Development of acute frost drought in Rhododendron ferrugineum at the alpine timberline

Summary Acute winter frost drought in Rhododendron ferrugineum at the alpine timberline was provoked by removing the snow that had covered the shrubs and by irradiating and rewarming the twigs in situ in a climatized chamber simulating snow melt conditions. Water potential decreased to -2.3 MPa within three days. After activation of photosynthesis on the 1st day, the COr-uptake decreased during the 2nd day and approached zero at the 3rd day. With advanced desiccation, the internal COr concentration of the leaves increased indicating an impairment of chloroplast functions. The procedure of the disturbances appears to be similar to that of other acute drought events.     

Changes in radial tree growth for Picea abies, Larix decidua, Pinus cembra and Pinus uncinata near the alpine timberline since 1750

Changes in radial growth of the four coniferous species growing in the French Alps near the upper treeline are investigated. Thirty-seven populations of Norway spruce [Picea abies (L.) Karst.], European larch (Larix decidua Mill.), Swiss stone pine (Pinus cembra L.) and mountain pine (Pinus uncinata Mill. ex Mirb.) were sampled by taking 1320 cores and analysing tree-ring widths. Sites were chosen in various climatic conditions (macroclimate and aspect) and on two kinds of bedrock in order to take into account the ecological behaviour of these species. Belledonne, Moyenne-Tarentaise, Haute-Maurienne and Briançonnais areas were sampled along increasing gradients of summer aridity and winter continentality. The calculation of time series after removing the age trend brings strong evidence for an increase in radial growth during the two last centuries, but with different stages and fluctuations for each species. This growth trend is significantly enhanced since 1860 for the spruce, and since 1920 for the two pine species. Furthermore, it also appears on Larix decidua with the same pattern despite periodical growth reduction due to attacks of the larch bud moth (Zeiraphera diniana Gn.). The analysis of ring-widths at a given cambial age reveals that this enhanced phenomenon is observed especially during the tree’s early years (25–75 years). The analysis of four regional climatic series, and three longer series of temperature (in farther single sites) reveals synchronous decadal fluctuations and an evident secular increase in minimum temperatures (especially in January and from July to October), that may be involved in tree-growth enhancement. Thermic amplitudes are significantly reduced during the whole growing period, what is more pronounced in Belledonne, the most oceanic region. Long term growth changes are well described by stepwise regression models, especially for the pine species. These models involved both a linear trend (CO₂ concentration or N-deposition) and low frequency of Turin monthly temperatures. However, they show different patterns than those observed from response functions at a yearly scale.     

The effects of wind and temperature on cuticular transpiration of Picea abies and Pinus cembra and their significance in dessication damage at the alpine treeline

Summary The importance of high winter winds and plant temperatures as causes of winter desiccation damage at the alpine treeline were studied in the Austrian Alps. Samples of 1- and 2-year twigs of Picea abies and Pinus cembra were collected from the valley bottom (1,000 m a.s.l.), forestline (1,940 m a.s.l.), kampfzone (2.090 m a.s.l.), wind-protected treeline (2,140 m a.s.l.), and wind-exposed treeline (2,140 m a.s.l.). Cuticular transpiration was measured at three different levels of wind speed (4, 10, and 15 ms^-1) and temperature (15°, 20°, and 25° C). At elevated wind speeds slight increases in water loss were observed, whereas at higher temperatures much greater increases occurred. Studies on winter water relations show a significant decline in the actual moisture content and osmotic potentials of twigs, especially in the kampfzone and at treeline. The roles of high winds and temperatures in depleting the winter water economy and causing desiccation damage in the alpine treeline environment are discussed.     

Hydraulic efficiency and safety of leader shoots and twigs in Norway spruce growing at the alpine timberline

Xylem within trees varies in its hydraulic efficiency and safety. Trees at the alpine timberline were expected to exhibit a hydraulic architecture protecting the leader shoot from winter embolism. Hydraulic and related anatomical parameters were compared as well as seasonal courses of winter embolism in leader shoots and twigs of Norway spruce trees growing at 2000 m. Leader shoots had a 1.4-fold higher specific hydraulic conductivity (ks) as well as a 4.9-fold higher leaf specific conductivity (kl) than side twigs. Vulnerability to drought-induced embolism was lower in leader shoots with a 50% loss of conductivity occurring at a water potential (Psi 50) 0.7 MPa lower than in twigs. Higher ks and kl were related to 1.2-fold wider tracheid diameters in leader shoots. Lower vulnerability corresponded to smaller pit dimensions but not to wood density. High ks and kl reflect the hydraulic dominance of the leader shoot, which is important for its water supply during summer. Low vulnerability protects the leader shoot from embolism during the winter season. In field measurements at the timberline during the winter of 2001/2002, conductivity losses of up to 56% were observed only in twigs while leader shoots showed little or no embolism. Results demonstrate that leader shoot xylem is both hydraulically efficient and safe.     

环境因子对辣椒光合与蒸腾特性的影响

采用灰色关联分析方法研究环境因子对辣椒(Capsicum annuum Linn.)光合和蒸腾特性的影响.结果表明,在强光照下影响净光合速率的环境因子大小顺序为相对湿度、CO2浓度、温度、光照强度、气孔导度;净光合速率与相对湿度、叶温呈显著的负相关,与CO2浓度呈极显著的正相关;蒸腾速率与气温呈极显著的正相关,与相对湿度呈极显著的负相关;水分利用效率与相对湿度呈极显著的正相关,与温度呈极显著的负相关.弱光照时影响净光合速率的环境因子大小顺序为光照强度、温度、相对湿度、CO2浓度;净光合速率与光照强度、CO2浓度呈显著的正相关,与相对湿度呈显著的负相关.在强光照下适当遮阳、灌水、增施CO2肥,在弱光照下减少荫蔽、通风透气、降低相对湿度、增加温度可促进辣椒光合作用,提高辣椒产量.     

A coupled model of stomatal conductance, photosynthesis and transpiration

A model that couples stomatal conductance, photosynthesis, leaf energy balance and transport of water through the soil–plant–atmosphere continuum is presented. Stomatal conductance in the model depends on light, temperature and intercellular CO₂ concentration via photosynthesis and on leaf water potential, which in turn is a function of soil water potential, the rate of water flow through the soil and plant, and on xylem hydraulic resistance. Water transport from soil to roots is simulated through solution of Richards’ equation. The model captures the observed hysteresis in diurnal variations in stomatal conductance, assimilation rate and transpiration for plant canopies. Hysteresis arises because atmospheric demand for water from the leaves typically peaks in mid‐afternoon and because of uneven distribution of soil matric potentials with distance from the roots. Potentials at the root surfaces are lower than in the bulk soil, and once soil water supply starts to limit transpiration, root potentials are substantially less negative in the morning than in the afternoon. This leads to higher stomatal conductances, CO₂ assimilation and transpiration in the morning compared to later in the day. Stomatal conductance is sensitive to soil and plant hydraulic properties and to root length density only after approximately 10 d of soil drying, when supply of water by the soil to the roots becomes limiting. High atmospheric demand causes transpiration rates, LE, to decline at a slightly higher soil water content, θ_s, than at low atmospheric demand, but all curves of LE versus θ_s fall on the same line when soil water supply limits transpiration. Stomatal conductance cannot be modelled in isolation, but must be fully coupled with models of photosynthesis/respiration and the transport of water from soil, through roots, stems and leaves to the atmosphere.     

Response of transpiration and photosynthesis to a transient change in illuminated foliage area for a Pinus radiata D. Don tree

Sudden but transient changes in the fraction or illuminated foliage area in a well-watered 7-year-old Pinus radiata D. Don tree were imposed by completely covering either the upper 22% or the lower 78% of the foliage for periods of up to 36 h. Measurements of transpiration flux density (E), tree conductance (g_t), stomatal conductance (g_s) and net photosynthesis (A) were made to test the hypothesis that compensatory responses would occur in the remaining illuminated foliage when the cover was installed. When the lower foliage was covered there was an immediate decrease in g_t. However, when tree conductance was normalized with respect to the illuminated leaf area (g_t'), it increased between 50 and 75%, depending on the value of air saturation deficit (D). The effect was also apparent from concurrent measurements of increases in g_s and A up to 59 and 24%, respectively, for needles in the top third of (he crown. When the cover was removed these effects were reversed. The changes in the lower foliage when the upper foliage was covered were much smaller. Changes in bulk needle water potential were small. It is suggested that the observed responses occurred because of a perturbation to the hydraulic pathway in the xylem that could have triggered the action of a chemical signal to regulate stomatal conductance and photosynthesis.     

Effects of guttation prevention on photosynthesis and transpiration in leaves of Alchemilla mollis

The ecophysiological function(s) and consequences of guttation, a phenomenon by which water is exuded by and accumulated as droplets along the leaf margins under high humidity in many plants that grow in wet soil, has been poorly studied and remains largely unknown. Thus, leaf gas exchange and chlorophyll fluorescence were examined, using two experimental approaches, in Alchemilla mollis plants under conditions that promoted guttation and those that prevented this phenomenon. Although results were variable, depending on the experimental approach, prevention of guttation effected reductions in photosynthesis and transpiration, as well as photochemical activity measured with fluorescence techniques. These findings lend partial support for a previously hypothesized function of guttation: prevention of excess water in leaves, yet they contradict those of several other studies. More work is required in order to adequately understand the function of guttation.     

Effect of water deficits on transpiration, photosynthesis and leaf conductance in cassava

Transpiration, net photosynthesis and leaf conductance decreased when leaf water potential dropped below -0.30 MPa. Both transpiration and net photosynthesis rates were considerably reduced before the leaves were visibly wilted at -0.95 MPa. Consequently, visual symptoms are unlikely to provide a useful index for characterizing water deficits in cassava ( Manihot esculenta Crantz cv. Llanera). Decreases in net photosynthesis closely followed decreases in transpiration and this suggests that stomatal closure controls both processes.     

Nonstomatal inhibition of photosynthesis by water stress. Reduction in photosynthesis at high transpiration rate without stomatal closure in field-grown tomato

Large underestimates of the limitation to photosynthesis imposed by stomata can occur because of an error in the standard method of calculating average substomatal pressures of carbon dioxide when heterogeneity of those pressures occurs across a leaf surface. Most gas exchange data supposedly indicating nonstomatal inhibition of photosynthesis by water stress could have this error. However, if no stomatal closure occurs, any reduction in photosynthesis must be due to nonstomatal inhibition of photosynthesis. Net carbon dioxide exchange rates and conductances to water vapor were measured under field conditions in upper canopy leaves of tomato plants during two summers in Beltsville, Maryland, USA. Comparisons were made near midday at high irradiance between leaflets in air with the ambient water vapor content and in air with a higher water content. The higher water content, which lowered the leaf to air water vapor pressure difference (VPD), was imposed either one half hour or several hours before measurements of gas exchange. In both seasons, and irrespective of the timing of the imposition of different VPDs, net photosynthesis increased 60% after decreasing the VPD from 3 to 1 kPa. There were no differences in leaf conductance between leaves at different VPDs, thus transpiration rates were threefold higher at 3 than at 1 kPa VPD. It is concluded that nonstomatal inhibition of photosynthesis did occur in these leaves at high transpiration rate.     

Effects of atmospheric humidity on net photosynthesis,transpiration, and stomatal resistance

Rhizomes ofHydrocotyle plants from three contrasting habitats were cloned and the ramets grown under controlled environmental conditions. Measurements of net photosynthesis, transpiration, and total leaf diffusion resistance were used to examine possible physiological adaptations to specific field environments. Increasing dryness of the growth chamber environment had large effects on gas exchange (CO₂ and water vapor) and on total diffusion resistance of plants from a pond, moderate effects on plants from a mesic forest, but plants from a coastal sand dune were unaffected by the experimentally imposed dryness. Thus the 3 Hydrocotyle types demonstrated adaptive physiological reponses to their specific field habitats. Periodic stomatal oscillations were induced in ramets from the pond by sharply increasing irradiance, but the adaptiveness of the oscillations cannot be determined with the evidence at hand.No stomatal closure could be induced by atmospheric dryness alone as long as soil and plant dessication were prevented. There were no observable differences in stomatal response to increasing atmospheric vapor pressure deficits.     

Comparative photosynthesis,growth and transpiration of two species of Atriplex

Summary Throughout a period of 23 days, during which the photosynthesis, growth and transpiration of two species of Atriplex were compared, A. spongiosa, a C₄ species (first products of photosynthesis = 4-C dicarboxylic acids), maintained net rates of leaf photosynthesis as high as, or higher than, those of A. hastata, a C₃ species (photosynthesis exhibiting the Calvin-type characteristics).However, as the experiment progressed, the proportion of photosynthate which was used to produce new leaf material declined progressively in A. spongiosa, so that total plant growth rate, initially more than twice as high as in A. hastata, declined to only 0.8 of the A. hastata value. This result demonstrated clearly that more efficient photosynthesis is only one factor, and in this case a relatively minor factor, in total growth rate.Transpiration rates were consistently lower in A. spongiosa than in A. hastata and the ratio declined slightly during the experiment. In consequence, water-use efficiency, both on a single-leaf and whole-plant basis, was much greater in the C₄ species.Levels of mesophyll resistance (r _m ) were consistently lower in A. spongiosa and increased from about 0.4–0.6 to 1.2–1.5 s cm^–1 during the experiment. In A. hastata there was more variability in r _m levels but little overall trend towards a higher r _m , initial and final values being of the order of 2.5–2.6 and 2.6–2.9 s cm^–1, respectively. Levels of stomatal resistance (r _l ) were higher in A. spongiosa (about 1.0–1.2 s cm^–1) than in A. hastata (about 0.7–0.8 s cm^–1) at the beginning of the experiment and increased to 2.0–2.6 s cm^–1, whereas they remained relatively constant in A. hastata.The combination of relatively low r _m levels and relatively high r _l levels provide the explanation for the substantially greater water use efficiency in A. spongiosa. The progressive changes in these levels and in the pattern of leaf area development in A. spongiosa provide an elegant example of adaptation to arid conditions by this species.     

The role of stem recharge in reducing the winter desiccation of Picea engelmannii (Pinaceae) needles at alpine timberline

The winter desiccation of needles is thought to limit tree growth and survival within alpine timberline ecotones of the southern Rocky Mountains, USA. To better understand the factors contributing to this desiccation damage, the extent to which stem water was available to needles of Picea engelmannii undergoing desiccation at timberline near Monarch Pass, Colorado, was monitored throughout the winter. Severed shoots experienced significantly greater desiccation than did intact shoots, indicating the availability of stem water to needles despite presumably frozen soil, roots, and stems. A model of water relations during winter predicted more extreme desiccation of severed shoots than observed. This suggests that one or more of the common assumptions concerning the winter water relations of timberline trees is in error. The influence of cold, dry conditions on the cuticular conductance of Picea engelmannii needles is not known and therefore not accounted for in current models of winter water relations. The assumption that cuticular conductance is not influenced by temperature or humidity is a likely source of error in such models.     

Effects of insect damage on photosynthesis,transpiration and SO2 uptake by sycamore

Summary The bilberry (Vaccinium myrtillus L.), is the dominant plant in the field layer of many boreal forests in northern Sweden. It is utilized by several herbivorous insect larvae as food (i.e. Lepidoptera: Geometridae, Tortricidae and Hymenoptera: Symphyta). Total density of insect larvae was 63% lower where birds had access to larvae compared to exclosures. Larvae with a feeding behaviour which exposes them to birds (geometrids and sawflies) showed a pronounced reduction in density when exposed to bird predators. Density of larvae with a concealed feeding behaviour (tortricids) was unaffected by birds. The frequency of larval damage to bilberry annual shoots was significantly higher in areas where larval density was high due to exclusion of birds. I concluded that birds have the potential to heavily modify the interaction between bilberry and insect larvae.     

Effects of polyethylene-glycol-induced osmotic stress on transpiration and photosynthesis in pinto bean leaf discs

A new leaf disc chamber allows measurements of chlorophyll fluorescence and CO₂ and H₂O vapor exchanges during infusion of solution into the cut edge of the disc. Polyethylene glycol (molecular weight, 6000) was used to apply a mild external osmotic stress to the leaf disc within this chamber. This stress rapidly caused a temporary increase in transpiration. This increase was reversed (5-6 minutes later) and after 20 to 25 min, the stomates nearly completely closed. Internal CO₂ (calculated) and leaf temperature followed the transpiration measurements. However, chlorophyll fluorescence (small rise) followed internal CO₂ (small rise). This complete sequence of events resembles those caused by exposure of leaves to certain air pollutants which have been seen to cause such a transient increase followed by a decrease in stomatal closure.     

Interannual variability of PSP outbreaks on the north east UK coast

Paralytic shellfish poisoning (PSP) occurs sporadically on theNB UK coast. The degree of toxicity shows considerable interannualvariability, but particularly severe events occurred in 1968and 1990. The time sequence of PSP toxin production in 1990is described and compared with 1989 when no significant PSPtoxin occurred. In 1990, PSP toxin was widespread in shellfishsamples taken on 300 km of coastline, from Berwick to Whitby,and toxin was present at high concentrations for >1 month.The distribution of Alerandrium tamarense cysts in the sedimentsis described. High concentrations were found in the Firth ofForth and also in a number of regions offshore of the Scottishand English coasts. A water transport model has been used toestimate back trajectories, with the aim of determining thesource of the A.tamarense bloom. The Firth of Forth has previouslybeen suggested as the seed bed for A.tamarense outbreaks inthe area, but the transport model clearly shows that A.tanwrensemoved inshore over a wide area in 1990; there was no singlesource of the bloom. Sea surface temperatures, estimated fromsatellite imagery, show that water temperatures were much higherat the end of April 1990, when the bloom occurred, than in 1989when PSP toxin incidence was very low. These conditions wouldhave resulted in early seasonal stratification and would havefavoured phytoplankton growth in the water column.