ENVIRONMENTAL AND PHYSIOLOGICAL FACTORS INFLUENCING THE NATURAL DISTRIBUTION OF EVERGREEN AND DECIDUOUS ERICACEOUS SHRUBS ON NORTHEAST- AND SOUTHWEST-FACING SLOPES OF THE SOUTHERN APPALACHIAN MOUNTAINS. II. WATER RELATIONS

Three species of shrubs (Ericaceae) were found to segregate upon the northeast and southwest slopes of spur ridges on Brush Mountain, in southwestern Virginia. Rhododendron maximum was found only in valleys and lower northeast slopes, Rhododendron periclymenoides = R. nudiflorum) was found on northeast slopes while Kalmia latifolia was most abundant on southwest slopes. Previous vegetation studies indicated that these partially segregated distributions were related to irradiance and water availability. In field studies of water potential, R. periclymenoides had the lowest diurnal leaf water potentials and the largest seasonal variation in midday leaf water potential. Kalmia latifolia had the highest leaf conductance in field and phytotron experiments. Rhododendron maximum had the greatest seasonal osmotic adjustment followed by R. periclymenoides and K. latifolia. In phytotron experiments, the photosynthetic capacity of R. maximum was the most sensitive to water stress followed by R. periclymenoides and K. latifolia. Kalmia latifolia was able to modify its conductance rates to reduce water loss and maintain constant leaf water potential minimizing photosynthetic inhibition. Rhododendron periclymenoides showed extreme luxury spending of water indicated by high conductance and low photosynthesis. The ecophysiological responses to water and irradiance provided an explanation for the distributions of the three species. For example, R. maximum leaves are sensitive to elevated irradiance, and carbon gain is strongly influenced by water stress. Thus, R. maximum will perform best in low irradiance environments with ample water, such as valley sites. Each species had a unique set of adaptations for performing best in their optimum habitat.     

INFLUENCE OF AGE AND MICROCLIMATE ON THE PHOTOCHEMISTRY OF RHODODENDRON MAXIMUM LEAVES II. CHLOROPLAST STRUCTURE AND PHOTOSYNTHETIC LIGHT RESPONSE

The influence of age on chloroplast structure and photosynthetic light response of Rhododendron maximum L. was studied in three different microhabitats. The three microhabitats constituted a gradient of low, intermediate, and high irradiance levels. The most dramatic change in chloroplast structure with increasing age was the proliferation of the number and size of plastoglobuli. The magnitude and age specific rate of chloroplast occlusion by plastoglobuli increased in habitats with higher irradiance. Photosynthetic responses to light differed among the age categories of leaves. Light saturated photosynthesis and quantum yield decreased as leaves aged. However, in high light environments the rate of reduction of quantum yield or light saturated photosynthetic rate was more rapid than in the low light environment. The quantity of plastoglobuli increased in association with reduced light reaction capacity. The presence and abundance of plastoglobuli in R. maximum chloroplasts and their association with reduced photosynthetic performance indicates that the photosynthetic apparatus of the R. maximum chloroplast is sensitive to photodestruction by high irradiance: commonly a winter phenomenon in these environments.     

QUANTITATIVE PHENOLOGY AND LEAF SURVIVORSHIP OF RHODODENDRON MAXIMUM IN CONTRASTING IRRADIANCE ENVIRONMENTS OF THE SOUTHERN APPALACHIAN MOUNTAINS

Seasonal plant growth patterns were compared for Rhododendron maximum L. in two contrasting subcanopy environments. The two subcanopy, above ground environments differed only in their quantity of irradiance by virtue of the relative dominance of evergreen or deciduous trees in the canopy. A third site had no canopy influence. Overall growth (shoot elongation, woody increment, leaf production) was maximized under the open (BMO) and deciduous dominated canopy (PCD). The leaf pool was significantly smaller under the evergreen dominated canopy (PCE) but average leaf area per leaf was slightly larger at PCE. Individual age-specific leaf cohorts, identified from shoot morphology, indicated increased leaf survivorship with a decreased irradiance environment. Leaf production was synchronous and rapid (1 week), followed by three weeks of leaf expansion, which created the even-aged leaf cohorts. Wood growth (diameter increment), in contrast, continued through the beginning of the winter. Reproductive effort increased with increasing irradiance environment. Significant variation in growth was observed between canopy shoot types at all three research sites. The significance of these phenological patterns is discussed in view of the variable subcanopy environment of southwestern Virginia.     

Leaf phenology and seasonal variation of photosynthesis of invasive <Emphasis Type="Italic">Berberis thunbergii</Emphasis> (Japanese barberry) and two co-occurring native understory shrubs in a northeastern United States deciduous forest

Early leafing and extended leaf longevity can be important mechanisms for the invasion of the forest understory. We compared the leaf phenology and photosynthetic characteristics of Berberis thunbergii, an early leafing invasive shrub, and two co-occurring native species, evergreen Kalmia latifolia and late leafing Vaccinium corymbosum, throughout the 2004 growing season. Berberis thunbergii leafed out 1 month earlier than V. corymbosum and approximately 2 weeks prior to the overstory trees. The photosynthetic capacity [characterized by the maximum carboxylation rate of Rubisco (V _cmax) and the RuBP regeneration capacity mediated by the maximum electron transport rate (J _max)] of B. thunbergii was highest in the spring open canopy, and declined with canopy closure. The 2003 overwintering leaves of K. latifolia displayed high V _cmax and J _max in spring 2004. In new leaves of K. latifolia produced in 2004, the photosynthetic capacity gradually increased to a peak in mid-September, and reduced in late November. V. corymbosum, by contrast, maintained low V _cmax and J _max throughout the growing season. In B. thunbergii, light acclimation was mediated by adjustment in both leaf mass per unit area and leaf N on a mass basis, but this adjustment was weaker or absent in K. latifolia and V. corymbosum. These results indicated that B. thunbergii utilized high irradiance in the spring while K. latifolia took advantage of high irradiance in the fall and the following spring. By contrast, V. corymbosum generally did not experience a high irradiance environment and was adapted to the low irradiance understory. The apparent success of B. thunbergii therefore, appeared related to a high spring C subsidy and subsequent acclimation to varying irradiance through active N reallocation and leaf morphological modifications.     

SEASONAL PRIMARY PRODUCTION OF UNDERSTORY RHODOPHYTA IN AN UPWELLING SYSTEM1

The in situ primary production of three common under-story members of the Rhodophyta in South African west coast kelp beds was determined monthly for a year using dissolved oxygen techniques. Strong seasonal patterns of photosynthesis and respiration were evident in all three species. Net photosynthesis of all three species was greatest in spring (October) and lowest in winter (June). Increasing photosynthesis in late winter coincided with increasing ambient irradiance and photoperiod, whereas decreasing photosynthesis in summer was not explained by changes in the environmental parameters measured. We suggest that this may he due to an innate pattern related to some other seasonal plant activity such as reproduction. Seasonal P_max and I_k values reveal that the obligate understory species, B. prolifera and E. obtusa, are shade-adapted whereas G. radula, a low intertidal and shallow subtidal dominant, is sun-adapted. Low C: X ratios consistent with a high nutrient environment and high rates of productivity were found in all three species. Net photosynthesis to respiration (Pn:R) ratios were fairly constant for B. prolifera and E. obtusa, implying that then photosynthetic processes were governed more by seasonal variations in irradiance than by instantaneous light availability. The Pn: R ratio of G. radula was variable, suggesting that this species is more responsive to rapid fluctuations in irradiance and may therefore be adapted for rapid growth during periods of high irradiance.     

Seasonal and diurnal leaf movements of Rhododendron maximum L. in contrasting irradiance environments

Summary Leaf orientation (azimuth and angle) and leaf curling were measured seasonally and diurnally on Rhododendron maximum L. under an evergreen and a deciduous canopy. The microclimatic conditions under the evergreen canopy (mixed pine and hemlock) were characterized by lower irradiance but similar temperature, and vapor pressure deficit (vpd) to that under the deciduous canopy (mixed oak and maple). Under both canopies irradiance was more intense during winter months.On a seasonal basis leaf angle was closer to horizontal under the evergreen canopy but there was no difference between leaf curling in the two sites. Stomatal conductance was higher under the deciduous canopy but stomata were closed in the winter (following canopy abscission) under the evergreen and deciduous canopies even during warm winter days. Leaf water potentials were lower in the winter and Rhododendron maximum had higher leaf water potentials under the evergreen canopy.Significant association between mean leaf angle and curling index were found above a mean leaf angle of 70°. Leaf curling was highly associated with leaf temperature where 0° C was a critical value stimulating leaf curling. Leaf angle was linearly related to leaf temperatures above 0° C although this relationship was different under the two canopy types as a result of differing irradiance or differing water potential.     

The impact of subcanopy light environment on the hydraulic vulnerability of Rhododendron maximum to freeze-thaw cycles and drought

The vulnerability of xylem to embolism development in Rhododendron maximum L., an evergreen diffuse-porous shrub, was investigated in relation to the frequency of winter freeze–thaw cycles in high and low light sites of the Eastern US. Though the frequency of freeze–thaw cycles during the winter was lower in North Carolina than in Virginia, the hydraulic conductivity of 3-year-old branches was reduced by up to 60% by winter embolism development in North Carolina compared to less than 30% in Virginia. Generally, small vessel diameters and volumes were associated with a significant resistance to embolism formation resulting from repeated freeze–thaws of xylem sap. In stems grown in high light sites (gaps), larger vessel volumes, and greater diameter growth of stems were associated with a significantly higher degree of freeze–thaw embolism development than in those grown in the low light sites. Thus, the growth patterns of R. maximum stems, under conditions of higher light availability, rendered them more susceptible to freeze–thaw-induced embolisms. Vulnerability to drought-induced embolism in stems was not affected by light environment. Rhododendron maximum was relatively sensitive to drought-induced embolism because 50% loss of hydraulic conductivity occurred at a water potential of -2.2 MPa. The distribution and gas exchange of R. maximum are constrained by the dual effects of freeze-thaw cycles and drought on vascular function.     

Light environment under Rhododendron maximum thickets and estimated carbon gain of regenerating forest tree seedlings

Canopy tree recruitment is inhibited by evergreen shrubs in many forests. In the southern Appalachian mountains of the USA, thickets of Rhododendron maximum L. restrict dominant canopy tree seedling survival and persistence. Using R. maximum as a model system, we examined available light under the thickets and the photosynthetic responses of seedlings of canopy tree species. We tested the hypothesis that the additional shading from under R. maximum drives carbon gain in seedlings below the threshold for growth and survival. A reduction in light under the thicket was found where canopy openness (derived from canopy photographs) under R. maximum was half the amount measured in forest without R. maximum. R.␣maximum also reduced direct radiation by 50% and diffuse radiation by 12–29% compared to forest without the shrub layer. Mean mid-day PPFD (photosynthetically active photon flux density between 1000 and 1400 h) under R. maximum (obtained from quantum sensors) was below 10 mol m⁻² s⁻¹ on both clear and overcast days and the amount of sunflecks greater than 10 mol m⁻² s⁻¹ PPFD was only 0–20 min per day. In contrast, forest without R. maximum received a mean PPFD of 18–25 mol m⁻² s⁻¹ on clear days and a cumulative sunfleck duration of 100–220 min per day in all sky conditions. Consistent with light availability between the sites, daily carbon gain in Quercus rubra L. seedlings was lower in forest with R. maximum compared to forest where the shrub was absent. The presence of the shrub layer also significantly suppressed average mid-day photosynthesis of both Q. rubra and Prunus serotina Ehrt. seedlings on 8 out of 11 measurement dates. However, parameters derived from light response curves between seedlings growing in forest sites with or without a thicket of R. maximum was significantly different only in A _max (maximum photosynthetic rate), indicating a lack of further acclimation to the deeper shade under R. maximum. While the additional shade cast by R. maximum is sufficient to prevent the regeneration of tree seedlings under this shrub, there was sufficient heterogeneity in light under the thicket to imply that deep shade only partially explains the complete inhibition of regenerating canopy trees under R. maximum.     

PHOTOACCLIMATION OF PHOTOSYNTHESIS IRRADIANCE RESPONSE CURVES AND PHOTOSYNTHETIC PIGMENTS IN MICROALGAE AND CYANOBACTERIA1

The photosynthesis‐irradiance response (PE) curve, in which mass‐specific photosynthetic rates are plotted versus irradiance, is commonly used to characterize photoacclimation. The interpretation of PE curves depends critically on the currency in which mass is expressed. Normalizing the light‐limited rate to chl a yields the chl a‐specific initial slope (α^chl). This is proportional to the light absorption coefficient (a^chl), the proportionality factor being the photon efficiency of photosynthesis (φ_m). Thus, α^chl is the product of a^chl and φ_m. In microalgae α^chl typically shows little (<20%) phenotypic variability because declines of φ_m under conditions of high‐light stress are accompanied by increases of a^chl. The variation of α^chl among species is dominated by changes in a^chl due to differences in pigment complement and pigment packaging. In contrast to the microalgae, α^chl declines as irradiance increases in the cyanobacteria where phycobiliproteins dominate light absorption because of plasticity in the phycobiliprotein:chl a ratio. By definition, light‐saturated photosynthesis (P_m) is limited by a factor other than the rate of light absorption. Normalizing P_m to organic carbon concentration to obtain P_m^C allows a direct comparison with growth rates. Within species, P_m^C is independent of growth irradiance. Among species, P_m^C covaries with the resource‐saturated growth rate. The chl a:C ratio is a key physiological variable because the appropriate currencies for normalizing light‐limited and light‐saturated photosynthetic rates are, respectively, chl a and carbon. Typically, chl a:C is reduced to about 40% of its maximum value at an irradiance that supports 50% of the species‐specific maximum growth rate and light‐harvesting accessory pigments show similar or greater declines. In the steady state, this down‐regulation of pigment content prevents microalgae and cyanobacteria from maximizing photosynthetic rates throughout the light‐limited region for growth. The reason for down‐regulation of light harvesting, and therefore loss of potential photosynthetic gain at moderately limiting irradiances, is unknown. However, it is clear that maximizing the rate of photosynthetic carbon assimilation is not the only criterion governing photoacclimation.     

EFFECTS OF CANOPY POSITION AND IRRADIANCE ON THE LEAF PHYSIOLOGY AND MORPHOLOGY OF PENTACLETHRA MACROLOBA (MIMOSACEAE)

Pentaclethra macroloba (Willd.) Kuntze (Mimosaceae) is a dominant late-successional tree species in the Atlantic lowland forests of Costa Rica. Leaves of P. macroloba from three heights in the forest canopy were compared with leaves of seedlings grown in controlled environment chambers under four different irradiance levels. Changes in leaf characteristics along the canopy gradient paralleled changes resulting from the light gradient under controlled conditions. The effect of light or canopy position on light-saturated photosynthesis was small, with maximum photosynthesis increasing from 5 to 6.5 μmol m^−-2 s^−-1 from understory to canopy. Both chamber grown and field leaves showed large adjustments in photosynthetic efficiency at low light via reductions in dark respiration rates and increases in apparent quantum yields. Light saturation of all leaves occurred at or below 500 μmol m^−-2 s^−-1. Leaf thickness, specific leaf weight, and stomatal density increased to a greater extent than saturated photosynthesis with higher irradiance during growth or height in the canopy. As a result, there was a poor correspondence between leaf thickness and light-saturated photosynthesis on an area basis. It is concluded that Pentaclethra macroloba possesses the characteristics of a typical shade-tolerant species.     

THE EFFECTS OF TEMPERATURE ON THE PHOTOSYNTHETIC PARAMETERS AND RECOVERY OF TWO TEMPERATE BENTHIC MICROALGAE,AMPHORA CF. COFFEAEFORMIS AND COCCONEIS CF. SUBLITTORALIS (BACILLARIOPHYCEAE)1

Temperature and irradiance are the most important factors affecting marine benthic microalgal photosynthetic rates in temperate intertidal areas. Two temperate benthic diatoms species, Amphora cf. coffeaeformis (C. Agardh) Kütz. and Cocconeis cf. sublittoralis Hendey, were investigated to determine how their photosynthesis responded to temperatures ranging from 5°C to 50°C after short‐term exposure (1 h) to a range of irradiance levels (0, 500, and 1,100 μmol photons · m^?2 · s^?1). Significant differences were observed between the temperature responses of maximum relative electron transport rate (rETRmax), photoacclimation index (E_k), photosynthetic efficiency (α), and effective quantum yield (ΔF/F_m’) in both species. A. coffeaeformis had a greater tolerance to higher temperatures than C. sublittoralis, with nonphotochemical quenching (NPQ) activated at temperatures of 45°C and 50°C. C. sublittoralis, however, demonstrated a more rapid rate of recovery at ambient temperatures. Temperatures between 10°C and 20°C were determined to be optimal for photosynthesis for both species. High temperatures and irradiances caused a greater decrease in ΔF/F_m’ values. These results suggest that the effects of temperature are species specific and that short‐term exposure to adverse temperature slows the recovery process, which subsequently leads to photoinhibition.     

THE EFFECT OF GROWTH IRRADIANCE ON THE COUPLING OF CARBON AND NITROGEN METABOLISM IN CHAETOMORPHA LINUM (CHLOROPHYTA)

The influence of growth irradiance on the non-steady-state relationship between photosynthesis and tissue carbon (C) and nitrogen (N) pools in Chaetomorpha linum (Muller) Kutzing in response to abrupt changes in external nitrogen (N) availability was determined in laboratory experiments. For a given thallus N content, algae acclimated to low irradiance consistently had a higher rate of light-saturated photosynthesis (P_max normalized to dry weight) than algae acclimated to saturating irradiance; for both treatments, P_max was correlated to thallus N. Both P_max and the photosynthetic efficiency (α_dw) were correlated in C. linum grown at either saturating or limiting irradiance over the range of experimental conditions, indicating that variations in electron transport were coupled to variations in C-fixation capacity despite the large range of tissue N content from 1.1% to 4.8%. Optimizing both α and P_max and thereby acclimating to an intermediate light level may be a general characteristic of thin-structured opportunistic algae that confers a competitive advantage in estuarine environments in which both light and nutrient conditions are highly variable. Nitrogen-saturated algae had the same photosynthesis–irradiance relationship regardless of light level. When deprived of an external N supply, photosynthetic rates did not change in C. linum acclimated to low irradiance despite a two-fold decrease in tissue N content, suggesting that the active pools of chlorophyll and Rubisco remained constant. Both α and P_max decreased immediately and continuously in algae acclimated to high irradiance on removal of the N supply even though tissue N content was relatively high during most of the N-starvation period, indicating a diversion of energy and reductant away from C fixation to support high growth rates. Carbon and nitrogen assimilation were equally balanced in algae in both light treatments throughout the N-saturation and -depletion phases, except when protein synthesis was limited by the depletion of internal N reserves in severely N-starved high-light algae and excess C accumulated as starch stores. This suggests that the ability for short-term adjustment of internal allocation to acquire N andC in almost constant proportions may be especially beneficial to macroalgae living in environments characterized by high variability in light levels and nutrient supply.     

Light acclimation, CO2 response and long-term capacity of underwater photosynthesis in three terrestrial plant species

To characterize underwater photosynthetic performance in some terrestrial plants, we determined (i) underwater light acclimation (ii) underwater photosynthetic response to dissolved CO₂, and (iii) underwater photosynthetic capacity during prolonged submergence in three species that differ in submergence tolerance: Phalaris arundinacea, Rumex crispus (both submergence-tolerant) and Arrhenatherum elatius (submergence-intolerant). None of the species had adjusted to low irradiance after 1 week of submergence. Under non-submerged (control) conditions, only R. crispus displayed shade acclimation. Submergence increased the apparent quantum yield in this species, presumably because of the enhanced CO₂ affinity of the elongated leaves. In control plants of the grass species P. arundinacea and A. elatius, CO₂ affinities were higher than for R. crispus. The underwater photosynthetic capacity of R. crispus increased during 1 month of submergence. In P. arundinacea photosynthesis remained constant during 1 month of submergence at normal irradiance; at low irradiance a reduction in photosynthetic capacity was observed after 2 weeks, although there was no tissue degeneration. In contrast, underwater photosynthesis of the submergence-intolerant species A. elatius collapsed rapidly under both irradiances, and this was accompanied by leaf decay. To describe photosynthesis versus irradiance curves, four models were evaluated. The hyperbolic tangent produced the best goodness-of-fit, whereas the rectangular hyperbola (Michaelis-Menten model) gave relatively poor results.     

INTERACTIVE EFFECTS OF IRRADIANCE AND TEMPERATURE ON THE PHOTOSYNTHETIC PHYSIOLOGY OF THE PENNATE DIATOM PSEUDO‐NITZSCHIA GRANII (BACILLARIOPHYCEAE) FROM THE NORTHEAST SUBARCTIC PACIFIC1

This study examined how light and temperature interact to influence growth rates, chl a, and photosynthetic efficiency of the oceanic pennate diatom Pseudo‐nitzschia granii Hasle, isolated from the northeast subarctic Pacific. Growth rates were modulated by both light and temperature, although for each irradiance tested, the growth rate was always the greatest at ～14°C. Chl a per cell was affected primarily by temperature, except at the maximum chl a per cell (at 10°C) where the effects of light were noticeable. At both ends of the temperature gradient, cells displayed evidence of chlorosis even at low light intensities. Chl fluorescence data suggested that cells at 8°C were significantly more efficient in their photosynthetic processes than cells at 20°C, despite having comparable concentrations of chl. Cells at low temperature showed photosynthetic characteristics similar to high‐irradiance‐adapted cells. The decline of growth rates beyond the optimum growth temperature coincided with the cell's inability to accumulate chl in response to increasing temperature. The decline in photosynthetic ability at 20°C was likely due to a combination of high‐temperature stress on cellular membranes and a decline in chl. Our results highlight the important interactions between light and temperature and the need to incorporate these interactions into the development of phytoplankton models for the subarctic Pacific.     

Photosynthetic limitation of several representative subalpine species in the Catalan Pyrenees in summer

Information on the photosynthetic process and its limitations is essential in order to predict both the capacity of species to adapt to conditions associated with climate change and the likely changes in plant communities. Considering that high‐mountain species are especially sensitive, three species representative of subalpine forests of the Central Catalan Pyrenees: mountain pine (Pinus uncinata Mill.), birch (Betula pendula Roth) and rhododendron (Rhododendron ferrugineum L.) were studied under conditions associated with climate change, such as low precipitation, elevated atmospheric [CO₂] and high solar irradiation incident at Earth's surface, in order to detect any photosynthetic limitations. Short‐term high [CO₂] increased photosynthesis rates (A) and water use efficiency (WUE), especially in birch and mountain pine, whereas stomatal conductance (g_s) was not altered in either species. Birch showed photosynthesis limitation through stomatal closure related to low rainfall, which induced photoinhibition and early foliar senescence. Rhododendron was especially affected by high irradiance, showing early photosynthetic saturation in low light, highest chlorophyll content, lowest gas exchange rates and least photoprotection. Mountain pine had the highest A, photosynthetic capacity (A_max) and light‐saturated rates of net CO₂ assimilation (A_sat), which were maintained under reduced precipitation. Furthermore, maximum quantum yield (F_v/F_m), thermal energy dissipation, PRI and SIPI radiometric index, and ascorbate content indicated improved photoprotection with respect to the other two species. However, maximum velocity of carboxylation of RuBisco (V_cmax) indicated that N availability would be the main photosynthetic limitation in this species.     

IMPORTANCE OF MACRO‐ VERSUS MICROSTRUCTURE IN MODULATING LIGHT LEVELS INSIDE CORAL COLONIES1

Adjusting the light exposure and capture of their symbiotic photosynthetic dinoflagellates (genus Symbiodinium Freud.) is central to the success of reef‐building corals (order Scleractinia) across high spatio‐temporal variation in the light environment of coral reefs. We tested the hypothesis that optical properties of tissues in some coral species can provide light management at the tissue scale comparable to light modulation by colony architecture in other species. We compared within‐tissue scalar irradiance in two coral species from the same light habitat but with contrasting colony growth forms: branching Stylophora pistillata and massive Lobophyllia corymbosa. Scalar irradiance at the level of the symbionts (2 mm into the coral tissues) were <10% of ambient irradiance and nearly identical for the two species, despite substantially different light environments at the tissue surface. In S. pistillata, light attenuation (90% relative to ambient) was observed predominantly at the colony level as a result of branch‐to‐branch self‐shading, while in L. corymbosa, near‐complete light attenuation (97% relative to ambient) was occurring due to tissue optical properties. The latter could be explained partly by differences in photosynthetic pigment content in the symbiont cells and pigmentation in the coral host tissue. Our results demonstrate that different strategies of light modulation at colony, polyp, and cellular levels by contrasting morphologies are equally effective in achieving favorable irradiances at the level of coral photosymbionts.     

THE INFLUENCE OF SALINITY AND TEMPERATURE COVARIATION ON THE PHOTOPHYSIOLOGICAL CHARACTERISTICS OF ANTARCTIC SEA ICE MICROALGAE1

The responses of sea ice microalgae to variation in ambient irradiance (0 to 150 μE · m^?2· s^?1), temperature (–6° to + 6° C), and salinity (0 to 100 ppt) were tested to determine whether these variables act independently or in concert to influence rates of microalgal photosynthesis. The photosynthetic efficiency and maximum photosynthetic rate for sea ice microalgae increased as a function of incubation temperature between -6° and + 6° C. Furthermore, photosynthetic efficiency, maximum photosynthetic rate, and quantum yield were greatest at salinities between SO and 50 ppt. In contrast, the mean specific absorption coefficients were lowest near seawater salinities, and the saturating irradiance, I_s, appeared to be inversely proportional to salinity. Results also suggest that the effects of salinity on the growth of sea ice microalgae are independent of those elicited by temperature or light, and that the functional relationship between salinity and light or temperature is multiplicative. This information is essential to the proper formulation of algorithms used to describe algal growth in environments where light, temperature, and salinity are changing simultaneously, such as within sea ice or within the water column at the marginal ice edge zone.     

青冈常绿阔叶林主要植物种叶片的光合特性及其群落学意义

 青冈(Quercus glauca)林乔木层主要树种叶片的净光合速率日进程在春夏季晴天均有明显午休。常绿树种的光合速率在秋季最高,大量换叶期最低,冬季仍有一定的净光合量。落叶树种的光合速率和光合产量低于常绿树种,随着群落的发育其地位将降低。青冈和石栎(Lithocarpus glaber)的光合速率接近,但青冈能够利用较弱的光,因而其高叶量的树冠具有较高的光合总量,以保持其在群落中的优势种地位;石栎主要利用较强的光,其较低叶量的树冠可以维持较高的光合总量,以保证其次优势种地位。灌木、草本层种类光合日进程均为单峰型。灌木种类对光的需求较高。蕨类植物耐荫强,因而在林中能占有稳定的伴生地位。     

MICROENVIRONMENTAL CONTROL OF PHOTOSYNTHESIS AND PHOTOSYNTHESIS-COUPLED RESPIRATION IN AN EPILITHIC CYANOBACTERIAL BIOFILM1

The photosynthetic performance of an epilithic cyano-bacterial biofilm was studied in relation to the in situ light field by the use of combined microsensor measurements of O₂, photosynthesis, and spectral scalar irradiance. The high density of the dominant filamentous cyanobacteria (Oscillatoria sp.) embedded in a matrix of exopolymers and bacteria resulted in a photic zone of < 0.7 mm. At the biofilm surface, the prevailing irradiance and spectral composition were significantly different from the incident light. Multiple scattering led to an intensity maximum for photic light (400–700 nm) of ca. 120% of incident quantum irradiance at the biofilm surface. At the bottom of the euphotic zone in the biofilm, light was attenuated strongly to < 5–10% of the incident surface irradiance. Strong spectral signals from chlorophyll a (440 and 675 nm) and phycobilins (phycoerythrin 540–570 nm, phycocyanin 615–625 nm) were observed as distinct maxima in the scalar irradiance attenuation spectra in the upper 0.0–0.5 mm of the biofilm. The action spectrum for photosynthesis in the cyanobacterial layer revealed peak photosynthetic activity at absorption wavelengths of phycobilins, whereas only low photosynthesis rates were induced by light absorption of carotenoids (450–550 nm). Respiration rates in light- and dark-incubated biofilms were determined using simple flux calculations on measured O₂ concentration profiles and photosynthetic rates. A significantly higher areal O₂ consumption was found in illuminated biofilms than in dark-incubated biofilms. Although photorespiration accounted for part of the increase, the enhanced areal O₂ consumption of illuminated biofilms could also be ascribed to a deeper oxygen penetration in light as well as an enhanced volumetric O₂ respiration in and below the photic zone. Gross photosynthesis was largely unaffected by increasing flow velocities, whereas the O₂ flux out of the photic zone, that is, net photosynthesis, increased with flow velocity. Consequently, the amount of produced O₂ consumed within the biofilm decreased with increasing flow velocity. Our data indicated a close coupling of photosynthesis and respiration in biofilms, where the dissolved inorganic carbon requirement of the photo-synthetic population may largely be covered by the respiration of closely associated populations of heterotrophic bacteria consuming a significant part of the photosynthetically produced oxygen and organic carbon.     

PHOTOSYNTHETIC PHYSIOLOGY AND CHEMICAL COMPOSITION OF SPORES OF THE KELPS MACROCYSTIS PYRIFERA,NEREOCYSTIS LUETKEANA,LAMINARIA FARLOWII,AND PTERYGOPHORA CALIFORNICA (PHAEOPHYCEAE)1

Recently released spores of the kelps Macrocystis pyrifera (L.) C. Ag., Nereocystis luetkeana (Mert.) Post. and Rupr., Laminaria farlowii Setch., and Pterygophora californica Rupr. had different levels of net photosynthesis. Spore-specific photosynthesis–irradiance relationships were similar in many respects for M. pyrifera, N. luetkeana, and L. farlowii spores. All three species had low rates of net light-saturated photosynthesis. In contrast, spores of P. californica had higher photosynthetic potential and overall net photosynthesis than the other three species. On a cell carbon basis, however, photosynthetic rates in N. luetkeana spores were similar to those of P. californica spores and higher than those of M. pyrifera spores. Chlorophyll a content of spores varied 10-fold among species. The rank order of significant differences in chlorophyll a content was P. californica > L. farlowii > N. luetkeana > M. pyrifera. As a result, chlorophyll-specific measurements suggest M. pyrifera and N. luetkeana spores had much higher quantum efficiency and photosynthetic potential than either P. californica or L. farlowii spores. Maternal carbon and nitrogen investment significantly differed in spores of M. pyrifera, N. luetkeana, and P. californica with P. californica > M. pyrifera > N. luetkeana. Carbon content in spores of each of these three species increased by about 30% during 12 h of saturating irradiance. We suggest that the photosynthetic capabilities of and maternal investment in spores may be related to the spore as a unit of dispersal, to the reproductive ecology of the parental sporophytic stages, and to the growth and physiology of the germling gametophyte stages.