The dynamics of leaf extension in plants with diverse altitudinal ranges

Summary Rates of leaf extension have been studied with electronic auxanometers at mid-altitude in the Austrian Alps, where both low and high altitude species co-occur. The results demonstrate a clear differentiation in the temperature responses of extension between these two groups of species. For the low or mid-altitude species of Achillea millefolium, Agrostis stolonifera, Poa alpina and Rumex arifolius, the average rate of leaf extension increases from 0.1 to 0.4 mm h^-1 between 10 and 20° C. For the high-alpine species of Achillea erba-rotta ssp moschata, Poa alpina ssp vivipara and Polygonum viviparum the average rate of leaf extension was considerably lower from 0.016 to 0.064 mm h^-1, between 10 and 20° C.Leaf extension in the lowland species was not observed below an average temperature of about 5° C, whilst no limit was observed for the upland species, down to a temperature of about 0° C.In the cases of the dicotyledons that were studied, leaf plus petiole shrinkage was observed to occur, for as much as 2 to 4 h, during periods of high water vapour pressure deficits. This response was not observed for the monocotyledons.The observations of leaf extension show that daily totals of extension in species from high altitudies will be much less sensitive to day, to day variations in local climate than will the species from low altitudes. The lowland species will have higher rates of extension during clear and warm weather conditions but lower rates in cold, cloudy weather.     

Elevational and age gradients in hawaiian montane rainforest: foliar and soil nutrients

Summary Soils and plants were sampled along an elevational gradient from 265–1675 m on a 133-and a 3100-year-old lava flow on Mauna Loa, Hawai'i. Soil organic matter and nutrients accumulated more rapidly at low elevation on the young flow, but reached higher levels at higher elevation on the old flow. Foliar nitrogen and phosphorus concentrations were less and specific leaf weight greater for Metrosideros polymorpha leaves collected at high versus low elevations and on the young versus the old flow. Foliar ¹³C was strongly correlated with specific leaf weight across the range of sites sampled.Published as a contribution to the Tropical Mountain Ecosystem Program of the IUBS Decade of the Tropics     

色季拉山急尖长苞冷杉叶片及细根性状随海拔的变异特征

叶片和细根是植物对环境变化响应的主要功能器官,在气候变化趋势下,研究其随环境梯度的变化,对理解植物对环境的适应策略具有重要意义。本文是在色季拉山急尖长苞冷杉分布区,按海拔梯度(3800、3900、4000、4100、4200、4300、4400m)选择7个典型样地,在样地内对其叶片基本特征、叶绿素含量、比叶重和细根特征(0—60cm)等参数进行了测量。结果表明:(1)随海拔梯度升高,急尖长苞冷杉叶片叶面积减小、叶片厚度增加、叶绿素含量逐渐降低、比叶重显著增大。3900m处急尖长苞冷杉的叶片形态特征表现最好,叶片长度、宽度和面积均显著优于其他海拔,海拔4200m叶片厚度达到最大值,叶片面积、叶绿素含量随海拔升高呈下降趋势,但在4200m处出现第二峰值。(2)随海拔梯度增加,细根各性状与海拔表现出非线性关系,其中细根生物量和细根体积在3900m和4200m处出现两次峰值,3900m处细根生物量达到最大值,4200m处细根体积达到最大值,4400m处细根生物量和细根体积均显著少于其他海拔。细根根长密度在海拔3900、4200、4300m较高,三者之间差异不显著,但显著高于其余海拔,4400m海拔细根根长密度最小。细根表面积在3900m海拔处达到最大值,显著高于其他海拔,4200和4300m次之,3800、4000和4400m海拔下细根表面积相对较小。4400m处细根比根长达到最大值。各海拔上细根均主要分布在0—20cm土层。随土层厚度增加,各海拔细根生物量和根系体积在0—60cm土层范围内均逐渐减小;细根根长密度、表面积在20—40cm和40—60cm显著提高;同一海拔细根比根长随土层深度增加呈明显的增加趋势。各海拔40—60cm土层细根比根长显著高于20—40和0—20cm土层。(3)综合叶片及细根特征,海拔3900m为急尖长苞冷杉的最适宜生长区域;随海拔升高,环境因子逐渐恶劣,环境胁迫加剧,急尖长苞冷杉最终形成在4400m处为其分布上限的海拔梯度格局。     

Variation and response to experimental warming in a New Zealand cushion plant species

Cold adapted plants, such as cushion plants, may be particularly sensitive to climate warming because of their compact growth form and high branch density. In the oceanic southern hemisphere, cushion communities tend to have large range distributions at low latitudes (sea level to low alpine), thus providing an opportunity to test the effects of temperature on plant morphology and reproduction across gradients. Using Donatia novae‐zelandiae as a model species, we compared the leaf morphology, reproduction and responses to warming. Two low‐alpine sites (Maungatua (880 m a.s.l.), Blue Mountains (1000 m a.s.l.)) and two sea‐level sites (Waituna 1 (0 m a.s.l.), Waituna 2 (0 m a.s.l.)) in South Island, New Zealand were used. Donatia novae‐zelandiae cushions differed significantly between the high‐elevation and sea‐level sites both morphologically and in terms of reproduction. High‐elevation cushions produced more flowers (threefold more flowers per plant) and seeds (sevenfold more seeds per capsule) than at sea level, but leaves were larger at sea level (in length and specific leaf area). The cushions were also twice as compact at the high‐elevation sites. After two growing seasons of artificial warming, seed production (35%), leaf length (7%) and width (13%), and specific leaf area (63%) significantly decreased in D. novae‐zelandiae plants; flower production was not significantly affected. Cushion plant morphology and reproduction were significantly affected by environmental drivers at their establishment sites, but all populations responded negatively to artificial warming of 1–3°C. Many cushion plants are considered keystone species because of their propensity to facilitate the growth and establishment of other plant species, the inferred negative effects of global warming on cushion plant species may have a cascading effect on other alpine plant groups.     

Water relations and growth responses of Uniola paniculata (sea oats) to soil moisture and water-table depth

Summary This study examined the water relations and growth responses of Uniola paniculata (sea oats) to (1) three watering regimes and (2) four controlled water-table depths. Uniola paniculata is frequently the dominant foredune grass along much of the southeastern Atlantic and Gulf coasts of the United States, but its distribution is limited in Louisiana. Throughout most of its range, U. paniculata tends to dominate and be well adapted to the most exposed areas of the dune where soil moisture is low. Dune elevations in Louisiana, however, rarely exceed 2 m, and as a result the depth to the water table is generally shallow. We hypothesized that if U. paniculata grows very near the water-table, as it may in Louisiana, it will display signs of water-logging stress. This study demonstrated that excessive soil moisture resulting from inundation or shallow water-table depth has a greater negative effect on plant growth than do low soil moisture conditions. Uniola paniculata's initial response to either drought or inundation was a reduction of leaf (stomatal) conductance and a concomitant decrease in leaf elongation. However, plants could recover from drought-induced leaf xylem pressures of less than-3.3 MPa, but prolonged inundation killed the plants. Waterlogging stress (manifested in significantly reduced leaf stomatal conductances and reduced biomass production) was observed in plants grown at 0.3 m above the water table. This stress was relieved, however, at an elevation of 0.9 m above the water table. As the elevation was increased from 0.9 to 2.7 m, there were no signs of drought stress nor a stimulation in growth due to lower soil moisture. We concluded that although U. paniculata's moisture-conserving traits adapt it well to the dune environment, this species can grow very well at an elevation of only 0.9 m above the water table. Field measurements of water-table depth in three Louisiana populations averaged about 1.3 m. Therefore, the observed limited distribution of U. paniculata along the Louisiana coast apparently cannot be explained by water-logging stress induced by the low dune elevations and the corresponding shallow water-table depth.     

Some effects of enhanced UV-B irradiation on the growth and composition of plants

Barley (Hordeum vulgare), corn (Zea mays), bean (Phaseolus vulgaris), and radish (Raphanus sativus) seedlings were continuously irradiated under a lighting device for 5–10 d at an increased ultraviolet (UV)-B fluence rate. In their growth parameters, composition, and leaf surface, these four species responded differently to the increased UV-B exposure. Bean seedlings suffered the most serious effects, radish and barley less, and corn was hardly influenced at all. In all plant species, the fresh weight, the leaf area, the amounts of chlorophylls, carotenoids and the galactolipids of the chloroplasts were reduced. The lipid content of the corn and bean seedlings also diminished. But all the irradiated plants showed a rise in their protein content compared to the control plants. The content of flavonoids increased in barley and radish seedlings by about 50%. The effects on growth parameters and composition were more extensive with increasing UV-B fluence rates, at least as shown in the case of barley seedlings. The fresh weights fell proportionally with the chlorophylls and carotenoids. In contrast, the flavonoid content of barley leaves rose parallel to the increasing UV-B fluence rates and reached 180% of the value in the control plants with the highest UV-B fluence rate. Scorching appeared regularly in the form of bronze leaf discoloration at the highest UV-B fluence rates. Scanning electron micrographs of the leaf surface of UV-B irradiated plants showed deformed epidermal structures.Abbreviations MGDG monogalactosyldiglyceride - DGDG digalactosyldiglyceride - SL sulfoquinovosyldiglyceride - PG phosphatidylglycerol - PC phosphatidylcholine - PE phosphatidylethanolamine - PI phosphatidylinositol - LA leaf are - FW fresh weight - DW dry weight - SEM scanning electron microscopy - C total carotenoids - Chl total chlorophyll     

Environmental and genetic control of insect abundance and herbivory along a forest elevational gradient

Environmental conditions and plant genotype may influence insect herbivory along elevational gradients. Plant damage would decrease with elevation as temperature declines to suboptimal levels for insects. However, host plants at higher elevations may exhibit traits that either reduce or enhance leaf quality to insects, with uncertain net effects on herbivory. We examined folivory, insect abundance and leaf traits along six replicated elevational ranges in Nothofagus pumilio forests of the northern Patagonian Andes, Argentina. We also conducted a reciprocal transplant experiment between low- and high-elevation sites to test the extent of environmental and plant genetic control on insect abundance and folivory. We found that insect abundance, leaf size and specific leaf area decreased, whereas foliar phosphorous content increased, from low-, through mid- to high-elevation sites. Path analysis indicated that changes in both insect abundance and leaf traits were important in reducing folivory with increasing elevation and decreasing mean temperature. At both planting sites, plants from a low-elevation origin experienced higher damage and supported greater insect loads than plants from a high-elevation origin. The differences in leaf damage between sites were twofold larger than those between plant origins, suggesting that local environment was more important than host genotype in explaining folivory patterns. Different folivore guilds exhibited qualitatively similar responses to elevation. Our results suggest an increase in insect folivory on high-elevation N. pumilio forests under future climate warming scenarios. However, in the short-term, folivory increases might be smaller than expected from insect abundance only because at high elevations herbivores would encounter more resistant tree genotypes.     

An invasive frog, Eleutherodactylus coqui, increases new leaf production and leaf litter decomposition rates through nutrient cycling in Hawaii

A frog endemic to Puerto Rico, Eleutherodactylus coqui, invaded Hawaii in the late 1980s, where it can reach densities of 50,000 individuals ha⁻¹. Effects of this introduced insectivore on invertebrate communities and ecosystem processes, such as nutrient cycling, are largely unknown. In two study sites on the Island of Hawaii, we studied the top-down effects of E. coqui on aerial, herbivorous, and leaf litter invertebrates; herbivory, plant growth, and leaf litter decomposition rates; and leaf litter and throughfall chemistry over 6 months. We found that E. coqui reduced all invertebrate communities at one of the two study sites. Across sites, E. coqui lowered herbivory rates, increased NH₄⁺ and P concentrations in throughfall, increased Mg, N, P, and K in decomposing leaf litter, increased new leaf production of Psidium cattleianum, and increased leaf litter decomposition rates of Metrosideros polymorpha. In summary, E. coqui effects on invertebrates differed by site, but E. coqui effects on ecosystem processes were similar across sites. Path analyses suggest that E. coqui increased the number of new P. cattleianum leaves and leaf litter decomposition rates of M. polymorpha by making nutrients more available to plants and microbes rather than through changes in the invertebrate community. Results suggest that E. coqui in Hawaii has the potential to reduce endemic invertebrates and increase nutrient cycling rates, which may confer a competitive advantage to invasive plants in an ecosystem where native species have evolved in nutrient-poor conditions.     

Plant transpiration at high elevations: Theory,field measurements,and comparisons with desert plants

Summary The influence of elevational changes on plant transpiration was evaluated using leaf energy balance equations and well-known elevational changes in the physical parameters that influence water vapor diffusion. Simulated transpirational fluxes for large leaves with low and high stomatal resistances to water vapor diffusion were compared to small leaves with identical stomatal resistances at elevations ranging from sea level to 4 km. The specific influence of various air temperature lapse rates was also tested. Validation of the simulated results was accomplished by comparing actual field measurements taken at a low elevation (300 m) desert site with similar measurements for a high elevation (2,560 m) mountain research site. Close agreement was observed between predicted and measured values of transpiration for the environmental and leaf parameters tested.Substantial increases in solar irradiation and the diffusion coefficient for water vapor in air (D _wv) occurred with increasing elevation, while air and leaf temperatures, the water vapor concentration difference between the leaf and air, longwave irradiation, and the thermal conductivity coefficient for heat in air decreased with increasing elevation. These changes resulted in temperatures for sunlit leaves that were further above air temperature at higher elevations, especially for large leaves. For large leaves with low stomatal resistances, transpirational fluxes for low-elevation desert plants were close to those predicted for high-elevation plants even though the sunlit leaf temperatures of these mountain plants were over 10°C cooler. Simulating conditions with a low air temperature lapse rate (0.003° C m^-1 and 0.004° C m^-1) resulted in predicted transpirational fluxes that were greater than those calculated for the desert site. Transpiration for smaller leaves decreased with elevation for all lapse rates tested (0.003° C m^-1 to 0.010° C m^-1). However, transpirational fluxes at higher elevations were considerably greater than expected for all leaves, especially larger leaves, due to the strong influence of increased solar heating and a greater D _wv. These results are discussed in terms of similarities in leaf structure and plant habit observed among low-elevation desert plants and high-elevation alpine and subalpine plants.     

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..     

Intraspecific variation in the response of Themeda triandra to defoliation: the effect of time of recovery and growth rates on compensatory growth

Summary The response to a single defoliation was studied on three clones of Themeda triandra collected in the short, mid, and tall grassland regions of the Serengeti National Park (Tanzania). These sites represent a gradient of decreasing grazing intensity. Growth, allocation pattern, and several morphometric traits were monitored during an 80-day period. Clipped plants of the short and medium clones fully compensated for the reduction of biomass, while plants of the tall clone showed overcompensation. During the first two weeks after clipping, clipped plants showed lower relative growth rates than unclipped ones, whereas the opposite was observed later on. Clipped plants compensated for the removal of leaf area by producing new leaves with lower specific weights and higher nitrogen content. They also produced more, smaller tillers. Although clipped plants mobilized nonstructural carbohydrates from roots and crowns, this did not account for a significant amount of growth. Relative growth rates of unclipped plants of the short clone were higher. The relative growth rate of the short clone diminished less after clipping, but also exhibited the lowest increase later. The tall clone was the most negatively affected early, but showed the highest compensation later. Compared to the other clones, the short ecotype showed many of the characteristics that defoliation induced in each individual of any clone: higher allocation to leaf area production, higher relative growth rate, higher number but smaller size of tillers, and lower leaf specific weights.     

Transitory growth control of apple seedlings with less persistent triazole derivatives

Paclobutrazol, triapenthenol (RSW0411), and BAS111 were applied to 4-week-old Delicious apple seedlings (Malus domestica Borkh.) as a root drench at 0.1, 1.0, and 10.0 mg per plant. Paclobutrazol eliminated shoot extension growth for 8 weeks at all three rates. RSW0411 controlled shoot elongation only at the highest rate. BAS111 produced the widest response, with shoot growth ranging from 38% to 93% of controls for the highest and lowest rates, respectively. Generally, leaf area decreased and leaf density increased with increasing rates of all chemicals. Root weight of plants treated with paclobutrazol nearly doubled but increased only slightly with RSW0411 and BAS111. Chemical analysis of the leaf tissue 8 weeks after treatment showed paclobutrazol levels highest, followed by RSW0411 and BAS111. Mention of a trademark, proprietary product, or vendor does not constitute a guarantee or warranty of the product by the U.S. Department of Agriculture and does not imply its approval to the exclusion of other products that may also be suitable.     

An experimental assessment of the upper elevational limit of flowering plants in the western Himalayas

Transplant experiments are a useful tool both for determining species’ ranges and understanding their cause, yet such experiments have seldom been performed in areas where plants reach extremely high elevations. We examined the position of the upper elevational limits of vascular plants in E Ladakh, India, by transplanting individuals of 14 subnival species from 5800–5850 m elevation to a control site at the same elevation, as well as to edaphically suitable sites at 5960 m (subnival belt with sparse plant cover), to 6030 m (the highest elevation reached by vascular plants in the area) and to 6160 m elevation (no vascular plants observed). Two years later, transplants of 13 species survived at the control elevation, whereas 5 species survived at 5960 m, 2 species (Waldheimia tridactylites, Poa attenuata) at 6030 m, and none at 6160 m. The highest elevation at which transplanted flowering plants survived corresponds well to the observed elevational limit. Soil temperature data at the sites suggest that the growing season, defined as the period with mean daily soil temperature above zero, lasted nearly 3 months at 5960 m, ca 1.5 month at 6030 m, but <3 weeks at 6160 m, moreover interrupted at the highest elevation by several days with temperatures remaining below zero. The experiment confirmed the observed sharp limit of vegetation, set not by any physical barrier per se (e.g. not by the top of the massif), but instead by physiological constraints of the species. The result provides support for the assumptions of mid‐domain effect models that domain limits are defined by shared organismal adaptations in relation to environmental gradients, in this case tolerance to freezing temperature.     

Temperature,water, and PAR influences on predicted and measured productivity of Agave deserti at various elevations

Summary To measure productivity of Agave deserti over its elevational range in the northwestern Sonoran Desert, leaf unfolding from its basal rosette was monitored on groups of 10 plants at 13 sites. Based on data from an intermediate elevation (840 m), leaf unfolding proved to be highly correlated (r ²=0.88) with an environmental productivity index (EPI) formed as the product of indices for water status, temperature, and photosynthetically active radiation (PAR); each of these latter indices indicated the fraction of maximum net CO₂ uptake expected for that parameter based on laboratory measurements of gas exchange and field microclimatic data. At 840 m, the main environmental variable influencing leaf unfolding for A. deserti over a 2-y period was soil water potential. On steep slopes, however, leaf unfolding during the winter ranged from 0.7 leaves per 10 plants for north-facing slopes to 7.3 for south-facing slopes, reflecting the importance of PAR. Summer and winter rainfall increased 3-fold from elevations of 300 m to 1,200 m. Temperatures were more optimal for net CO₂ uptake at high elevations in the summer and at low elevations in the winter. Hence EPI increased with elevation in the summer but was maximal at an intermediate elevation in the winter. Moreover, measured leaf unfolding in both the summer and the winter closely followed the changes in EPI with elevation, indicating that productivity could be closely predicted for A. deserti based on physiological CO₂ responses and changes in environmental conditions with elevation.     

三峡水库长期水淹条件下耐淹植物甜根子草的资源分配特征

为研究历经三峡水库长期水淹的驯化后,不同海拔高程之间相同种源的甜根子草的生物量分配特征是否发生改变,进而探讨该物种对水淹胁迫表现出的适应性进化特征,试验于2008年初选育相同种源的甜根子草同龄幼苗栽植于三峡水库消落区甜根子草种植试验示范区,并考察了2012年、2013年不同海拔高程甜根子草植株的形态和生物量特征。试验共设置3个海拔高程,即水淹高程168、172m和不受水淹对照高程176m。试验结果表明:(1)较低高程的甜根子草植株较矮小细弱,168m高程的甜根子草植株主茎长和主茎基径显著低于对照176m高程(P0.05);平均节间长度随高程的降低而缩短;与之相反,主茎长/主茎基径随高程的降低而增大。(2)甜根子草的叶片厚度、叶片长/叶片宽、叶片长/叶鞘长均随海拔高程的降低而减小;与之相反,比叶面积随高程的降低而增大。(3)水淹前,甜根子草近端成熟节间的质量密度随高程的降低而增大;水淹后,其地上存活茎段基部成熟节间的质量密度在各高程之间无显著差异(P0.05)。以上研究结果表明,甜根子草历经三峡水库长期水淹的驯化后,生物量分配特征在不同海拔高程之间发生了改变,表现出了相应的驯化特征。相较于高高程的甜根子草植株而言,低高程的植株生长缓慢,采取低株高下的高向生物量投资策略;对叶的物质投资大部分分配到叶面积的增加、叶鞘的伸长生长和叶片的直立生长上,以加强植株的光合生产。     

Leaf growth, stomatal diffusion resistances and photosynthesis during droughting of Lolium perenne populations selected for contrasting stomatal length and frequency

Lolium perenne selection lines with high calculated stomatal resistances to diffusion (r_s) as a result of either few or short stomata, maintained leaf extension rates and photosynthetic rates longer than selection lines with low resistances when deprived of water. There were no significant differences between high and low r_s plants in light saturated CO₂ uptake of turgid attached leaves. When grown in soil drying to 21% moisture, plants with low calculated r_s exhibited lower minimum leaf resistances (r_l) than those with high, measured with a diffusion porometer, on all except the last day. The daily maximum r_l (1.5 h after the start of the light period) became greater among low than high r_s plants as the difference in rate of leaf extension between the two groups of plants increased with drying soil. Rate of leaf extension was negatively correlated with daily maximum r_l and started to decline when relative leaf water content (RLWC), at 5 h after the start of the light period, fell below about 88%. Transpiration rate of plants grown in different soil moisture regimes was correlated (r=+ 0.83, P < 0.01) with mean maximum adaxial leaf conductance (reciprocal of resistance). There was a highly significant correlation (r=+ 0.62, P < 0.01) between calculated adaxial r_s and mean minimum measured r_l among plants growing at high or intermediate soil moisture, but not at low. Therefore, some random variation in minimum r_l, even with adequate moisture, seemed to be unaccounted for by variation in stomatal numbers or size. Selection for increased numbers of adaxial stomata also resulted in more on the abaxial surface, but mean adaxial/abaxial ratio in the ‘frequent’ stomata plants was still only about 9:1.     

Effects of leaf age,nitrogen nutrition and photon flux density on the distribution of nitrogen among leaves of a vine (Ipomoea tricolor Cav.) grown horizontally to avoid mutual shading of leaves

Effects of leaf age, nitrogen nutrition and photon flux density (PFD) on the distribution of nitrogen among leaves were investigated in a vine, Ipomoea tricolor Cav., which had been grown horizontally so as to avoid mutual shading of leaves. The nitrogen content was highest in newly developed young leaves and decreased with age of leaves in plants grown at low nitrate concentrations and with all leaves exposed to full sunlight. Thus, a distinct gradient of leaf nitrogen content was formed along the gradient of leaf age. However, no gradient of leaf nitrogen content was formed in plants grown at a high nitrate concentration. Effects of PFD on the distribution of nitrogen were examined by shading leaves in a manner that simulated changes in the light gradient of an erect herbaceous canopy (i.e., where old leaves were placed under increasingly darker conditions with growth of the canopy). This canopy-type shading steepened the gradient of leaf nitrogen content in plants grown at a low nitrogen supply, and created a gradient in plants grown at high concentrations of nitrate. The steeper the gradient of PFD, the larger the gradient of leaf nitrogen that was formed. When the gradient of shading was inverted, that is, younger leaves were subjected to increasingly heavier shade, while keeping the oldest leaves exposed to full sunlight, an inverted gradient of leaf nitrogen content was formed at high nitrate concentrations. The gradient of leaf nitrogen content generated either by advance of leaf age at low nitrogen availability, or by canopy-type shading, was comparable to those reported for the canopies of erect herbaceous plants. It is concluded that both leaf age and PFD have potential to cause the non-uniform distribution of leaf nitrogen. It is also shown that the contribution of leaf age increases with the decrease in nitrogen nutrition level.     

Whorl and pollen-shed stage application of Beauveria bassiana for suppression of adult western corn rootworm

The corn rootworm complex is the most damaging insect pest of corn (Zea mays L.). This study was conducted to determine the efficacy of whorl and pollen-shed stage applications of a granular formulation of Beauveria bassiana (Balsamo) Vuillemin for control of adult western corn rootworm (Diabrotica virgifera virgifera Le Conte). The effect of application time (whorl-stage, pollen-shed) and plant surface exposed (leaves and leaf collars; silks; leaves, leaf collars, and silks) on level of beetle fungal infection were investigated. In addition, the number of colony forming units of B. bassiana in the corn leaf collar area was quantified. In the three years (1998–2000) of the study, application of B. bassiana at whorl-stage did not significantly increase beetle fungal infection. Beauveria bassiana applied to plants at pollen-shed in 1998 resulted in a significant increase in beetle infection with 51% of beetles from treated plants infected and 6.0% from control plants. Similar applications at pollen-shed in 1999 and 2000 resulted in very low infection levels. Beauveria bassiana application at pollen-shed stage significantly increased the number of colony forming units per leaf collar during all years of the study. Beetle infection with B. bassiana did not differ consistently among plant surface to which beetles were exposed for either application. Increased fungal load in leaf collars was not correlated with increased levels of adult infection. Increased rates of B. bassiana and application when beetles are present on the plants are likely needed to significantly increase infection rates.     

Growth of Trifolium alpinum: Effects of soil properties, symbionts and pathogens

The lowland cultivation of Trifolium alpinum, a clover species found on acid soils in the Alps and suitable for the restoration of erosion areas at high altitudes, failed repeatedly in previous experiments. Three experiments were carried out in a controlled environment to elucidate the reasons for the failure and to develop possible cultivation strategies. In experiment I, T. alpinum was grown in an autochthonous soil from the Alps (high elevation) and in two allochthonous soils, a grassland soil from the Hercynian mountains (medium elevation), and an arable soil (low elevation), in which the seed propagation of T. alpinum had failed previously. The two allochthonous soils had lower contents of soil organic C and ergosterol, an indicator for fungal biomass, than the autochthonous high-elevation soil, but higher levels of exchangeable Ca and extractable P. Plants grown in the allochthonous soils achieved higher biomass and total N amounts per plant than those from the high elevation soil if inoculated with this autochthonous material to establish rhizobial infection. In the allochthonous high elevation soil, the growth of T. alpinum was P-limited as shown in experiment II. In experiment I, plants grown in the low elevation soil had a lower biomass and smaller number of active leaves at 120 days after emergence than those grown on the medium elevation soil. This difference can be explained by strong colonization with the phytophagous nematode Pratylenchus sp., as demonstrated in experiment III by comparing plant growth either in untreated or in autoclaved low-elevation soil. Successful propagation of T. alpinum at low elevation may be achieved through suitable inoculation with autochthonous soil biota, especially Rhizobia, and avoidance of soils infested by Pratylenchus species by choosing sites with acidic soil and ensuring adequate P-availability.     

Fatty acid synthesis by slices from developing leaves

Fatty acid synthesis was studied in successive leaf sections from the base to the tip of developing barley (Hordeum vulgare L.), maize (Zea mays L.), rye grass (Lolium perenne L.) and wheat (Triticum aestivium L.) leaves. The basal regions of the leaves had the lowest rates of fatty acid synthesis and accumulated small amounts of very long chain fatty acids. Fatty acid synthesis was highest in the middle leaf sections in all four plants. Linolenic acid synthesis from [1-¹⁴C]acetate was highest in the distal leaf sections of rye grass. The labelling of the fatty acids of individual lipids of rye grass was examined and it was found that [¹⁴C]linolenic acid was highest in the galactolipids. Synthesis of this acid in the galactolipids was most active in leaf segment C. Only traces of [¹⁴C]linolenic acid were ever found in phosphatidylcholine and it is concluded that this phospholipid cannot serve as a substrate for linoleic acid desaturation in rye grass. The synthesis of fatty acids was sensitive to arsenite, fluoride and the herbicide EPTC. The latter was only inhibitory towards those leaf segments which made very long chain fatty acids. Formation of fatty acids from [1-¹⁴C]acetate was also studied in chloroplasts prepared from successive leaf sections of rye grass. Chloroplasts isolated from the middle leaf sections had the highest activity. Palmitic and oleic acids were the main fatty acid products in all chloroplast preparations. Linolenic acid synthesis was highest in chlorplasts isolated from the distal leaf sections of rye grass.