range from arid habitat, drought-deciduous shrubs to mesic or wet forest evergreen trees and shrubs, were investigated under uniform greenhouse conditions. Nine species exhibited CO₂ response curves typical of C₄ plants, but differed markedly in photosynthetic capacity. Light-saturated CO₂ uptake rates ranged from 48 to 52 mol m^-2 s^-1 in arid habitat species to 18 to 20 mol m^-2 s^-1 in mesic and wet forest species. Two possessed unusual CO₂ response curves in which photosynthesis was not saturated above intercellular CO₂ pressures [p(CO₂)] of 10 to 15 Pa, as typically occurs in C₄ plants.Both leaf (g₁) and mesophyll (g_m) conductances to CO₂ varied widely between species. At an atmospheric p(CO₂) of 32 Pa, g₁ regulated intercellular p(CO₂) at 12–15 Pa in most species, which supported nearly maximum CO₂ uptake rates, but did not result in excessive transpiration. Intercellular p(CO₂) was higher in the two species with unusual CO₂ response curves. This was especially apparent in E. remyi, which is native to a bog habitat. The regulation of g₁ and intercellular p(CO₂) yielded high photosynthetic water use efficiencies (P/E) in the species with typical CO₂ response curves, whereas P/E was much lower in E. remyi.Photosynthetic capacity was closely related to leaf nitrogen content, whereas correlations with leaf morphological characteristics and leaf cell surface area were not significant. Thus, differences in photosynthetic capacity may be determined primarily by investment in the biochemical components of the photosynthetic apparatus rather than by differences in diffusion limitations. The lower photosynthetic capacities in the wet habitat species may reflect the lower light availability. However, other factors, such as reduced nutrient availability, may also be important.     

Field measurement of net photosynthesis of mosses at Langhovde,East Antarctica

Net photosynthesis and dark respiration (CO₂ flux) of Antarctic mosses were measured at Langhovde, East Antarctica, from 9 to 17 January 1988. Moss blocks were taken from communities in the Yukidori Valley (69°14′30″S, 39°46′00″E) at Langhovde. Each block was composed ofCeratodon purpureus andBryum pseudotriquetrum, orB. pseudotriquetrum. The upper part of the block was used to measure net photosynthesis and dark respiration. The net photosynthesis of each sample was measured in the field for one or three days with two infrared CO₂ gas analyzers and an assimilation chamber. The relationships of net photosynthetic rate and dark respiration rate, to the water content of the sample, the intensity of solar radiation and the moss temperature were estimated from the field data. The maximum rate of net photosynthesis was about 4 μmol CO₂ m⁻²s⁻¹ at saturating radiation intensity and at optimum temperature, about 10°C. Environmental features of moss habitats in the Yukidori Valley are discussed in relation to these results.     

The effect of light quality and gibberellic acid (GA3) on photosynthesis and respiration rates of pea seedlings

CO₂ exchange were measured on pea seedlings (Pisum sativum L. var. Bördi) cultivated from seeds imbibed either in water (C-plants) or in gibberellic acid (GA₃) at the concentration of 25 g/1 (GA-plants), and then grown under 17 W/m² blue light (B-plants) or 11 W/m² red light (R-plants).When measured under the same light conditions as during growth the net photosynthesis (APS) rate in B-plants was about twice higher than that in R-plants. Dark respiration (DR) rate was 70% higher in B- than in R-plants. Red light retarded the development of photosynthetic activity, but GA₃ suppressed this effect. The hormone enhanced net photosynthesis and dark respiration to the same extent.When measured under saturating white light net photosynthesis rate of C-plants was also two times higher in B-plants than in R-plants. Growth conditions had only a slight effect on the APS of GA-plants under white light. APS rates of GA-plants grown under red light were higher under white light than those of C-plants, but lower than those of plants grown under blue light.We assume that blue light induced formation of plants that were adapted to higher light intensity: red light had an opposite effect, whereas gibberellic acid induced formation of plants that were adapted to medium light intensity.     

Predicting the impact of climatic warming on the carbon balance of the moss <Emphasis Type="Italic">Sanionia uncinata</Emphasis> on a maritime Antarctic island

The effects of climatic factors, especially those of temperature, on the carbon balance of the moss Sanionia uncinata were examined on King George Island in the maritime Antarctic. Net photosynthesis (P _n ) and dark respiration rates of two colonies (A and B) were measured with a portable infrared gas analyzer. Colony A showed small P _n compared with its dark respiration rates throughout the growing season. Colony B showed much higher net photosynthetic rates, but the dark respiration rates of the two colonies did not differ significantly. Net photosynthetic rate determined at light saturation was almost constant over a wide temperature range, from 5° to 15°C, while the dark respiration was strongly affected by temperature. To assess the impact of warming on the carbon balance of the moss, cumulative carbon gain of the moss was calculated using a simulation model for the main part of the growing season. The results suggest that climatic warming may cause a reduction of carbon gain in some relatively photosynthetically inactive moss colonies. Received: April 13, 2001 / Accepted: November 5, 2001     

Photosynthetic performance of transplanted ecotypes ofEcklonia cava(Laminariales, Phaeophyta)

Young sporophytes of short-stipe ecotype ofEcklonia cavafrom a warmer locality (Tei, Kochi Pref., southern Japan) and those of long-stipe ecotype from a cooler locality (Nabeta, Shizuoka Pref., central Japan) were transplanted in 1995 to artificial reefs immersed at the habitat of long-stipe ecotype in Nabeta Bay, Shizuoka Pref., central Japan. The characteristics of photosynthesis and respiration of bladelets of the transplanted sporophytes of the two ecotypes were compared in winter and summer 1997; the results were assessed per unit area, per unit chlorophyllacontent and per unit dry weight. In photosynthesis-light curves at 10–29 °C, light saturation occurred at 200–400 mol photon m^–2s^–1in sporophytes from both Tei and Nabeta. The maximum photosynthetic rate (P _max) at 10–29 °C and the light-saturation index (I _k) at 25–29 °C in sporophytes from both localities were generally higher in winter than in summer.P _maxat 25–29 °C (per unit area and chlorophylla) were higher in sporophytes from Tei than those from Nabeta in both seasons. The optimum temperature for photosynthesis was 25 °C in winter and 27 °C in summer at high light intensities of 100–400 mol photon m^–2s^–1. However, at lower light intensities of 12.5–50 mol photon m^–2s^–1, it was 20 °C in winter and 25–27 °C in summer for sporophytes from both locations. Dark respiration increased with temperature rise in the range of 10–29 °C in sporophytes from both locations in summer and winter. The sporophytes transplanted from Tei (warmer area) showed higher photosynthetic activities than those from Nabeta (cooler area) at warmer temperatures even under the same environmental conditions. This indicates that these physiological ecotypes have arisen from genetic differentiation.     

Microhabitat,water relations,and photosynthesis of a desert fern,Notholaena parryi

Summary Interrelationships between morphology, microhabitat, water relations, and photosynthesis of a xeric fern, Notholaena parryi D.C. Eat. (Pteridaceae), were examined in the western Colorado desert. In its typical microhabitat rock outcroppings protected N. parryi from direct sunlight and moderated the diurnal variations in air temperature. For example, frond temperature at noon in late winter was 15.3° C, which was 7.3° C cooler than an energy budget simulation predicting frond temperature at an exposed site. The lowest soil water potential leading to daytime stomatal opening was about-1.5 MPa (-15 bars). Rainfall runoff that was channeled to the periphery of the rocks caused ^soil near the fern roots to rise above-1.5 MPa even after light rainfalls, and it remained above-1.5 MPa longer after rainfall than in non-rocky sites.The water potential gradient along the stipe necessary to support the observed rates of transpiration was about-10 MPa m^-1; such a large gradient reflected the small conducting area in the xylem. The water vapor conductance decreased as the frond temperature was raised, an effect that became proportionally greater as the soil dried out. The daytime water-use efficiency (mass CO₂ fixed/mass water transpired) was 0.0058 for a spring day. Individual fronds reached 90% of light saturation for photosynthesis at only 100 Einsteins m^-2 s^-1, a photosynthetically active radiation similar to that from the diffuse sunlight incident on the generally north-facing microhabitat of the fern. Below 50 E m^-2 s^-1 the quantum requirement was 13 Einsteins absorbed/mole CO₂ fixed. The ratio of chlorophyll to P700 was 552, indicating a fairly large photosynthetic unit that is characteristic of plants adapted to shaded habitats. The temperature optimum for net photosynthesis shifted from 13° C in midwinter (mean daily air temperature of 11° C) to 19° C in early fall (air temperature of 23° C).