Ecophysiological studies of sonoran Desert plants

Summary Photosynthetic responses to light and temperature are compared for two genetically related chromosomal races of Machaeranthera gracilis. The ancestral foothills race occurs in cooler, more mesic environments, while the derived desert race occurs in more arid environments. The desert race exhibited greater rates of net photosynthesis at all levels of irradiance than did plants of the foothills race. This enhancement is due primarily to a greater quantum yield and leaf density. High light pretreatment significantly increased the photosynthetic capacity of the desert race, with little or no effect on the foothills race. Furthermore, the leaf density of the desert race was affected proportionately more than the foothills race by light pretreatment. The desert race also possessed higher stomatal and mesophyll conductances to CO₂. Both races exhibited enhanced photosynthetic capacities when grown in a warm thermoperiod (35/25 C), relative to a cool thermoperiod (25/15 C), concomitant with slight increases in leaf density. There was a lack of thermal acclimation for both photosynthesis and respiration of the two races. The greater photosynthetic capactity of the desert race is suggested as an evolutionary modification which could account for invasion into and survival in the sonoran Desert.     

Life cycle characteristics of Machaeranthera gracilis (Compositae) in desert habitats

Summary Plant population dynamics and life cycle characteristics are presented for two genetically related, yet ecologically distinct, chromosomal races of Machaeranthera gracilis (Nutt.) Shinners. Seedling mortality was very high for a field population of the desert race (n=2) as only 8% of the maximum number of seedlings produced did survive to the period of peak reproduction. Lower mortality occurred in the foothills race (n=4) as 15% of the maximum number of seedlings produced did survive until the period of peak reproduction. The desert race also exhibited the capacity to assume the perennial growth habit. Perennial plants of the desert race produced 4.4 heads/plant (116 heads/m²) versus 0.2 heads/plant (4 heads/m²) for annual plants of the desert race under field conditions. Annual plants of the foothills race produced 2.7 heads/plant (75 heads/m²). The perennial habit did not occur in field populations of the foothills race. When grown in identical glasshouse conditions the desert race reached anthesis of the first four flower heads ca. 25 days prior to the foothills race. This acceleration to anthesis was accompanied by a greater potential reproductive allocation. The latter response was due in part to greater biomass allocation to reproductive structures at the expense of vegetative structures, and the uncoupling of net photosynthesis from phenological events.     

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     

Light,temperature and tocopherol status influence foliar vascular anatomy and leaf function in Arabidopsis thaliana

This study addressed whether the winter annual Arabidopsis thaliana can adjust foliar phloem and xylem anatomy both differentially and in parallel. In plants acclimated to hot vs cool temperature, foliar minor vein xylem‐to‐phloem ratio was greater, whereas xylem and phloem responded concomitantly to growth light intensity. Across all growth conditions, xylem anatomy correlated with transpiration rate, while phloem anatomy correlated with photosynthetic capacity for two plant lines (wild‐type Col‐0 and tocopherol‐deficient vte1 mutant) irrespective of tocopherol status. A high foliar vein density (VD) was associated with greater numbers and cross‐sectional areas of both xylem and phloem cells per vein as well as higher rates of both photosynthesis and transpiration under high vs low light intensities. Under hot vs cool temperature, high foliar VD was associated with a higher xylem‐to‐phloem ratio and greater relative rates of transpiration to photosynthesis. Tocopherol status affected development of foliar vasculature as dependent on growth environment. The most notable impact of tocopherol deficiency was seen under hot growth temperature, where the vte1 mutant exhibited greater numbers of tracheary elements (TEs) per vein, a greater ratio of TEs to sieve elements, with smaller individual sizes of TEs, and resulting similar total areas of TEs per vein and transpiration rates compared with Col‐0 wild‐type. These findings illustrate the plasticity of foliar vascular anatomy acclimation to growth environment resulting from independent adjustments of the vasculature's components.     

Photosynthetic compensation by the reproductive structures in the spring ephemeral Gagea lutea

Growth and reproduction of spring ephemerals inhabiting deciduous forests progress simultaneously during a short period from snowmelt to canopy closure. To clarify the mechanism to mitigate the cost of reproduction, contributions of foliar and non-foliar photosynthetic products to seed production were examined in a spring ephemeral Gagea lutea. Leaf growth, foliar and non-foliar photosynthetic activities, and total assimilated products were compared among reproductive-intact, floral bud-removal, and vegetative plants. Translocation of current photosynthetic products to individual organs was quantified by ¹³CO₂-trace experiment. Bulb growth was compared between hand-pollination and floral bud-removal treatments. Finally, seed set was compared between intact, leaf-clipping, and bract-clipping treatments. Fruit-forming plants retained leaves longer than vegetative and floral bud-removal plants, but the assimilative contribution of extended leaf longevity was negligible. Carbon supply by bract photosynthesis was large enough for fruit development, while carbon supply by fruit photosynthesis was offset by the high respiration loss. Foliar photosynthetic products were largely transported to bulbs, while translocation to reproductive functions was negligible. Because the floral bud-removal increased the bulb growth, lack of reproduction could lead to more storage. The leaf-clipping had no effect on seed production, while the bract-clipping significantly reduced the seed production. Therefore, current photosynthesis of leafy bracts might be a major carbon source for fruit development. This self-compensative mechanism of reproductive structure enables the continuous reproductive activity in this species.     

Shoot growth dynamics and photosynthetic response to increased nitrogen availability in the alpine willow Salix glauca

Plants subjected to increases in the supply of resource(s) limiting growth may allocate more of those resources to existing leaves, increasing photosynthetic capacity, and/or to production of more leaves, increasing whole-plant photosynthesis. The responses of three populations of the alpine willow, Salix glauca, growing along an alpine topographic sequence representing a gradient in soil moisture and organic matter, and thus potential N supply, to N amendments, were measured over two growing seasons, to elucidate patterns of leaf versus shoot photosynthetic responses. Leaf-(foliar N, photosynthesis rates, photosynthetic N-use efficiency) and shoot-(leaf area per shoot, number of leaves per shoot, stem weight, N resorption efficiency) level measurements were made to examine the spatial and temporal variation in these potential responses to increased N availability. The predominant response of the willows to N fertilization was at the shoot-level, by production of greater leaf area per shoot. Greater leaf area occurred due to production of larger leaves in both years of the experiment and to production of more leaves during the second year of fertilization treatment. Significant leaflevel photosynthetic response occurred only during the first year of treatment, and only in the dry meadow population. Variation in photosynthesis rates was related more to variation in stomatal conductance than to foliar N concentration. Stomatal conductance in turn was significantly related to N fertilization. Differences among the populations in photosynthesis, foliar N, leaf production, and responses to N fertilization indicate N availability may be lowest in the dry meadow population, and highest in the ridge population. This result is contrary to the hypothesis that a gradient of plant available N corresponds with a snowpack/topographic gradient.     

Non-structural carbohydrate dynamics associated with antecedent stem water potential and air temperature in a dominant desert shrub

Non-structural carbohydrates (NSCs) are necessary for plant growth and affected by plant water status, but the temporal dynamics of water stress impacts on NSC are not well understood. We evaluated how seasonal NSC concentrations varied with plant water status (predawn xylem water potential, Ψ) and air temperature (T) in the evergreen desert shrub Larrea tridentata. Aboveground sugar and starch concentrations were measured weekly or monthly for ~1.5 years on 6–12 shrubs simultaneously instrumented with automated stem psychrometers; leaf photosynthesis (A_net) was measured monthly for 1 year. Leaf sugar increased during the dry, premonsoon period, associated with lower Ψ (greater water stress) and high T. Leaf sugar accumulation coincided with declines in leaf starch and stem sugar, suggesting the prioritization of leaf sugar during low photosynthetic uptake. Leaf starch was strongly correlated with A_net and peaked during the spring and monsoon seasons, while stem starch remained relatively constant except for depletion during the monsoon. Recent photosynthate appeared sufficient to support spring growth, while monsoon growth required the remobilization of stem starch reserves. The coordinated responses of different NSC fractions to water status, photosynthesis, and growth demands suggest that NSCs serve multiple functions under extreme environmental conditions, including severe drought.     

Seasonal response of photosynthesis to elevated CO2 in loblolly pine (Pinus taeda L.) over two growing seasons

Trees growing in natural systems undergo seasonal changes in environmental factors that generate seasonal differences in net photosynthetic rates. To examine how seasonal changes in the environment affect the response of net photosynthetic rates to elevated CO₂, we grew Pinus taeda L. seedlings for three growing seasons in open-top chambers continuously maintained at either ambient or ambient + 30 Pa CO₂. Seedlings were grown in the ground, under natural conditions of light, temperature nd nutrient and water availability. Photosynthetic capacity was measured bimonthly using net photosynthetic rate vs. intercellular CO₂ partial pressure (A-C_i) curves. Maximum Rubisco activity (Vc_max) and ribulose 1,5-bisphosphate regeneration capacity mediated by electron transport (J_max) and phosphate regeneration (PiRC) were calculated from A-C_i curves using a biochemically based model. Rubisco activity, activation state and content, and leaf carbohydrate, chlorophyll and nitrogen concentrations were measured concurrently with photosynthesis measurements. This paper presents results from the second and third years of treatment. Mean leaf nitrogen concentrations ranged from 13.7 to 23.8 mg g^?1, indicating that seedlings were not nitrogen deficient. Relative to ambient CO₂ seedlings, elevated CO₂ increased light-saturated net photosynthetic rates 60–110% during the summer, but < 30% during the winter. A relatively strong correlation between leaf temperature and the relative response of net photosynthetic rates to elevated CO₂ suggests a strong effect of leaf temperature. During the third growing season, elevated CO₂ reduced Rubisco activity 30% relative to ambient CO₂ seedlings, nearly completely balancing Rubisco and RuBP-regeneration regulation of photosynthesis. However, reductions in Rubisco activity did not eliminate the seasonal pattern in the relative response of net photosynthetic rates to elevated CO₂. These results indicate that seasonal differences in the relative response of net photosynthetic rates to elevated CO₂ are likely to occur in natural systems.     

Hydraulic limitation not declining nitrogen availability causes the age‐related photosynthetic decline in loblolly pine (Pinus taeda L.)

Declining net primary production (NPP) with forest age is often attributed to a corresponding decline in gross primary production (GPP). We tested two hypotheses explaining the decline of GPP in ageing stands (14–115 years old) of Pinus taeda L.: (1) increasing N limitation limits photosynthetic capacity and thus decreases GPP with increasing age; and (2) hydraulic limitations increasingly induce stomatal closure, reducing GPP with increasing age. We tested these hypotheses using measurements of foliar nitrogen, photosynthesis, sap‐flow and dendroclimatological techniques. Hypothesis (1) was not supported; foliar N retranslocation did not increase and declines were not observed in foliar N, leaf area per tree or photosynthetic capacity. Hypothesis (2) was supported; declines were observed in light‐saturated photosynthesis, leaf‐ and canopy‐level stomatal conductance, concentration of CO₂ inside leaf air‐spaces (corroborated by an increase in wood δ¹³C) and specific leaf area (SLA), while stomatal limitation and the ratio of sapwood area (SA) to leaf area increased. The sensitivity of radial growth to inter‐annual variation in temperature and drought decreased with age, suggesting that tree water use becomes increasingly conservative with age. We conclude that hydraulic limitation increasingly limits the photosynthetic rates of ageing loblolly pine trees, possibly explaining the observed reduction of NPP.     

Effects of desiccation and CO2 concentrations on emersed photosynthesis in Porphyra haitanensis (Bangiales,Rhodophyta), a species farmed in China

The effects on photosynthesis of CO₂ and desiccation in Porphyra haitanensis were investigated to establish the effects of increased atmospheric CO₂ on this alga during emersion at low tides. With enhanced desiccation, net photosynthesis, dark respiration, photosynthetic efficiency, apparent carboxylating efficiency and light saturation point decreased, while the light compensation point and CO₂ compensation point increased. Emersed net photosynthesis was not saturated by the present atmospheric CO₂ level (about 350?ml?m^?3), and doubling the CO₂ concentration (700?ml?m^?3) increased photosynthesis by between 31% and 89% at moderate levels of desiccation. The relative enhancement of emersed net photosynthesis at 700?ml?m^?3 CO₂ was greater at higher temperatures and higher levels of desiccation. The photosynthetic production of Porphyra haitanensis may benefit from increasing atmospheric CO₂ concentration during emersion.     

PHOTOINHIBITION AND PHOTOACCLIMATION OF TURF ALGAL COMMUNITIES ON A TEMPERATE REEF,AFTER IN SITU TRANSPLANTATION EXPERIMENTS1

The photophysiology of turf algal communities was studied in situ on a temperate reef off the coast of South Australia. Algal communities were grown on artificial substrate at depths of 2, 4, and 10 m. To investigate the response of the algal communities to changing light environments in both the short and long term, reciprocal transplantation experiments were conducted among these depths on a seasonal basis. The extent of photoinhibition was assessed every 3 h for the first 2 days following transplantation and then on a daily basis for 16 days after transplantation. Photosynthetic acclimation was assessed using photosynthesis–light curves obtained from transplanted and non‐transplanted turfs after the acclimation period. Transplanted turfs responded very quickly to the light shift. Algae acclimated to low light (10 m depth) were highly susceptible to photoinhibition and photodamage, having greater decreases in maximum and effective quantum yields than turfs from shallower depths. Yield recovery and acclimation usually occurred very rapidly in algae from all depths (3–5 days), but were faster in spring and summer compared with winter. Changes in photosynthetic capacity (across seasons, depths, and after transplantation to a different depth) were accompanied by changes in respiration, so that the ratio of net to gross photosynthetic capacity (P_mnet : P_mgross) remained high and constant over the whole range of light levels. We discuss the possible acclimation strategies of turfs, taking into account the balance between photoacclimation, production, and growth strategy.     

Seasonal variation in foliar carbon exchange in Pinus radiata and Populus deltoides: respiration acclimates fully to changes in temperature but photosynthesis does not

We investigated seasonal variation in dark respiration and photosynthesis by measuring gas exchange characteristics on Pinus radiata and Populus deltoides under field conditions each month for 1 year. The field site in the South Island of New Zealand is characterized by large day-to-day and seasonal changes in air temperature. The rate of foliar respiration at a base temperature of 10 °C ( R ₁₀) in both pine and poplar was found to be greater during autumn and winter and displayed a strong downward adjustment in warmer months. The sensitivity of instantaneous leaf respiration to a 10 °C increase in temperature ( Q ₁₀) was also greater during the winter period. The net effect of this strong acclimation was that the long-term temperature response of respiration was essentially flat over a wide range of ambient temperatures. Seasonal changes in photosynthesis were sensitive to temperature but largely independent of leaf nitrogen concentration or stomatal conductance. Over the range of day time growth temperatures (5–32 °C), we did not observe strong evidence of photosynthetic acclimation to temperature, and the long-term responses of photosynthetic parameters to ambient temperature were similar to previously published instantaneous responses. The ratio of foliar respiration to photosynthetic capacity ( R _d/ A _sat) was significantly greater in winter than in spring/summer. This indicates that there is little likelihood that respiration would be stimulated significantly in either of these species with moderate increases in temperature – in fact net carbon uptake was favoured at moderately higher temperatures. Model calculations demonstrate that failing to account for strong thermal acclimation of leaf respiration influences determinations of leaf carbon exchange significantly, especially for the evergreen conifer.     

Short-term effects of fertilization on loblolly pine (Pinus taeda L.) physiology

Fertilization commonly increases biomass production in loblolly pine (Pinus taeda L.). However, the sequence of short‐term physiological adjustments allowing for the establishment of leaf area and enhanced growth is not well understood. The effects of fertilization on photosynthetic parameters, root respiration, and growth for over 200 d following the application of diammonium phosphate were intensively investigated in an effort to establish a relative sequence of events associated with improved growth. Root respiration, foliar nitrogen concentration [N]_f, and light‐saturated net photosynthesis (A_sat) temporarily increased following fertilization. A_sat was correlated positively with [N]_f when non‐fertilized and fertilized treatments were pooled (R² = 0.47). Increased photosynthetic capacity following fertilization was due to both improved photochemical efficiency and capacity and enhanced carboxylation capacity of Rubisco. Positive effects of fertilization on growth were observed shortly after A_sat increased. Fertilized seedlings had 36.5% more leaf area and 36.5% greater total dry weight biomass at 211 d following fertilization. It is concluded that fertilization temporarily increased photosynthetic capacity, which resulted in a pool of photo‐assimilate used to build leaf area. The N from fertilizer initially invested in photosynthetic structures and enzymes probably re‐translocated to newly developing foliage, explaining the reduction in [N]_f and A_sat that was observed after peak levels were achieved following fertilization.     

Carbon balance,productivity, and water use of cold-winter desert shrub communities dominated by C3 and C4 species

Summary Common generalizations concerning the ecologic significance of C₄ photosynthesis were tested in a study of plant gas exchange, productivity, carbon balance, and water use in monospecific communities of C₃ and C₄ salt desert shrubs. Contrary to expectations, few of the hypotheses concerning the performance of C₄ species were supported. Like the C₃ species, Ceratoides lanata, the C₄ shrub, Atriplex confertifolia, initiated growth and photosynthetic activity in the cool spring months and also exhibited maximum photosynthetic rates at this time of year. To compete successfully with C₃ species, Atriplex may have been forced to evolve the capacity for photosynthesis at low temperatures prevalent during the spring when moisture is most abundant. Maximum photosynthetic rates of Atriplex were lower than those of the C₃ species. This was compensated by a prolonged period of low photosynthetic activity in the dry late summer months while Ceratoides became largely inactive. However, the annual photosynthetic carbon fixation per ground area was about the same in these two communities composed of C₃ and C₄ shrubs. The C₄ species did not exhibit greater leaf diffusion resistance than the C₃ species. The photosynthesis/transpiration ratios of the two species were about the same during the period of maximum photosynthetic rates in the spring. During the warm summer months the C₄ species did have superior photosynthesis/transpiration ratios. Yet, since Ceratoides completed a somewhat greater proportion of its annual carbon fixation earlier in the season, the ratio of annual carbon fixation/transpiratory water loss in the two communities was about the same. Atriplex did incorporate a greater percentage of the annual carbon fixation into biomass production than did Ceratoides. However, this is considered to be a reflection of properties apart from the C₄ photosynthetic pathway. Both species displayed a heavy commitment of carbon to the belowground system, and only about half of the annual moisture resource was utilized in both communities.     

Physiological behaviors of <Emphasis Type="Italic">Acer mono</Emphasis> under drought and low light

The seedlings of Acer mono Maxim. were exposed to two watering regimes (well watered (100% of field capacity) and drought (30% of field capacity)) and two light levels (high light (100% of full sunlight) and low light (15% of full sunlight)) in a greenhouse to assess growth, photosynthesis, and foliar nutrient traits of woody plants under drought and low light. Drought significantly reduced growth and gas exchange characteristics of A. mono, including net photosynthetic rate (P _N), stomatal conductance (g _s), intercellular CO₂ concentration (C_i), and photosynthetic nitrogen use efficiency (PNUE). Also, drought reduced relative water content (RWC) and foliar C and N concentrations, but increased the C/N ratio. P _N and C concentration were lower under drought and low light than in any other treatment, indicating that deep shade might seriously decrease C assimilation. However, the negative effect induced by drought was alleviated by improving RWC and maintaining C balance, and therefore low-light seedlings accumulated more biomass than those under high light when they were exposed to drought. Our results showed that trade-off and facilitation effects of drought and low light might be complementary and alter in different species.     

COMPARISONS OF SALT-MARSH FUCOID PRODUCTION ESTIMATED FROM THREE DIFFERENT INDICES1

Production of Ascophyllum nodosum (L.) LeJolis ecads and Fuscus vesiculosus L. was calculated from measurements of in situ growth, seasonal variations in standing-crops and seasonal variations in photosynthetic capacity. A computer model for predicting daily, monthly and yearly net production from photosynthesis data was constructed. This model used daily irradiation, actual biomass of algae/m² contributing to production and photosynthesis vs. light intensity relationships as data inputs. Comparison of production estimated from in situ growth, standing-crops and photosynthesis indicated that both marsh fucoids turn over biomass twice per year. Total net production of both fucoids, estimated from photosynthesis data, was ca. 315 g C · m^?2· yr^?1. On the other hand, production of both fucoids calculated from standing-crop data was only 155 g C · m^?2· yr^?1.     

COTYLEDON PHOTOSYNTHESIS DURING SEEDLING GROWTH OF COTTON,GOSSYPIUM HIRSUTUM L.

Respiration and net photosynthetic O₂ production by cotton cotyledons were determined from an early age through the senescent stage. Various treatments were applied to cotyledons to assess the importance of current photosynthesis as compared to translocation of reserves to seedling development. Rates of respiration and net photosynthesis per cm² were high on 1-day-old cotyledons, but the rates decreased sharply with rapid expansion to reach a fairly stable rate. Respiration per cotyledon decreased linearly with age until the onset of senescence, then exhibited a distinct climacteric rise followed by a sharp decrease. Net photosynthesis per cotyledon increased until expansion was completed and then decreased linearly and steeply with age. Excision of cotyledons, inhibition of photosynthesis either chemically or by covering, and removal of the terminal bud indicated that current photosynthesis is a potent force behind early epicotyl growth.     

Contribution of stem CO2 fixation to whole-plant carbon balance in nonsucculent species

In many plant species that remain leafless part of the year, CO₂ fixation occurring in green stems represents an important carbon gain. Traditionally, a distinction has been made between stem photosynthesis and corticular photosynthesis. All stem photosynthesis is, sensu stricto, cortical, since it is carried out largely by the stem cortex. We proposed the following nomenclature: stem net photosynthesis (SNP), which includes net CO₂ fixation by stems with stomata in the epidermis and net corticular CO₂ fixation in suberized stems, and stem recycling photosynthesis (SRP), which defines CO₂ ling in suberized stems. The proposed terms should reflect differences in anatomical and physiological traits. SNP takes place in the chlorenchyma below the epidermis with stomata, where the net CO₂ uptake occurs, and it resembles leaf photosynthesis in many characteristics. SRP is found in species where the chlorenchyma is beneath a well-developed stomata-free periderm and where reassimilation of internally respired CO₂ occurs. SNP is common in plants from desert ecosystems, rates reaching up to 60% of the leaf photosynthetic rate. SRP has been demonstrated in trees from temperate forests and it offsets partially a carbon loss by respiration of stem nonphotosynthetic tissues. Reassimilation can vary between 7 and 123% of respired CO₂, the latter figure implying net CO₂ uptake from the atmosphere. Both types of stem photosynthesis contribute positively to the carbon economy of the species, in which they occur; they are advantageous to the plant because they allow the maintenance of physiological activity during stress, an increase of integrated water use efficiency, and they provide the carbon source used in the production of new organs.     

Influence of Thiourea on Photosynthesis, Nitrogen Metabolism and Yield of Clusterbean (Cyamopsis tetragonoloba (L.) Taub.) under Rainfed Conditions of Indian Arid Zone

A 2-year field trial was conducted to study the effects of seed treatment (500 ppm) or foliar application of 1000 ppm thiourea (at 25 and 40 days after sowing) or a combination of these, on growth, yield, net photosynthesis and nitrogen metabolism of clusterbean (Cyamopsis tetragonoloba (L.) Taub.) grown for two consecutive years (1999 and 2000) under rainfed conditions of the Indian arid zone. Thiourea application either as pre-sowing seed treatment or as foliar spray significantly increased plant height, leaf area, dry matter production and seed yield as compared to the untreated control plants during both the years. However, maximum favourable effects were obtained with combined application of seed treatment and foliar spray. The beneficial effects of thiourea were attributed to its role in significantly increasing the net photosynthetic rates and the concentrations of total chlorophyll and starch in the leaves. Thiourea also reflected a positive role in enhancing nitrogen metabolism as it significantly increased nitrate reductase activity and concentration of soluble protein in the treated plants. It has been concluded that seed treatment with thiourea followed by foliar spray could significantly improve growth, yield and water use efficiency of rainfed clusterbean under arid conditions due to enhanced photosynthesis and more efficient nitrogen metabolism.     

Effect of soil temperature on stem sap flow,shoot gas exchange and water potential of Picea engelmannii (Parry) during snowmelt

Summary The effect of cold soils on stem sap flow, shoot gas exchange and water potential of Picea engelmannii (Parry) was investigated during the snowmelt period in the Medicine Bow Mountains, Wyoming, USA. Shoot net photosynthetic rates were higher in young trees (1.5–1.8 m in height) growing in cold soils (<3.5° C) associated with snowpack, than trees in warm soils until about 1500 h. Higher shoot photosynthetic rates of trees in cold soils continued after snow was removed and could not be completely explained by higher visible irradiance over highly reflective snow. Following soil warming higher photosynthetic rates were evident in these trees for five days. High nutrient availability associated with snowmelt may improve shoot nutrient status leading to higher gas-exchange rates during snowmelt. Shoot conductance to water vapor was higher in trees in cold soil until midday, when declining shoot conductance led to lower intercellular CO₂ concentrations. Midday through afternoon shoot water potentials of trees in cold soils were similar or higher than those of trees in warm soils and the lower afternoon shoot conductances in cold soils were not the result of lower bulk shoot water potentials. Decline in net photosynthesis of trees in cold soils at 1500 h paralleled increases in intercellular CO₂ concentrations, implying a nonstomatal limitation of photosynthesis. This scenario occurred consistently in mid-afternoon following higher morning and midday photosynthesis in cold soils, suggesting a carbohydrate feedback inhibition of photosynthesis. Diurnal patterns in stem sap flow of all trees (cold and warm soils) reflected patterns of shoot conductance, although changes in stem sap flow lagged 1–3 h behind shoot conductance apparently due to stem water storage. Total daily stem sap flow was similar in trees in cold and warm soils, although diel patterns differed. The morning surge and night-time drop in sap flow commenced 1–2 h earlier in trees in cold soils. Overnight stem sap flow was lower in trees in cold soils, possibly due to higher resistance to root water uptake in cold soils, which may explain lower predawn shoot water potentials. However, midday shoot water potentials of trees in cold soils equalled or exceeded those of trees in warm soils. Higher resistance to root water uptake in P. engelmannii in cold soils was apparently overshadowed by transpirational forces and significant shoot water deficits did not develop.