PHOTOSYNTHESIS AND CARBON ALLOCATION IN TIPULARIA DISCOLOR (ORCHIDACEAE), A WINTERGREEN UNDERSTORY HERB

Seasonal patterns of photosynthesis and carbon allocation were determined for Tipularia discolor, a summer-deciduous wintergreen orchid of the southeastern United States, to assess the effects of environmental conditions and leaf age on carbon acquisition and allocation patterns. There was no shift in the optimum temperature for photosynthesis (T_opt) on a seasonal basis and T_opt (≈26 C) was at least 10 C higher than daily maximum air temperature during most of the growing season. Lack of photosynthetic adjustment in Tipularia to seasonal fluctuations in temperature and light suggested that the photosynthetic characteristics of this wintergreen were more similar to those of spring ephemerals than to those of evergreens and summer-active herbs. The decline in photosynthetic capacity during the winter growing season for Tipularia, largely due to leaf age effects, gradually reduced net photosynthetic rates in the field despite more favorable light and temperature conditions. Photosynthesis in the field was primarily limited by environmental conditions in early- and mid-season and by photosynthetic capacity in late-season. A ¹⁴CO₂ labelling experiment demonstrated that patterns of carbon allocation to vegetative structures were affected by the season of photosynthetic carbon fixation, whereas reproductive structures received 21% of the recovered labelled carbon regardless of the period of labelling. Carbon acquired and stored during all periods of the growing season was used to produce new vegetative and reproductive structures.     

A hemiparasite in the forest understorey: photosynthetic performance and carbon balance of Melampyrum pratense

• Melampyrum pratense is an annual root‐hemiparasitic plant growing mostly in forest understorey, an environment with unstable light conditions. While photosynthetic responses of autotrophic plants to variable light conditions are in general well understood, light responses of root hemiparasites have not been investigated.
• We carried out gas exchange measurements (light response and photosynthetic induction curves) to assess the photosynthetic performance of M. pratense in spring and summer. These data and recorded light dynamics data were subsequently used to model carbon balance of the hemiparasite throughout the entire growth season.
• Summer leaves had significantly lower rates of saturated photosynthesis and dark respiration than spring leaves, a pattern expected to reflect the difference between sun‐ and shade‐adapted leaves. However, even the summer leaves of the hemiparasite exhibited a higher rate of light‐saturated photosynthesis than reported in non‐parasitic understorey herbs. This is likely related to its annual life history, rare among other understorey herbs. The carbon balance model considering photosynthetic induction still indicated insufficient autotrophic carbon gain for seed production in the summer months due to limited light availability and substantial carbon loss through dark respiration.
• The results point to potentially high importance of heterotrophic carbon acquisition in M. pratense, which could be of at least comparable importance as in other mixotrophic plants growing in forests – mistletoes and partial mycoheterotrophs. It is remarkable that despite apparent evolutionary pressure towards improved carbon acquisition from the host, M. pratense retains efficient photosynthesis and high transpiration rate, the ecophysiological traits typical of related root hemiparasites in the Orobanchaceae.

     

The seasonal pattern of CO2 exchange of Festuca rubra L. in a montane meadow community in Northern Germany

Summary Completely climatized cuvettes were used to follow the CO₂ gas exchange of red fescue (Festuca rubra L.), growing on a fertilized and an unfertilized plot, during a growing season from May through October. Objective of the study was to determine the effect of environmental factors on the seasonal CO₂ gas exchange.Gas exchange rates were calculated on the basis of leaf dry weight, surface area and chlorophyll content. Photosynthetic rates differed between the fertilized and unfertilized plants when based on leaf dry weight or leaf surface area but were similar when based on chlorophyll.Multiple regression analysis was used to related photosynthetic rates to radiation, temperature, water vapor concentration difference, chlorophyll content and time. A cubic regression equation based on daily radiation alone explained 85% of the variation for the fertilized plants and 87% of the variation for the unfertilized plants.During the growing season the unfertilized plants had a continual decline in their photosynthetic rates. The fertilized plants had high photosynthetic rates in the spring and in the fall.Light response curves indicated greater photosynthetic rates at light saturation as well as in the light limited portion of the light response curve for the fertilized plants. Photosynthetic rates of the fertilized plants were generally depressed during periods of warm temperature and high light intensity in June and July.Photosynthetic rates declined at temperatures above 24°C. The decline was greater for the more mesomorphic fertilized plants. A similar response was noted to increasing water vapor difference, although it was difficult to separate from the temperature effect. Maximum photosynthetic rates were found between 14°C and 22°C, although there was considerable variation in the maximum rates.The effects of cutting (mowing) on the gas exchange were difficult to determine due to the interaction of the environmental factors.Chlorophyll content showed significant correlation with photosynthetic rates.     

Improved growth and water use efficiency of cherry saplings under reduced light intensity

Cherry (Prunus avium L.) saplings were grown under natural sunlight (controls) or moderate shading (up to 30%, depending on the incident light intensity and the hour of the day). Reduced light intensity increased the dry mass of each of the plant components studied. Consequently, the total dry mass of shaded plants was significantly greater than that of controls at the end of the growing season. However, the diurnal trend in the level of photosynthesis (per unit of leaf area) of shaded plants was similar to the controls in August, but lower in September. As the growing season proceeded, reduced photosynthetic rates, thinner mesophyll and larger specific leaf area in the shaded plants indicated that leaf development had adapted to shaded conditions throughout the growing season. It is suggested that increased growth of shaded plants was caused by a higher initial relative growth rate and a greater whole-plant photosynthesis. Shading consistently reduced transpiration over the season, therefore improving water use efficiency of shaded leaves. Our results suggest that a moderate reduction in light intensity can be a useful method for improving growth and saving water in hot and dry environments.     

Seasonal patterns of acid fluctuations and resource storage in the arborescent cactus Opuntia excelsa in relation to light availability and size

Summary We investigated relationships between light availability, diel acid fluctuation, and resource storage in the arborescent cactus Opuntia excelsa growing in western Mexico. We compared canopy and understory individuals from a deciduous forest as well as open-grown plants of the same approximate size as those in the understory. During the wet season light availability and daily fluctuations in titratable acidity (an index of carbon uptake) were lower in the understory than in unshaded habitats. In the dry season all plants had reduced levels of acid fluctuation, with the smallest individuals, regardless of habitat, showing the greatest reduction. These data suggest that light availability in the forest understory constrains carbon assimilation during the wet season, but that a factor associated with plant size, possibly water status, limits carbon gain during the dry season. Plants in all habitats remained physiologically active for at least five months into the dry season. We suggest that this was possible due to the maintenance of constant concentrations of water and nitrogen in the photosynthetically active chlorenchyma. Parenchyma in terminal cladodes showed a different seasonal pattern of resource storage; water content and nitrogen concentration were reduced from the wet to the dry season in the parenchyma. Using the parenchyma to supply photosynthetic tissues during times of reduced resource availability allows O. excelsa to assimilate carbon during times of the year when most other trees in the forest are leafless.     

Photosynthesis inCeratophyllum demersum. Carbon fixation rates in relation to the plants' physiological stage,and the contents of chlorophylland non-structural carbohydrates

Summary Since the role of the submerged aquatic macrophytes in the carbon cycle of lake Vechten (the Netherlands) is in study, attention is paid to several aspects of their productivity.Ceratophyllum demersum is a predominant macrophytic species in the littoral zone of this lake, occurring mainly from three to five metres depth. In situ measurements of its photosynthetic rate demonstrated a maximum in spring (May), followed by a distinct decrease early in summer and a second maximum in late summer. The spring maximum of the photosynthetic rate coincided with the maximum increase of plant biomass. Maximum biomass occurred in early autumn (August–September).Althoughinsitu measurements pointed to the upper plant portions (70 cm) as the most important with respect to photosynthetic activity, it was demonstrated under standard experimental conditions that lower portions might be equally active depending on the age of the plant. In the lake, light reduction with depth is one of the major causes in the commonly observed decrease in photosynthetic activity in lower plant parts.Comparison of the photosynthetic rates in plant tipsin situ and under standard experimental conditions indicated that in spring, both water temperature and the chlorophyll content of the plant limit photosynthesis. This was concluded since under experimental conditions (a) photosynthetic rate and chlorophyll content were strongly correlated during the first half of the growing season and (b) higher rates were measured thanin situ. In the subsequent stage, however, light and the distribution of non-structural carbohydrates (TNC) in the plant also play an important role.At the second half the growing season, the lower parts of plants growing in the lake lose gradually their buoyancy, caused by a deteriorating light climate resulting in a decreasing photosynthetic activity.     

Respiration and photosynthesis in cones of Norway spruce (Picea abies (L.) Karst.)

Summary Dark respiration and photosynthetic carbon dioxide refixation in purple and green Picea abies cones were investigated from budbreak to cone maturity. The rate of dark respiration per unit dry weight and CO₂ refixation capacity decreased during cone maturation. At the beginning of the growing season, photosynthetic CO₂ refixation could reduce the amount of CO₂ released by respiration in green and purple cones by 50% and 40%, respectively. The seasonal performance of the components of the cone carbon balance was calculated using information on the seasonal course of respiration, refixation capacity and the light response curves of cone photosynthesis, as well as the actual light and temperature regime in the field. The daily gain of CO₂ refixation reached 28%–34% of respiration in green and 22%–26% in purple cones during the first month of their growth, but decreased later in the season. Over the entire growth period refixation reduced carbon costs of cone production in both cone colour polymorphs by 16%–17%.     

Seasonal variation in crassulacean acid metabolism by the aquatic isoetid Littorella uniflora

The seasonal temperature acclimation in crassulacean acid metabolism (CAM) and photosynthetic performance were investigated in the aquatic isoetid, Littorella uniflora. Plants were collected monthly from January to September, and CAM capacity and photosynthesis rates were measured at 5, 10, 15, and 20?°C. Seasonal acclimation was observed for CAM (Q (10) range: 0.6-1.8), and CAM was optimised close to ambient temperature throughout the season. Thus, in winter acclimated L. uniflora, the short-term response to raised temperature resulted in a decline in CAM capacity. Even though the ambient CAM increased from winter to spring/summer, CAM was present in cold acclimated plants, thus indicating an ecophysiological role for CAM even in winter. Similar to CAM, seasonal acclimation was observed in the light and carbon-saturated photosynthesis (Q (10) values ranged from 1.4 to 2.3), and the photosynthetic capacity was generally higher during the winter at all temperatures, indicating compensatory investments in the photosynthetic apparatus. Thus, L. uniflora displayed seasonal temperature acclimation with respect to both CAM and photosynthesis. The estimated in situ contribution of CAM to the carbon budget in L. uniflora was independent of season and varied from 23 to 46?%. A positive correlation between photosynthetic capacity and CAM capacity (both measured in the lab at temperature close to ambient temperature) was found, and the ratio of CAM activity to photosynthetic capacity was higher in summer compared with winter plants. Overall, the results from the present study support the suggested role of CAM as a carbon conserving mechanism of importance for survival in a carbon-limited habitat.     

Gas exchange and resource-use efficiency of Leymus cinereus (Poaceae): diurnal and seasonal responses to naturally declining soil moisture

We examined factors that limit diurnal and seasonal photosynthesis in Leymus cinereus, a robust tussock grass from shrub-steppes of western North America. Data from plants in a natural stand and in experimental field plots indicate that this bunchgrass has 1) a high photosynthetic capacity, 2) high leaf nitrogen content and high nitrogen-use efficiency, 3) a steep leaf-to-air diffusion gradient for carbon dioxide, which enhances intrinsic water-use efficiency, and 4) photosynthetic tissues that tolerate severe water stress and recover quickly from moderate water stress. Midday depressions of CO₂ assimilation (A) and stomatal conductance were slight in plants with plentiful water, but marked in plants subject to moderate water stress. Midday stomatal closure in moderately stressed plants reduced intercellular carbon dioxide concentration (c_i) by ≈40 μl liter^-1. The maximum rate of A achieved during the day for severely stressed plants (predawn water potential = -4 MPa) was one-third and daily carbon gain per unit leaf area was about one-fourth that of well-watered plants. For plants in the natural stand, CO₂-saturated photosynthesis declined almost linearly with decreasing soil water availability over the growing season, whereas there was little effect on A at CO₂ ambient levels or on carboxylation efficiency until predawn water potentials reached -1.8 MPa. Nitrogen-use efficiency declined with diminishing soil moisture, but there was no seasonal change in stomatal limitation or instantaneous water-use efficiency as estimated from A vs. c_i curves at optimal leaf temperature and moderate atmospheric evaporative demand. Thus, reduced stomatal conductance in response to increased evaporative demand may increase stomatal limitation diumally, but over the growing season, stomatal limitation estimated from A vs. c_i curves is relatively constant because maximum stomatal conductance is closely tuned to the CO₂ assimilatory capacity of the mesophyll.     

A model of the seasonal pattern of carbon acquisition in two woodland herbs,Mercurialis perennis L. and Geum urbanum L.

Summary Seasonal changes in the light and temperature dependence of photosynthesis were investigated in field grown plants of Mercurialis perennis and Geum urbanum. In both species changes in photosynthetic capacity were closely related to the development of the overstorey canopy. In G. urbanum there was a marked shift in the temperature dependence of photosynthesis through the season whereas no such pattern was found in M. perennis. Model predictions of field rates of photosynthesis were made using the measurements of light and temperature dependence in the laboratory and validated against field observations. Long term continuous records of light and temperature in the field were used in conjunction with the model to make predictions of carbon acquisition in shoots of the two species through the season. These calculations indicated that G. urbanum was able to take advantage of high light levels just prior to canopy closure through a combination of high photosynthetic capacity, the ability to maintain photosynthesis at relatively low temperatures and the presence of overwintering leaves. In M. perennis leaf development was early enough to utilise the high spring light period. After canopy closure M. perennis maintained a higher average rate of CO₂ flux due to a combination of high apparent quantum efficiency and low rates of respiration.     

Morphological and Physiological Acclimation Responses to Contrasting Light and Water Regimes in <Emphasis Type="Italic">Primula sieboldii</Emphasis>

The effects of the availability of light (high, medium and low) and soil water (wet and dry) on morphological and physiological traits responsible for whole plant carbon gain and ramet biomass accumulation were examined in a splitter-type clonal herbaceous species Primula sieboldii, a spring plant inhabiting broad range of light environments including open grassland and oak forest understory. Growth experiments were conducted for three genets originated from natural microhabitats differing in light and soil water availability. Ramets of a genet from high light and wet microhabitat, which were grown in low light (relative photon flux density: R-PPFD of 5%) showed 41% less light-saturated photosynthetic rate, 50% less dark respiration rate and earlier defoliation than the ramets in high light (R-PPFD of 61%). The estimation of daily photosynthesis revealed that the light acclimation response in leaf gas exchange contributes to efficient carbon gain of whole plants, irrespective of experimental light conditions. Water stress increased root weight ratio, decreased ramet leaf area, petiole length and photosynthetic capacity. These morphological effects of water stress were larger in high and medium light regimes than in low light regime. The consequence of the above responses was recognized in the relative growth rate of the ramets. The relative growth rate of the ramets in high light with wet regime was four-fold of that in low light plus wet regime, and was 1.5-fold of that in high light plus dry regime. However, even in low light and/or dry regimes, ramets kept positive relative growth rates and produced gemma successfully. We could not detect significant variation in growth responses among genets. The high photosynthetic plasticity revealed in the present study should enable Primula sieboldii to inhabit in a broad range of light and soil water availability.     

An empirical model of net photosynthesis for the desert plant Hammada scoparia (Pomel) Iljin

Summary An empirical model for describing daily courses of net photosynthesis in Hammada scoparia is being developed. The model is based on the functional relationships, by which various environmental factors affect the photosynthetic activity and which can be measured by experiment in the field. In a sequence of steady-states daily courses of net photosynthesis are predicted during a growing season considering the variability of the physiological states and the capacity for regulative adaptations. The rate of net photosynthesis at a certain date is calculated from the maximal rate of CO₂ uptake being expected at that season and from the effects of light, temperature, and air humidity which are scaled from 0 to 1. All factors are connected multiplicatively. The light function accounts for the seasonal changes in the light curve, the temperature function is based on the seasonal shift of the temperature optimum, and the humidity function considers the increasing sensitivity of the stomatal humidity response at increasing water stress. The model is built to be a submodel of a general ecosystem model, where various other submodels (i.e. water stress model, phenology model) are supplied. The present model is tested by predicting daily courses at extreme climatic conditions during the year and by comparing the predicted values of gas exchange with values being measured in an independent experimental procedure. The result shows that the model is able to simulate the natural behaviour of Hammada scoparia during the growing and dry season of a desert habitat. The problems of incorporating the influence of water stress, the interaction of the various factors, and the phenological aspect of the photosynthetic activity is being discussed.     

Early-spring gas exchange and uptake of deuterium-labelled water in the poikilohydric fernPolypodium virginianum

This study was carried out to determine if the desiccation-tolerant fernPolypodium virgimanum L. ecologically resembles lower plants by absorbing atmospheric water through its fronds and actively growing in early spring when the soil along cliff edges is still frozen. Three times between February and April, 1991,P. virginianum clonal mats were treated with deuterium-labelled water. Following each application, fronds were collected over several days and analyzed for the presence of deuterium. Two treatment groups plus a control were used: fronds were sprayed directly while covering the soil, or the roots were watered while protecting the fronds. The control mats were left untreated. Soil, air, and frond temperatures, plus photosynthesis and frond conductance were monitored throughout the study period. At subfreezing temperatures in February, no labelled water was taken up from the soil and no photosynthesis took place. Small amounts of label were absorbed from the soil in March during freeze-thaw cycles when rates of photosynthesis and stomatal conductance were both low. Large amounts of label were taken up from the soil in April when the soil was fully thawed and gas exchange was at normal seasonal levels. Label was not absorbed directly through the fronds when the plants were actively growing. Despite the desiccation tolerance ofP. virginianum, the timing and patterns of its water uptake and gas exchange in early spring resemble those found in higher vascular plants, not poikilohydric lower plants.     

Measuring and modelling environmental influences on photosynthetic gas exchange in Sphagnum and Pleurozium

A mechanistic model has been used to examine the environmental regulation of photosynthetic gas exchange in moss. The effects of water content on conductance to CO₂ and on photosynthetic capacity during desiccation were calculated from the carbon isotope discrimination data of Williams & Flanagan (1996 , Oecologia 108, pp. 38–46) and combined with the biochemical model of Farquhar et al. (1980 , Planta 149, pp. 78–90). The model includes a simple light attenuation function that imparts curvature to the light response curve for net assimilation, enabling the use of physiologically realistic values for the biochemical parameters. Measurements of gas exchange for Sphagnum and Pleurozium were made in an old black spruce ecosystem over a growing season in order to assign values to parameters in the model. The calculated maximum rates of carboxylation by Rubisco ( V _max) were 5, 14 and 6 μ mol m^–2 s^–1 for Sphagnum during the spring, summer and autumn seasons of 1996, respectively. The increase in V _max during the summer was consistent with an increased allocation of resources to the photosynthetic apparatus. In contrast, no seasonal variation in V _max was observed in Pleurozium with average values of 7, 5 and 7 μ mol m^–2 s^–1 during the spring, summer and autumn, respectively.     

CONTRASTING STOMATAL RESPONSES TO VARIABLE SUNLIGHT IN TWO SUBALPINE HERBS

Gas exchange was measured in two subalpine herbs during alternating periods of sun and shade that simulated natural cloud patterns. Stomatal conductance (g) in the subalpine herb Helianthella quinquenervis was reduced by as much as 75% during 5-min shade periods that ranged in photosynthetic photon flux density (PPFD) from 100 to 1,100 μmol m^–2 sec^–1. In contrast, responses in g in another herb, Frasera speciosa, to fluctuations in sunlight were small, apparently due to slower stomatal responses. Based on an earlier hypothesis that water stress may strongly influence nonsteady state gas exchange responses to fluctuations in PPFD, net photosynthesis and g were measured in these herbs as seasonal water stress increased and compared with responses in irrigated plants. Stomatal conductance was relatively unresponsive to changes in PPFD in F. speciosa regardless of water stress. In contrast, substantial decreases in g occurred for H. quinquenervis during shade only when this species experienced water stress later in the season. Little response in g was measured early in the season or in irrigated plants. The seasonal shift in nonsteady state responses in H. quinquenervis from little response in g to shade when plants were nonstressed to more rapid reductions in g as water stress increased would maximize carbon gain early in the season when soil water was abundant while conserving water during periods of soil drought.     

Influence and predictive capacity of climate anomalies on daily to decadal extremes in canopy photosynthesis

Significant advances have been made over the past decades in capabilities to simulate diurnal and seasonal variation of leaf-level and canopy-scale photosynthesis in temperate and boreal forests. However, long-term prediction of future forest productivity in a changing climate may be more dependent on how climate and biological anomalies influence extremes in interannual to decadal variability of canopy ecosystem carbon exchanges. These exchanges can differ markedly from leaf level responses, especially owing to the prevalence of long lags in nutrient and water cycling. Until recently, multiple long-term (10+ year) high temporal frequency (daily) observations of canopy exchange were not available to reliably assess this claim. An analysis of one of the longest running North American eddy covariance flux towers reveals that single climate variables do not adequately explain carbon exchange anomalies beyond the seasonal timescale. Daily to weekly lagged anomalies of photosynthesis positively autocorrelate with daily photosynthesis. This effect suggests a negative feedback in photosynthetic response to climate extremes, such as anomalies in evapotranspiration and maximum temperature. Moisture stress in the prior season did inhibit photosynthesis, but mechanisms are difficult to assess. A complex interplay of integrated and lagged productivity and moisture-limiting factors indicate a critical role of seasonal thresholds that limit growing season length and peak productivity. These results lead toward a new conceptual framework for improving earth system models with long-term flux tower observations.     

Effects of an Experimental Increase of Temperature and Drought on the Photosynthetic Performance of Two Ericaceous Shrub Species Along a North–South European Gradient

Plant ecophysiological changes in response to climatic change may be different in northern and southern European countries because different abiotic factors constrain plant physiological activity. We studied the effects of experimental warming and drought on the photosynthetic performance of two ericaceous shrubs (Erica multiflora and Calluna vulgaris) along a European gradient of temperature and precipitation (UK, Denmark, The Netherlands, and Spain). At each site, a passive warming treatment was applied during the night throughout the whole year, whereas the drought treatment excluded rain events over 6–10 weeks during the growing season. We measured leaf gas exchange, chlorophyll a fluorescence, and leaf carbon isotope ratio (¹³C) during the growing seasons of 1999 and 2000. Leaf net photosynthetic rates clearly followed a gradient from northern to southern countries in agreement with the geographical gradient in water availability. Accordingly, there was a strong correlation between net photosynthetic rates and the accumulated rainfall over the growing season. Droughted plants showed lower leaf gas exchange rates than control plants in the four sites. Interestingly, although leaf photosynthetic rates decreased along the precipitation gradient and in response to drought treatment, droughted plants were able to maintain higher leaf photosynthetic rates than control plants in relation to the accumulated rainfall over the months previous to the measurements. Droughted plants also showed higher values of potential photochemical efficiency (F _v/F _m) in relation to controls, mainly at midday. The warming treatment did not affect significantly any of the studied instantaneous ecophysiological variables..     

Seasonal changes in photosynthetic characteristics of Anemone raddeana,a spring-active geophyte,in the temperate region of Japan

Summary Seasonal changes in the photosynthetic characteristics of intact involucral leaves of Anemone raddeana were investigated under laboratory conditions. Net photosynthesis and constant water vapor pressure deficit showed almost the same seasonal trend. They increased rapidly from mid-April immediately after unfolding of the leaves and reached the maximum in late-April, before the maximum expansion of the leaves. They retained the maximum values until early-May and then decreased toward late-May with a progress of leaf senescence. The calculated values of intercellular CO₂ concentration and relative stomatal limitation of photosynthesis showed no significant change throughout the season. The carboxylation efficiency as assessed by the initial slope of Ci-photosynthesis curve and the net photosynthesis under a high Ci regime varied seasonally in parallel with the change of the light-saturated photosynthesis. The results indicate that the seasonal changes in light-saturated net photosynthesis are not due to a change of stomatal conductance, but to a change in the photosynthetic capacity of mesophyll. Nevertheless, leaf conductance changed concomitantly with photosynthetic capacity, indicating that the seasonal change in stomatal conductance is modulated by the mesophyll photosynthetic capacity such that the intercellular CO₂ concentrations is maintained constant. The shape of light-photosynthesis curve was similar to that of sun-leaf type. The quantum yield also changed simultaneously with the photosynthetic capacity throughout the season.Contribution No. 2965 from the Institute of Low Temperature Science     

Fruit Development in Trillium : Dependence on Stem Carbohydrate Reserves

Leaves are the main source of carbon for fruit maturation in most species. However, in plants seeing contrasting light conditions such as some spring plants, carbon fixed during the spring could be used to support fruit development in the summer, when photosynthetic rates are low. We monitored carbohydrate content in the rhizome (a perennating organ) and the aboveground stem of trillium (Trillium erectum) over the entire growing season (May–November). At the beginning of the fruiting stage, stems carrying a developing fruit were harvested, their leaves were removed, and the leafless stems were maintained in aqueous solution under controlled conditions up to full fruit maturation. These experiments showed that stem carbohydrate content was sufficient to support fruit development in the absence of leaves and rhizome. This is the first reported case, to our knowledge, of complete fruit development sustained only by a temporary carbohydrate reservoir. This carbohydrate accumulation in the stem during the spring enables the plant to make better use of the high irradiances occurring at that time. Many other species might establish short-term carbohydrate reservoirs in response to seasonal changes in growing conditions.     

Photosynthesis and water relations of well-watered orange plants as affected by winter and summer conditions

The aim of this study was to evaluate how the summer and winter conditions affect the photosynthesis and water relations of well-watered orange trees, considering the diurnal changes in leaf gas exchange, chlorophyll (Chl) fluorescence, and leaf water potential (Ψ) of potted-plants growing in a subtropical climate. The diurnal pattern of photosynthesis in young citrus trees was not significantly affected by the environmental changes when compared the summer and winter seasons. However, citrus plants showed higher photosynthetic performance in summer, when plants fixed 2.9 times more CO₂ during the diurnal period than in the winter season. Curiously, the winter conditions were more favorable to photosynthesis of citrus plants, when considering the air temperature (< 29 °C), leaf-to-air vapor pressure difference (< 2.4 kPa) and photon flux density (maximum values near light saturation) during the diurnal period. Therefore, low night temperature was the main environmental element changing the photosynthetic performance and water relations of well-watered plants during winter. Lower whole-plant hydraulic conductance, lower shoot hydration and lower stomatal conductance were noticed during winter when compared to the summer season. In winter, higher ratio between the apparent electron transport rate and leaf CO₂ assimilation was verified in afternoon, indicating reduction in electron use efficiency by photosynthesis. The high radiation loading in the summer season did not impair the citrus photochemistry, being photoprotective mechanisms active. Such mechanisms were related to increases in the heat dissipation of excessive light energy at the PSII level and to other metabolic processes consuming electrons, which impede the citrus photoinhibition under high light conditions.