Can early wilting of old leaves account for much of the ABA accumulation in flooded pea plants?

When pea plants (Pisum sativum L.) were subjected to flooding,abscisic acid (ABA) content in shoots and roots increased upto 8-fold in the following days and stomatal conductance significantlydecreased. Although young leaves of flooded plants had a slightlyhigher water potential than those of the unflooded plants, oldleaves had lower water potential and lost turgor at the timewhen a substantial ABA increase was detected. In plants wherethe old leaves were clipped off, flooding did not cause anyABA increase during 7 d of the experimental period, except underconditions of higher transpiration demand, when the increasein ABA content was both delayed and small in scale (only I-fold).When intact plants were flooded and ABA was assayed separatelyin both old and young leaves, the ABA increase in old leavespreceded that in young leaves. Evidence here suggests that theflooding-induced ABA increase mainly results from the wiltingof old leaves. This suggests that young leaves may be protectedfrom wilting by ABA originating in old leaves under unfavourableenvironmental conditions. Key words: Waterlogging, soil flooding, ABA, leaf water relations, pea, Pisum sativum     

Water translocation in Kalanchoë daigremontiana during periods of drought

Abstract. Kalanchoë daigremontiana strongly reduced daily water loss within 6 d of drought using CAM to restrict transpiration and net CO₂ uptake to the dark period.
Water translocation from old to young leaves of the plant was an additional mechanism which reduced the negative effects of drought on the water relations of young leaves. Excision of old leaves after 7–9 d of drought resulted in a decrease in the water content of young leaves. This was observed despite a decrease in transpirational water loss from young leaves. Water content in young leaves increased slightly in plants with all their leaves in place.
The dry weight of young leaves clearly increased during the experimental period when old leaves were present, but it remained relatively constant in plants without old leaves. Obviously, in addition to water, solutes were transported from old to young leaves of the plant via the phloem. Xylem tension was higher in young compared to old leaves; thus, water translocation could have occurred via xylem elements.
Since transport of organic matter in the phloem is also linked to water flow, phloem transport additionally may contribute effectively to the balance of the water budget in young leaves.     

Water recycling in leaves of Lithops (Aizoaceae)

Lithops plants consist of a pair of succulent leaves inserted on a short stem; in each growing season, young leaves develop in a cavity formed between the older pair. Young leaves can take up water from the older pair allowing the plant to maintain growth and leaf expansion even without external supply of water. Recycling water between vegetative organs is one of the possible adaptation strategies of plants under drought stress, but it had never been demonstrated experimentally in Lithops. The methodology used to verify the existence of water redistribution from old leaves to young leaves was fluorescence microscopy, using two dyes to follow the water pathway inside the plant: Sulforhodamine G (SRG) and 5(6)-carboxyfluoroscein diacetate (CFDA). In Lithops fluorescent tracers loaded into old leaves were found in young leaves, in 74% of the cases for SRG, in 59% of the cases for CFDA. Our data demonstrate that young leaves take up water from the old ones following both a symplastic and an apoplastic pathway. Water recycling is therefore one of the adaptive responses of these plants allowing them to perform at least a complete growth cycle even during prolonged drought stress periods, using the water stored in the older leaves.     

Mineral nutrition and the water relations of plants

Summary When young tomato plants were transferred from nutrient solution to mineral-free water, reductions in transpiration, water content of the shoots and stomatal aperture were not accompanied by a reduction in the relative water content or an increase in the suction pressure of the leaves. The relative water content of the leaves was increased and the suction pressure was little affected.Following transfer of the plants to mineral-free water, the mineral content of the shoots and the osmotic pressure of expressed leaf sap were reduced. It was concluded that mineral salts were necessary for maintaining the osmotic pressure of the leaf cell sap and that this was achieved, at least in part, by maintaining the mineral concentration of the sap. The amount of water that could be taken up by leaves and their turgor pressure were related to the osmotic pressure of the sap and calculations of turgor pressure showed that it was less in the leaves of plants with their roots in mineral-free water than in the leaves of plants in nutrient solution.Evidence was obtained that in leaflets detached from plants with their roots in mineral-free water, stomatal closure could occur at a higher water content than in leaflets detached from plants in nutrient solution, indicating a further role of minerals in leaf water relations. It is suggested that this role may be related to the properties of the cell walls.     

Changes in photosynthetic and transpiration rates of cotton leaves infested with the cotton aphid, Aphis gossypii: Unrestricted infestation

The influence of aphid, Aphis gossypii, feeding on photosynthesis and transpiration in cotton plants was investigated under greenhouse conditions. Four population densities of 0, 5, 10, and 25 aphids were used to infest individual cotton leaves. Gas exchange rates were determined for single attached cotton leaves after 9, 18, and 27 days of aphid infestation. Aphid feeding changed photosynthetic rates and transpiration rates. These changes were proportional to the number of aphids and the length of infestation period. Photosynthetic rates were significantly reduced in infested leaves with 25 aphids over 18 days, whereas significant reduction in photosynthetic rates was recorded within 27 days in infested leaves with 5, 10, and 25 aphids in comparison to their respective control. Initial population of 10 aphids increased significantly the transpiration rate of infested leaves over 9 and 27 days. Leaves of plants with 25 aphids had significantly greater transpiration rate than the control at all times.     

Sexual dimorphism and seasonal changes of leaf gas exchange in the dioecious tree Ilex paraguariensis grown in two contrasted cultivation types

Yerba maté (Ilex paraguariensis, Aquifoliaceae) is a subtropical, evergreen, dioecious, South American tree. In one preliminary study, it was observed that the functional strategy of yerba mate females, aiming to finish reproductive process, was increased transpiration relative to photosynthetic rates compared with males, on self‐shaded leaves. We hypothesised that the long‐term gas exchange response of males and females can evolve independently of phenological stage and cultivation type. In this spirit, the primary aim of the study was to analyse the physiological sexual dimorphism of this species, evaluating fluctuations of gas exchanges related to microclimate and phenological stages. A field study was conducted on adult plants of yerba maté cultivated in monoculture (MO) and in forest understorey (FUS), and measurements carried out in situ on microclimate and leaf gas exchange parameters. The photosynthetic photon flux density that was attained at leaf level in FUS was reduced roughly 10‐fold compared with that in MO. Various leaf age populations were observed during a 2‐year period at 2‐month intervals and grouped into four categories: young, young‐fully‐expanded, fully‐expanded and old. Young and young‐fully‐expanded leaves were the most active in photosynthesis. Leaves of female plants showed greater photosynthetic rate than those of male plants, which was expressed on all leaf age categories in MO, but only during vegetative stages previous to flowering and fruit ripening. The photosynthesis of young‐fully‐expanded leaves of females grown in FUS was superior to males but only during winter growth pause. The stomatal conductance differed in relation to cultivation type and leaf age but did not show the sexual differentiation. Physiological sexual dimorphism in yerba mate is shown to be plastic, responding to environmental conditions. The cost associated to the reproduction of yerba maté could be most easily met showing physiological differentiation of both sexes. A higher reproductive investment of females might be compensated for by exhibiting greater leaf photosynthesis than males that occurs in vegetative stages that precede flowering and fruit ripening.     

Spectral properties of heavily glaucous and non-glaucous leaves of a succulent rosette-plant

Summary Comparisons of reflection, transmittance, and absorptance spectra of heavily glaucous leaves, glaucous leaves from which the glaucescence was experimentally removed, and naturally occurring non-glaucous leaves of a single species (Dudleya brittonii Johansen) reveal that glaucescence, a powdery wax coating on the leaf surface, is responsible for very high reflectance of ultraviolet radiation (UV) and high reflectances of visible and near-infrared radiation. UV reflectance (up to 83% in UV-B) measured for glaucous leaves of D. brittonii, a succulent rosette-plant occurring in fissures in steep volcanic outcrops, is higher than that reported for any other plant species. Non-glaucous leaves of the species reflect about 10% in UV. I hypothesize that the high UV reflectance of glaucous leaves is ecologically significant in reducing damage to dehydrated leaves from visible and UV-B radiation, thus promoting longevity of the leaves important both in conservation of the mineral nutrient capital of these succulent plants and in their acquisition and storage of water. This and other demonstrated and hypothesized functions of glaucescence are discussed in terms of the evolutionary significance of glaucescence in succulent plants.     

Apoplastic Transport of 14C-Photosynthates Measured under Drought and Nitrogen Supply

Using water infiltration of the plant and individual shoots with the subsequent intercellular liquid extraction by the pressure chamber, dynamics of the movement ¹⁴C-photosynthates from cell to apoplast, and ¹⁴C distribution among photosynthetic products in mesophyll cells and apoplast were studied. The relative quantity of ¹⁴C-photosynthetes in leaf apoplast depended on growing conditions; drought increased, and nitrate supply decreased it. When the middle leaves absorbed ¹⁴CO₂, photosynthates moving down in stem phloem appeared in intercellular space, where they were transported up by transpiration stream. ¹⁴C-photosynthates entering to the apex and young leaves were utilized a accumulated, and photosynthates transported to the mature leaves were reloaded into the phloem and reexported. Thus, photosynthates circulated through the plant and were redistributed to the plant organs according to their transpiration. In leaf apoplast photosynthetic sucrose was partly hydrolyzed to glucose and fructose. This increased under high nitrogen supply. The result indicate that apoplast sucrose hydrolysis is the basic cause of the reduction of photosynthate flux from leaves when the nitrate concentration in soil increases.     

Effects of varying air and soil moisture on the water relations and growth of sugar beet

Sugar beet were grown for short periods with different amounts of moisture in the soil and air. Growing plants in wet soil (23 % moisture on dry weight) compared with dry soil (15% moisture) increased growth of the shoots and roots and plant dry weights by 15% in young plants and 10% in mature plants. Growing plants in wet air containing 10.9 g m^-3 of water (equivalent to a saturation deficit of 2.5 mb) compared with dry air containing 6.4 g m^-3 of water (saturation deficit = 8.5 mb) increased the dry weights of both young and mature plants by 8%, mostly by increasing the sizes of their storage roots. Wet air and wet soil increased the net assimilation rates of both young and mature plants. Wet soil, but not wet air, increased leaf areas of young plants by accelerating leaf expansion, and both increased the leaf area of mature plants by slowing senescence of the older leaves. Wet soil increased the water potential of the leaves of both young and mature plants and, by doing so, increased their stomatal conductances and rates of photosynthesis. Wet air also increased stomatal conductances and rates of photosynthesis of leaves of plants of both ages, but without changing their water potentials. Stomatal conductances and photosynthetic rates were greater for young leaves than mature on the same plant and at the same water potential. It is suggested that at certain stages in the crops growth photosynthetic efficiency could be increased by applying additional water as a mist to increase the moisture content of the air around the crop.     

Phosphorus concentrations in the leaves of defoliated white clover affect abscisic acid formation and transpiration in drying soil

Increased leaf phosphorus (P) concentration improved the water-use efficiency (WUE) and drought tolerance of regularly defoliated white clover plants by decreasing the rate of daily transpiration per unit leaf area in dry soil. Night transpiration was around 17% of the total daily transpiration. The improved control of transpiration in the high-P plants was associated with an increased individual leaf area and WUE that apparently resulted from net photosynthetic assimilation rate being reduced less than the reductions in the transpiration (27% vs 58%). On the other hand, greater transpiration from low-P plants was associated with poor stomatal control of transpirational loss of water, less ABA in the leaves when exposed to dry soil, and thicker and smaller leaf size compared with high-P leaves. The leaf P concentration was positively related with leaf ABA, and negatively with transpiration rates, under dry conditions ( P < 0.001). However, leaf ABA was not closely related to the transpiration rate, suggesting that leaf P concentration has a greater influence than ABA on the transpiration rates.     

Enhancement of photosynthesis in post-disturbance resprouts of two co-occurring Mediterranean <Emphasis Type="Italic">Erica</Emphasis> species

The higher growth rates of resprouting shoots compared with those of mature plants in resprouter woody species are supported by higher rates of photosynthesis and transpiration. In this contribution we hypothesize that species with higher resprouting vigour will show a larger enhancement of photosynthesis in resprouting shoots. We test this hypothesis by comparing gas exchange and leaf parameters between resprouting and mature plants in Erica scoparia and E. australis. These two Erica species co-occur in Mediterranean heathlands of the Strait of Gibraltar. Erica scoparia has a higher rate of post-disturbance starch recovery than E. australis, which makes it more resistant to recurrent disturbance. We tested the hypothesis that enhancement of photosynthesis and water use characteristics of resprouting shoots compared with mature plants should be more pronounced in E. scoparia. In both species, resprouts had higher efficiency in the use of light and higher maximum net photosynthesis than mature shoots. However, contrary to expectations, differences in the photosynthetic performance between resprouts and mature plant shoots were larger in E. australis. Higher root to shoot ratios in resprouting E. australis plants, determined by their slower above-ground recovery, together with stronger demand from carbon sinks might explain this result.     

Structural and hydraulic correlates of heterophylly in Ginkgo biloba

This study investigates the functional significance of heterophylly in Ginkgo biloba, where leaves borne on short shoots are ontogenetically distinct from those on long shoots. Short shoots are compact, with minimal internodal elongation; their leaves are supplied with water through mature branches. Long shoots extend the canopy and have significant internodal elongation; their expanding leaves receive water from a shoot that is itself maturing. Morphology, stomatal traits, hydraulic architecture, Huber values, water transport efficiency, in situ gas exchange and laboratory-based steady-state hydraulic conductance were examined for each leaf type. Both structure and physiology differed markedly between the two leaf types. Short-shoot leaves were thinner and had higher vein density, lower stomatal pore index, smaller bundle sheath extensions and lower hydraulic conductance than long-shoot leaves. Long shoots had lower xylem area:leaf area ratios than short shoots during leaf expansion, but this ratio was reversed at shoot maturity. Long-shoot leaves had higher rates of photosynthesis, stomatal conductance and transpiration than short-shoot leaves. We propose that structural differences between the two G. biloba leaf types reflect greater hydraulic limitation of long-shoot leaves during expansion. In turn, differences in physiological performance of short- and long-shoot leaves correspond to their distinct ontogeny and architecture.     

Inducibility of crassulacean acid metabolism (CAM) in Clusia species; physiological/biochemical characterisation and intercellular localization of carboxylation and decarboxylation processes in three species which exhibit different degrees of CAM

The biochemical basis for photosynthetic plasticity in tropical trees of the genus Clusia was investigated in three species that were from contrasting habitats and showed marked differences in their capacity for crassulacean acid metabolism (CAM). Physiological, anatomical and biochemical measurements were used to relate changes in the activities/amounts of key enzymes of C₃ and C₄ carboxylation to physiological performance under severe drought stress. On the basis of gas-exchange measurements and day/night patterns of organic acid turnover, the species were categorised as weak CAM-inducible (C.aripoensis Britt.), C₃-CAM intermediate (C. minor L.) and constitutive CAM (C.␣rosea Jacq. 9.). The categories reflect genotypic differences in physiological response to drought stress in terms of net carbon gain; in C. aripoensis net carbon gain was reduced by over 80% in drought-stressed plants whilst carbon gain was relatively unaffected after 10 d without water in C. rosea. In turn, genotypic differences in the capacity for CAM appeared to be directly related to the capacities/amounts of phosphoenolpyruvate carboxylase (PEPCase) and phosphoenolpyruvate carboxykinase (PEPCK) which increased in response to drought in both young and mature leaves. Whilst measured activities of PEPCase and PEPCK in well-watered plants of the C₃-CAM intermediate C. minor were 5–10 times in excess of that required to support the magnitude of organic acid turnover induced by drought, close correlations were observed between malate accumulation/PEPCase capacity and citrate decarboxylation/PEPCK capacity in all the species. Drought stress did not affect the amount of ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco) protein in any of the species but Rubisco activity was reduced by 35% in the weak CAM-inducible C. aripoensis. Similar amounts of glycine decarboxylase (GDC) protein were present in all three species regardless of the magnitude of CAM expression. Thus, the constitutive CAM species C. rosea did not appear to show reduced activity of this key enzyme of the photorespiratory pathway, which, in turn, may be related to the low internal conductance to CO₂ in this succulent species. Immuno-histochemical techniques showed that PEPCase, PEPCK and Rubisco were present in cells of the palisade and spongy parenchyma in leaves of species performing CAM. However, in leaves from well-watered plants of C. aripoensis which only performed C₃ photosynthesis, PEPCK was localized around latex-producing ducts. Differences in leaf anatomy between the species suggest that the association between mesophyll succulence and the capacity for CAM in these hemi-epiphytic stranglers has been selected for in arid environments. Received: 4 July 1997 / Accepted: 27 November 1997     

毛竹的无性系生长与立竹密度和叶龄结构的关系

毛竹为单轴型散生竹,属典型的无性系植物,原产我国亚热带地区。由于其个体高大、生长迅速、产量高、材质好、分布广,长期以来,一直是我国最为重要的经济竹种。本文应用无性系生长生理整合的理论,从种群统计学的角度,探讨了毛竹林立竹密度与叶龄结构对其无性系生长潜力的影响。结果表明：由于毛竹叶的生活期为两年,１龄新叶的光合能力比２龄老叶高,每样地的出笋数、活笋数与带１龄新叶的立竹数呈正相关,而与带２龄老叶的立竹数相关性不显著。另外,竹笋的死亡率是非密度制约的。本研究结果合理地解释了常见的毛竹林产量大小年交替变化的现象。     

Assimilate Transport in the Xylem Sap of Pedunculate Oak (Quercus robur) Saplings

Abstract: The rates of photosynthesis and transpiration, as well as the concentrations of organic compounds (total soluble non-protein N compounds [TSNN], soluble carbohydrates), in the xylem sap were determined during two growth seasons in one-year-old Quercus robur saplings. From the data, the total C gain of the leaves, by both photosynthesis and the transpiration stream, was calculated. Large amounts of C were allocated to the leaves by the transpiration stream; depending on the time of day and the environmental conditions the portion of C originating from xylem transport amounted to 8 to 91% of total C delivery to the leaves. Particularly under conditions of reduced photosynthesis, e.g., during midday depression of photosynthesis, a high percentage of the total C delivery was provided to the leaves by the transpiration stream (83 to 91 %). Apparently, attack by phloem-feeding aphids lowered the assimilate transport from roots to shoots; as a consequence the portion of C available to the leaves from xylem transport amounted to only 12 to 16 %. The most abundant organic compounds transported in the xylem sap were sugars (sucrose, glucose, fructose) with concentrations of ca. 50 to 500 μmol C ml^-1, whereas C from N compounds was of minor significance (3 to 20 μmol ml^-1 C). The results indicate a significant cycling of C in the plants because the daily transport of C with the transpiration stream exceeded the daily photosynthetic CO₂ fixation in several cases. This cycling pool of C may sustain delivery of photosynthate to heterotrophic tissues, independent of short time fluctuations in photosynthetic CO₂ fixation.     

The role of sink demand in carbon partitioning and photosynthetic reinvigoration following shoot decapitation

Photosynthesis, growth, and carbon partitioning of vigorous coppice shoots were compared with the slower growing intact shoots of Populus maximowiczii × nigra L. MN9 to determine the relationship between carbon partitioning and photosynthetic rate. Relative height growth rate of coppice shoots was 2.2 times that of intact shoots with net photosynthetic rate 1.9 times that of intact shoots. Coppice leaves exported a larger proportion of newly-fixed assimilate (11% compared with 6%) after a 4-h chase. The greater export from coppice leaves was correlated with a greater proportion of [¹⁴C]-labelled photosynthate deposited as starch in stems 4 cm below the point of label application. Coppice leaf assimilate levels were reduced to 15% that of leaves on intact plants, but coppice leaves had twice the concentration of labelled sucrose. Carbohydrates constituted 55% of the water-soluble [¹⁴C]-labelled photosynthate in leaves of coppice shoots compared with 40% in intact shoots. The results suggest that carbon allocation and partitioning in coppice shoots were altered towards production and export of new assimilate, and support the hypothesis that photosynthetic rate is responsive to sink demand for assimilates.     

三峡库区岸生植物秋华柳对水淹的光合和生长响应

为阐明三峡库区岸生植物秋华柳(Salix variegata)对水淹的耐受机制,模拟三峡库区消落带水淹发生的情况,研究了在不同水淹时间和水淹深度处理下秋华柳的光合和生长特性。实验设置了对照(不进行水淹,常规供水管理)、水淹根部(植株置于水中,植株地下部分被淹没)、水下0.5 m(植株置于水中,植株顶部在水面下0.5 m)和水下2 m(植株置于水中,植株顶部在水面下2 m)4个不同的水淹深度和0、10、20、40、60和90 d 6个不同的水淹时间处理,并测定了在不同水淹时间和水淹深度处理下秋华柳的光合作用、叶绿素荧光和生长。研究结果发现:随着水淹时间的延长,对照和水淹根部植株都具有高的净光合速率、表观量子效率和羧化效率。水淹40 d后,相同水淹深度处理秋华柳植株的净光合速率显著高于耐水湿环境的垂柳(Salix babylonica)(p<0.05)。水淹90 d后,全淹处理植株的光合能力较对照有显著的下降(p<0.05),对照、水下0.5 m和水下2 m植株的净光合速率分别为13.2、10.1和8.05 μmol·m^-2·s^-1,同时全淹植株PSII的最大光化学效率也有一定程度的下降,显著低于对照和水淹根部处理的植株(p<0.05)。水淹40、60和90 d后,全淹植株的胞间CO₂浓度都高于对照和水淹根部植株。随着水淹时间的增加,水淹根部植株不定根数量不断增加,而全淹植株只有极少量的不定根产生。水淹根部植株的主茎长的增量、分枝数的增量、主茎新生叶片数、根生物量的积累和总生物量的积累都高于全淹植株,全淹植株在水淹过程中,其主茎长、分枝数、主茎新叶数、根生物量和总生物量都有增加,同时其凋落叶片较多。水淹90 d后,秋华柳植株的存活率为100%。研究结果表明,秋华柳在经过较长时间的水淹后,表现出较强的光合和生长适应性,可以考虑将秋华柳列为三峡库区消落带植被构建的物种之一。     

Root distribution and seasonal production in the northwestern Sonoran Desert for a C3 subshrub, a C4 bunchgrass, and a CAM leaf succulent

To investigate root distribution with depth, which can affect competition for water, surface areas of young and old roots were determined in 4-cm-thick soil layers for the C3 subshrub Encelia farinosa Torrey and A. Gray, the C4 bunchgrass Pleuraphis rigida Thurber, and the CAM (crassulacean acid metabolism) leaf succulent Agave deserti Engelm. At a site in the northwestern Sonoran Desert these codominant perennials had mean rooting depths of only 9-10 cm for isolated plants. Young roots had mean depths of 5-6 cm after a winter wet period, but 11-13 cm after a summer wet period. Young roots were most profuse in the winter for E. farinosa, which has the lowest optimum temperature for root growth, and in the summer for P. rigida, which has the highest optimum temperature. Roots for interspecific pairs in close proximity averaged 2-3 cm shallower for A. deserti and a similar distance deeper for the other two species compared with isolated plants, suggesting partial spatial separation of their root niches when the plants are in a competitive situation. For plants with a similar root surface area, the twofold greater leaf area and twofold higher maximal transpiration rate of E. farinosa were consistent with its higher root hydraulic conductivity, leading to a fourfold higher estimated maximal water uptake rate than for P. rigida. Continuous water uptake accounted for the shoot water loss by A. deserti, which has a high shoot water-storage capacity. A lower minimum leaf water potential for P. rigida than for A. deserti indicates greater ability to extract water from a drying soil, suggesting that temporal niche separation for water uptake also occurs.     

Causes and consequences of high osmotic potentials in epiphytic higher plants

Past reports of the water relations of epiphytes, particularly bromeliads, indicate that tissue osmotic potentials in these tropical and subtropical plants are very high (close to zero) and are similar to values for aquatic plants. This is puzzling because several ecophysiological studies have revealed a high degree of drought stress tolerance in some of these epiphytes. The goal of this study was two-fold: (1) to increase the number of epiphytic taxa sampled for tissue osmotic potentials; and (2) to explain the apparent discrepancy in the significance of the tissue water relations and tolerance of drought stress in epiphytes. Tissue osmotic potentials of 30 species of epiphytic ferns, lycophytes, and orchids were measured in a subtropical rain forest in northeastern Taiwan. Nearly all values were less negative than -1.0 MPa, in line with all previous data for epiphytes. It is argued that such high osmotic potentials, indicative of low solute concentrations, are the result of environmental constraints of the epiphytic habitat on productivity of these plants, and that low rates of photosynthesis and transpiration delay the onset of turgor loss in the tissues of epiphytes such that they appear to be very drought-stress tolerant. Maintenance of photosynthetic activity long into drought periods is ascribed to low rates of transpiration and, hence, delayed tissue desiccation, and hydration of the photosynthetic tissue at the expense of water from the water-storage parenchyma.     

Nitrogen allocation in response to partial shading of a plant: Possible mechanisms

The significance of photosynthetic and transpiration rates for the perception by plants of light gradients in leaf canopies has been investigated with regard to nitrogen allocation and re-allocation. A gradient of photon flux density (PFD) over a plant's foliage was simulated by shading one leaf of a pair of primary leaves of bean ( Phaseolus vulgaris L. cv. Rentegever). Photosynthetic rate was manipulated independently of PFD and, to some extent, also of transpiration, by subjecting the leaf to different CO₂ concentrations. Transpiration rate was changed independently of PFD and photosynthetic rate by subjecting the leaf to different vapour pressure differences (VPD). A reduced partial pressure of CO₂ reduced specific leaf mass (SLM) as did a decreased PFD, but did not change leaf N per unit area (N_LA) and light saturated rate of photosynthesis (A_max). A reduced VPD caused several effects consistent with the effect of PFD. It decreased N_LA and A_max and increased the chlorophyll to N ratio in old and young leaves. Furthermore, it decreased the chlorophyll a to b ratio and inhibited leaf growth in young leaves. The transpiration stream is partitioned among the leaves of a plant according to their transpiration rates. The results suggest that relative rates of import of xylem sap into leaves of a plant play an important role in the perception of partial shading of a plant, a situation normally found in dense vegetations. The possible role of cytokinin influx into leaves as controlled by transpiration rate, is discussed.