Water-use efficiency in cork oak (Quercus suber) is modified by the interaction of water and light availabilities

We studied the interaction of light and water on water-use efficiency in cork oak (Quercus suber L.) seedlings. One-year-old cork oak seedlings were grown in pots in a factorial experiment with four light treatments (68, 50, 15 and 5% of full sunlight) and two irrigation regimes: well watered (WW) and moderate drought stress (WS). Leaf predawn water potential, which was measured at the end of each of two cycles, did not differ among the light treatments. Water-use efficiency, assessed by carbon isotope composition (delta(13)C), tended to increase with increasing irradiance. The trend was similar in the WW and WS treatments, though with lower delta(13)C in all light treatments in the WW irrigation regime. Specific leaf area increased with decreasing irradiance, and was inversely correlated with delta(13)C. Thus, changes in delta(13)C could be explained in part by light-induced modifications in leaf morphology. The relationship between stomatal conductance to water vapor and net photosynthesis on a leaf area basis confirmed that seedlings in higher irradiances maintained a higher rate of carbon uptake at a particular stomatal conductance, implying that shaded seedlings have a lower water-use efficiency that is unrelated to water availability.     

Growth,biomass partitioning,and water-use efficiency of a leguminous shrub (<Emphasis Type="Italic">Bauhinia faberi</Emphasis> var. <Emphasis Type="Italic">microphylla</Emphasis>) in response to various water availabilities

Responses of the endemic leguminous shrub Bauhinia faberi var. microphylla, to various soil water supply regimes were studied in order to assess water stress tolerance of seedlings. Two-month-old seedlings were grown under water supply regimes of 100, 80, 60, 40, and 20% water field capacity (FC), respectively, in a temperature and light-controlled greenhouse. Plant height and leaf number were measured monthly over a 4-month period, while water use (WU), water-use efficiency (WUE), leaf relative water content (RWC), biomass production and its partitioning were recorded at the end of the experiment. Seedlings exhibited the greatest biomass production, height, basal diameter, branch number, leaf number, and leaf area when soil content was at 100% FC, and slightly declined at 80% FC. These parameters declined significantly under 60% FC water supply, and severely reduced under 40 and 20% FC. RWC, WU and WUE decreased, while the ratio of root mass to stem mass (R:S) increased in response to decreasing water supply. Water stress caused leaf shedding, but not plant death. The results demonstrated that B. faberi var. microphylla seedlings could tolerate drought by reducing branching and leaf area while maintaining a high R:S ratio. However, low dry mass and WUE at 40 and 20% FC suggested that the seedlings did not produce significant biomass under prolonged severe water deficit. Therefore, before introducing B. faberi var. microphylla in vegetation restoration efforts, water supply above 40% FC is recommended for seedlings to maintain growth.     

Effects of root trenching of overstorey Norway spruce (Piceaabies) on growth and biomass of underplanted beech (Fagussylvatica) and Douglas fir (Pseudotsugamenziesii) saplings

In Central Europe, the conversion of pure Norway spruce stands (Picea abies [L.] Karst.) into mixed stands with beech (Fagus silvatica L.) and other species like e.g. Douglas fir (Pseudotsuga menziesii [Mirb.] Franco) is accomplished mainly by underplanting of seedlings beneath the canopy of overstorey spruce trees after partial cutting treatments what means exposure to shade and below-ground root competition by the overstorey to the seedlings. Particularly about the second factor, our knowledge is limited. Therefore, we carried out a below-ground competition exclusion experiment by root trenching and investigated the effects on soil resources, growth, and biomass partitioning of underplanted beech and Douglas fir saplings under target diameter and strip cutting treatments. The exclusion of overstorey root competition by trenching increased the soil water potential in the second year that had a fairly dry growing season and led to significantly higher foliar concentrations of most nutrients, particularly in Douglas fir, indicating an amended nutrient supply. Both improvements were accompanied by an increase in length and diameter increment of the underplanted saplings, appearing in both species only after having surpassed a species-specific threshold light value (Douglas fir 16% of above canopy radiation, beech 22%). We also found significant interactions between trenching and light for specific fine root length and further biomass and morphological parameters. Judged by the much steeper increase in height and diameter growth with increasing light after release from below-ground competition, Douglas fir saplings appeared to be more sensitive to root competition than beech saplings what conforms to older findings for beech. According to our results, a strip cutting seems to be more appropriate than a target diameter cutting treatment to replace a pure spruce stand by a mixed stand with beech and Douglas fir.     

Effects of shading and removal of plant parts on growth of Trema micrantha seedlings

Effects of artificial shading and removal of plant parts on growth of Trema micrantha (L.) Blume (Ulmaceae) seedlings were studied. Seedlings were grown in pots in a greenhouse in 45, 30, 10.6, 4.8 and 1.8% of full sunlight. Shading for 60 days had no effect on survival, but it influenced all growth parameters measured. Total biomass decreased with decreasing irradiance, reflecting reductions in dry mass of leaves, stems and roots. In response to shading, allocation of biomass to leaves increased, while allocation of biomass to roots decreased. Specific leaf area, leaf area ratio and leaf mass ratio increased with decreasing irradiance. Decreases in relative growth rate were caused by reductions in net assimilation rate rather than leaf area ratio. Photosynthetic efficiency, as determined by the Fv/Fm ratio (Fv = variable fluorescence, Fm = maximal fluorescence), was unaffected by the shading treatments. Partial removal of leaves, stem or roots did not affect seedling survival. Seedlings responded to removal of plant parts by compensatory growth. Topophysis was observed when the apex was removed: the lateral buds developed only as new plagiotropic lateral shoots; consequently, the decapitated plant ceased height growth and was unable to compete with its neighbors for light.     

Morphological response of Vitex negundo var. heterophylla and Ziziphus jujuba var. spinosa to the combined impact of drought and shade

Light and soil moisture availability are two important abiotic factors influencing plant growth in an agroforestry system. Different soil moisture and light treatments were applied to examine the combined impact of drought and shade on the morphological plasticity of Vitex negundo var. heterophylla (Chinese chastetree) and Ziziphus jujuba var. spinosa (Spine jujube). We found that the interaction of light and soil moisture was orthogonal in the two species. V. negundo captured irradiance efficiently with relatively long petiole and petiolule, while Z. jujuba maintained higher branches to absorb light than V. negundo. Compared to the seedlings under full sunlight, the palmately compound leaves of V. negundo under low light showed larger specific leaf area (SLA), lower ratio of leaflet length to width and higher leaf mass ratio (leaf biomass to total biomass); in contrast, the simple leaves of Z. jujuba under low light showed larger SLA and ratio of leaf length to petiole length. In both species, drought reduced the branch number, mean internode length of stem, and increased root mass ratio (root biomass to total biomass), but leaf morphology showed little variation to the decreased soil moisture. Between the two species, V. negundo is more shade-tolerant, while Z. jujuba is more drought-tolerant. In an alley cropping system, the spacing can be smaller in the V. negundo-crop system than the Z. jujuba-crop system when soil moisture is adequate. However, the V. negundo-crop system should be managed with more caution under the soil moisture limited conditions.     

Seasonal changes in photosynthesis and growth of Zizyphus attopensis seedlings in three contrasting microhabitats in a tropical seasonal rain forest

We hypothesized that photosynthesis and growth of tropical vegetation at its most northern distribution in Asia (Xishuangbanna, SW China) is adversely affected by seasonal drought and chilling temperatures. To test this hypothesis, we measured photosynthetic and growth characteristics of Zizyphus attopensis Pierre seedlings grown in three contrasting forest microhabitats: the understory, a small gap and a large gap. Photosynthetic capacity (light-saturated photosynthetic rate (A(max)), maximum rate of carboxylation and electron transport rate) and partitioning of leaf nitrogen (N) into carboxylation and electron transport differed significantly among seasons and microhabitats. Specific leaf area (SLA) did not change seasonally, but differed significantly among microhabitats and showed a negative linear relationship with daily integrated photon flux (PPF(i)). In contrast, leaf N concentration per unit area (N(a)) changed seasonally but did not differ among microhabitats. Measurements of maximum PSII photochemical efficiency (F(v)/F(m)) indicated that chronic photoinhibition did not occur in seedlings in any of the microhabitats during the study. Photosynthetic capacity was greatest in the wet season and lowest in the cool season. During the cool and dry seasons, the reduction in A(max) was greater in seedlings grown in the large gap than in in the understory and the small gap. Close logarithmic relationships were detected between PPF(i), leaf N(a) and photosynthetic capacity. Stem mass ratio decreased, and root mass ratio increased, in the dry season. We conclude that seasonal acclimation in growth and photosynthesis of the seedlings was associated with changes in biochemical features (particularly N(a) and partitioning of total leaf N between the different photosynthetic pools) and biomass allocation, rather than with changes in leaf morphological features (such as SLA). Local irradiance is the main factor driving seasonal variations in growth and photosynthesis in the study area, where the presence of heavy fog during the cool and dry seasons limits irradiance, but supplies water to the soil surface layers.     

Effects of drought and rewatering on growth and transpiration in European beech seedlings late in the growing season

Drought stress is one of the most important environmental factors affecting plant growth and survival. To date, most studies aim at understanding of post-stress physiological and anatomical adaptation to drought stress; however only few studies focus on plant recovery. In the present study, transpiration, shoot water potential, and anatomical and morphological measurements were performed on 4-year-old European beech seedlings with fully developed leaves. The seedlings were exposed to three levels of soil water potential (well-watered, moderate drought stress and severe drought stress) and followed by rewatering under greenhouse conditions. Reduced transpiration rates were observed in the stressed seedlings as a response to drought stress, whereas anatomical and morphological variables remained unchanged. Three days after rewatering, transpiration rates in both moderately and severely stressed seedlings recovered to the levels of those of well-watered seedlings. Drought stress promoted leaf budding, resulting in higher shoot dry mass of stressed seedlings. Our findings indicate that anatomical and morphological adaptations of European beech seedlings to drought stress are visibly limited during late-season growth stages. These results will help us to further understand factors involved in drought adaptation potential of European beech seedlings faced with expected climate-related environmental changes. To complete our findings, further experiments on plant recovery from drought stress should be focused on different periods of growing season.     

Genotypic variation and phenotypic plasticity in the drought response of fine roots of European beech

How temperate trees respond to drier summers, as predicted by climate change models for parts of Europe and eastern North America, will depend on the drought susceptibility of the root systems. We investigated the importance of the genetic constitution for the belowground drought response of European beech (Fagus sylvatica L.), in four populations from regions differing in precipitation (520-970 mm year(-1)). Saplings were grown at ample (10 vol.%; well-watered) or reduced (5 vol.%; drought treatment) soil water content in the G?ttingen Rhizolab Facility for two consecutive summers, and the responses of fine root biomass, root morphology, root depth distribution, and fine root production and turnover were investigated by a combined mini-rhizotron and harvest technique approach. In the drought treatment, total root mass per plant was reduced by 30-40% as a result of: (1) a reduction in median fine root lifespan by roughly 50% and hence an increase in fine root turnover; and (2) a 10-fold reduction in relative fine root growth rate (productivity per standing root biomass). The root:shoot ratio did not increase with drought. Although beech plants originating from drier climates tended to reduce their root biomass in response to drought less than those from wetter climates, analyses of variance revealed no significant influence of genotype on root mass, morphology, growth rate or turnover. However, most fine root traits showed marked differences between the well-watered and drought treatments. We conclude that beech saplings respond to summer drought primarily by shortening root lifespan, whereas root system structure and root:shoot carbon partitioning pattern are unaltered. Beech fine root growth and turnover exhibited high phenotypic plasticity, but genotypic variation was of minor importance. In contrast, genotype had a strong influence on leaf and shoot morphogenesis and growth.     

Effects of water stress on growth,dry matter allocation and water-use efficiency of a leguminous species, <Emphasis Type="Italic">Sophora davidii</Emphasis>

The adaptation responses to different water conditions and the drought tolerance of Sophora davidii seedlings were assessed in a greenhouse experiment. Two-month-old seedlings were subjected to the following water supplies for 95 days: 100, 80, 60, 40 and 20% of field water capacity. The seedlings at 100% FC had the greatest productivity, height, basal diameter, branch number, leaf number and leaf area. Water supply <80% FC was the threshold of drought-initiated negative effects on seedling growth, yield and physiological processes; these parameters were severely reduced at 20% FC, however, there was no plant death during the experiment. Moreover, water stress decreased leaf relative water content, specific leaf area, leaf area ratio, and water-use efficiency (WUE), whereas it increased the biomass allocation to roots, which resulted in a higher root:stem mass ratio under drought. The S. davidii seedlings tolerated drought by maintaining high leaf relative water content and by reducing branching and leaf expansion. However, low productivity and WUE at 20% FC suggested that seedlings did not produce high biomass under severe drought. Therefore, prior to introducing S. davidii in forestation efforts, a water supply >40% FC is recommended for seedlings to maintain growth and productivity. These results provide insights into limitations and opportunities for establishment of S. davidii in arid regions.     

Response of Fagus sylvatica Seedlings to Root Trenching of Overstorey Picea abies

In a trenching experiment the impact of root competition by overstorey Norway spruce on growth and allocation of above-ground woody biomass of advanced planted beech was investigated in two stands over two growing seasons after trenching. Measurements of soil water potential were taken inside and outside trenched areas by tensiometers at 16 measuring points in one stand from May to September 1999. Height and diameter of the seedlings, as well as branch length and diameter, were measured of beech growing in trenched and untrenched conditions in September before trenching (1998) and afterwards (1999 and 2000). Tensiometers inside the trenches reached the expiring value of -0.085 Mpa 30-40 days later, or never, compared with those in untrenched areas. Whereas relative growth rate (RGR) in height was not affected by trenching, RGR both in diameter and in estimated dry weight of the main stem were substantially enhanced in the first growing season after trenching (1999). A different result was obtained in 2000, when RGR in seedling diameter and estimated dry weight of the main stem was slightly reduced by trenching. The only variable that showed increased RGR of the trenched seedlings in both years was the estimated dry weight of the branches, indicating different above-ground biomass allocation patterns of seedlings with and without root competition.     

Growth and biomass production in Azadirachta indica seedlings in response to nutrients (N and P) and moisture stress

Production of quality seedlings is an important aspect of successful tree planting. No information is available on the effect of nutrients and water on the growth and development of the seedlings of neem (Azadirachta indica A. Juss), an important component of many tropical agroforestry systems. In an experiment in central India, the growth and nutrient-use efficiency of neem seedlings grown at various levels of light, nutrients (N and P), and water stress were determined. Seedlings were given light (diffused and complete) and nutrient (N and P) treatments comprising either high N-high P, high N-low P, low N-high P, or low N-low P. Seedlings grown in complete light (800 mol/m²/s) had four times more biomass than those grown in diffused light (200 mol/m²/s). Significant increase in seedling biomass was observed with nitrogen application, while phosphorus had no effect on biomass. Leaves contributed maximum biomass, followed by roots and stem. Nutrient use efficiency decreased with an increase in the supply of nutrients. In another experiment, containerized and bare-root seedlings were subjected to five watering treatments, viz., watering twice a week, watering weekly, watering every alternate week, watering every third week, and natural precipitation. Plant moisture stress affected both growth and survival of neem seedlings. Only 50 per cent of seedlings survived in severe drought treatment (no. 4) whereas 90 to 95 per cent seedlings showed growth in all other treatments. Plant moisture stress in severe drought treatment averaged –21 bar while in other treatments it ranged from –9 to –12 bar. Shoot-root ratio was high in bare-root seedlings as compared to containerized seedlings. Containerized seedlings had shown better endurance against drought than the bare-root seedlings. The results suggest that neem seedlings adjust their nutrient use efficiency and can be grown even under limited available resources. It is also inferred that the species can tolerate soil resource depletion caused by competitor species.This revised version was published online in November 2005 with corrections to the Cover Date.     

Spacing of silver birch seedlings grown in containers of equal size affects their morphology and its variability

Silver birch (Betula pendula Roth) seedlings were grown individually in containers arranged in rows radiating from a central point (Nelder plot) at densities spanning the range from 207 to 891 plants m(-2). Height of one set of seedlings was measured at weekly intervals and additional seedlings were harvested each week for dry mass and leaf area measurements. Height and shoot:root dry mass ratio increased with plant density. Seedling-to-seedling variability in dry mass, but not height, increased with increasing plant density. The red to far-red (R:FR) photon ratio in horizontally propagated radiation decreased with increasing density, even when plant densities and leaf area index values were low. In a separate experiment, elongating internodes of seedlings were irradiated locally by red and far-red light emitting diodes and stem elongation measured. Far-red light markedly increased stem elongation, suggesting that changes in light quality sensed by growing internodes are involved in the observed responses to growth density.     

Survival and growth of drought hardened <Emphasis Type="Italic">Eucalyptus pilularis</Emphasis> Sm. seedlings and vegetative cuttings

Forestry requires low mortality of transplanted seedlings. Mortality shortly after planting is often associated with inadequate hydration of transplants. Seedlings can be hardened to the drought conditions they may experience after transplanting by exposing them to controlled drought conditions in the nursery. Eucalyptus pilularis Sm. seedlings were drought hardened by providing nil (severe treatment) or half (mild treatment) the daily irrigation routinely received (control treatment) for up to two non-consecutive days per week during the last 4 weeks of growth in the nursery. Drought hardening reduced stem diameter, seedling leaf area, leaf area per root biomass and seedling quality measured by the Dickson quality index, but increased root:shoot ratio. Hardened seedlings had lower stomatal conductance and leaf water potential on the days they received less irrigation that the control treatment. Hardened seedlings had greater stomatal conductance and were less water stressed than seedlings experiencing drought for the first time indicating hardened seedlings had adjusted physiologically to drought. Survival after transplanting in the controlled drought environment in a glasshouse was enhanced by the hardening treatments. Non hardened seedlings that had had their upper leaves manually removed immediately prior to transplanting to reduce leaf area (top-clipped) had similar survival to hardened seedlings. Stomatal conductance and leaf water potential after transplanting were higher in hardened and top-clipped seedlings than unhardened control seedlings or vegetative cuttings. Survival in the field trial was over 95% for all treatments, possibly as rain fell within 4 days of planting and follow-up rain occurred in the subsequent weeks. Neither the hardened or top-clipped seedlings planted in the field trial had reduced growth, increased propensity to form double leaders or worse stem form than control seedlings when measured at age 3 years.     

Responses to ultraviolet-B radiation by purely symbiotic and NO3-fed nodulated tree and shrub legumes indigenous to southern Africa

Purely symbiotic and NO3-fed nodulated seedlings of Virgilia oroboides (Bergius) T.M. Salter, Cyclopia maculata (L.) Vent and Podalyria calyptrata Willd. were exposed to biologically effective ultraviolet-B radiation (UV-B) to assess the effects of above- and below-ambient UV-B on growth, symbiotic function and metabolite concentrations. Seedlings were grown outdoors either on tables under ambient or 34 or 66% above-ambient UV-B conditions (UV-B100 control, UV-B134 and UV-B166, respectively), or in chambers providing below-ambient (22% of ambient) UV-B (UV-B22) along with a UV-A control and a photosynthetically active radiation (PAR) control. Exposure of seedlings to UV-B166 radiation reduced (P < or = 0.05) leaf and stem dry mass by 34 and 39%, respectively, in C. maculata, and reduced leaf nitrogen concentration (%N) by 12% in V. oroboides. Nodule %N in C. maculata and stem %N in P. calyptrata also decreased (P < or = 0.05) in response to UV-B22 radiation compared with the UV-A control, but not compared with the PAR control. Concentrations of flavonoids, soluble sugars and starch were unaltered by the UV-B treatments. Application of 1 mM NO3 to UV-B166-treated seedlings increased whole-plant dry mass of V. oroboides and P. calyptrata by 47 and 52%, respectively. Dry mass of organs, nodule %N and total N concentration of these species also increased with NO3 application. However, NO3 supply decreased (P < or = 0.05) nodule dry mass, stem %N and leaf %N as well as root and leaf anthocyanin concentrations in C. maculata. In terms of UV-B x N interactions, dry mass of stems, roots, nodules and total biomass of NO3-fed C. maculata seedlings were reduced, and nodule %N, total N and leaf anthocyanins were depressed by the UV-B134 and UV-B166 treatments relative to UV-B100-treated seedlings. Although we found that above-ambient UV-B had no effects on growth and symbiotic function of V. oroboides and P. calyptrata seedlings, feeding NO3 to these species increased (P < or = 0.05) seedling growth. In contrast, purely symbiotic C. maculata seedlings were sensitive to the UV-B166 radiation treatment, and adding NO3 further increased their sensitivity to both the UV-B134 and UV-B166 treatments.     

干旱胁迫对榆树幼苗生物量分配的影响

干旱是主要的环境胁迫因子之一,严重影响植物的分布与生长发育。通过选取辽宁阜新章古台沙地1年生榆树幼苗为材料,采用盆栽控水方法,设置田间持水量80%、60%、40%和20%等4个水分处理梯度,研究干旱胁迫对榆树幼苗株高、基径及叶片、茎和根系生物量分配的影响。结果表明:随着干旱胁迫程度增加,幼苗茎和根系的生物量逐渐降低;但轻度干旱胁迫增加了株高和基径的生长,提高了叶片生物量,以抵御干旱胁迫的影响。中度干旱胁迫下,增加了根系生物量分配的比例,从而更利于获取水分资源适应干旱环境;重度干旱胁迫下幼苗无法存活。在科尔沁沙地,栽植榆树幼苗保证存活生长的最低田间持水量为40%。     

Influence of drought stress and low irradiance on plant water relations and structural constituents in needles of Pinus ponderosa seedlings

The influence of low light on tolerance to prolonged drought was tested on unshaded and shaded seedlings of ponderosa pine (Pinus ponderosa var. scopulorum Dougl. ex Laws.). Unshaded seedlings of P. ponderosa var. ponderosa were also drought stressed to compare varietal responses to drought. The maximum irradiance received by shaded seedlings was 10% of full light. Seedlings were progressively drought stressed until predawn water potentials (Psi(x)) were -5.0 MPa. Relative water content (RWC) and the reciprocal of Psi(x) were analyzed by means of an unusual application of the pressure-volume relationship for determination of RWC of the apoplast (RWC(a)), osmotic potential at full turgor (Psi(oft)), and ratio of fully turgid weight to dry weight. Major varietal differences in drought response were in RWC(a) and needle cellulose content. The shaded seedlings showed tissue damage at relative water contents < 60%, and were killed by water deficits from which unshaded seedlings recovered. Correspondingly, shaded plants had significantly higher cell volume/cell mass ratio, Psi(oft), less cellulose in needle tissue, and lower RWC(a) than unshaded plants. These differences suggest that low irradiance restricts drought adaptation in ponderosa pine.     

Comparison of water-use efficiency of seedlings from two sympatric oak species: genotype x environment interactions

Seedlings of two sympatric oak species, Quercus robur L. and Quercus petraea (Matt.) Liebl., were grown in common garden conditions to test for potential interspecific differences in intrinsic water-use efficiency (WUE). Intrinsic water-use efficiency was estimated based on carbon isotope composition of shoots (delta13C) and on gas exchange measurements (ratio of net CO2 assimilation rate to stomatal conductance (A/g(sw))). In addition, genotype x environment interactions were tested by subjecting the seedlings to four irradiance treatments (8, 18, 48 and 100% of incident solar irradiance) imposed by neutral shading nets, and, in the 100% irradiance treatment, two watering regimes. In all treatments, initial growth of Q. robur was faster than that of Q. petraea. In both species, there was a tight correlation between delta13C and A/g(sw). Intrinsic water-use efficiency increased with increasing irradiance (almost doubling from 8 to 100% irradiance), and this effect paralleled the increase in A with increasing irradiance. In full sun, WUE of Q. petraea seedlings was 10-15% higher than in Q. robur seedlings, with the difference attributable to a difference between the species in g(sw). The interspecific difference in WUE was maintained during drought, despite the appreciable increase in WUE and decrease in growth imposed by drought. No interspecific differences in WUE were observed at low irradiances, suggesting a strong genotype x environment interaction for WUE. These findings confirm the existence of interspecific genetic differences in WUE, but also show that there is large intraspecific variability and plasticity in WUE. The initially greater height and biomass increments in Q. robur seedlings illustrate the ability of this species to out-compete Q. petraea in the early stages of forest regeneration. For adult trees growing in closed canopies, the high WUE of Q. petraea may contribute significantly to its survival during dry years, whereas the low WUE of Q. robur may account for the frequently observed declines in adult trees of this species following drought.     

Gas exchange,biomass, whole-plant water-use efficiency and water uptake of peach (Prunus persica) seedlings in response to elevated carbon dioxide concentration and water availability

We examined the interactive effects of elevated CO2 concentration ([CO2]) and water stress on growth and physiology of 1-year-old peach (Prunus persica L.) seedlings grown in 10-dm3 pots in open-top chambers with ambient (350 micromol mol-1) or elevated (700 micromol mol-1) [CO2]. Seedlings were supplied weekly with a non-limiting nutrient solution. Water was withheld from half of the plants in each treatment for a 4-week drying cycle, to simulate a sudden and severe water stress during the phase of rapid plant growth. Throughout the growing season, seedlings in elevated [CO2] had higher assimilation rates, measured at the growth [CO2], than seedlings in ambient [CO2], and this caused an increase in total dry mass of about 33%. Stomatal conductance, total water uptake, leaf area and leaf number were unaffected by elevated [CO2]. Because seedlings in the two CO2 treatments had similar transpiration despite large differences in total dry mass, water-use efficiency (WUE) of well-watered and water-stressed seedlings grown in elevated [CO2] was an average of 51 and 63% higher, respectively, than WUE of comparable seedlings grown in ambient [CO2]. Elevated [CO2] enhanced total biomass of water-stressed seedlings by 31%, and thus ameliorated the effects of water limitation. However, the percentage increases in total dry mass between well-watered and water-stressed seedlings were similar in ambient (53%) and elevated (58%) [CO2], demonstrating that there was no interaction between elevated [CO2] and water stress. This finding should be considered when predicting responses of trees to global climate change in hot and dry environments, where predicted temperature increases will raise evaporative demands and exacerbate the effects of drought on tree growth.     

Impacts of drought on mineral macro- and microelements in provenances of beech (Fagus sylvatica L.) seedlings

Beech seedlings originating from 11 German provenances with different climatic conditions were grown in pots and cultivated in a greenhouse. The composition of macro- and microelements in roots, axes and leaves was measured after half of the seedlings were subjected to a simulated summer drought. The recently described sensitivity of these provenances to drought was compared with drought-mediated changes in the elemental and ionic composition in organs of the seedlings; in addition, partitioning between roots and shoots was evaluated. A number of element concentrations were decreased in roots due to drought (K 94% of control, Mg 94%, Mn 75% and Zn 85%). However, chloride concentration increased in all organs (115-125%) and was the only element affected in leaves. Some changes in ionome can be related to sensitivity of provenances, but it is difficult to decide whether these changes are a result of, or a reason for, drought tolerance or sensitivity. Observed increases in chloride concentration in all plant parts of drought-treated beech seedlings can be explained by its function in charge balance, in particular since the level of phosphate was reduced. As a result of chloride accumulation, the sum of added charges of anions (and cations) in water extracts of leaf and root material was similar between drought and control plants. Since only the partitioning of Ca and Al (both only in axis) as well as Mn was affected and other elements (together with previously observed effects on C, N, S and P) remained unaffected by drought in all provenances, it can be concluded that direct effects by means of mass flow inhibition in xylem and phloem are unlikely. Secondary effects, for example on the pH of transport sap and the apoplastic space, cannot be excluded from the present study. These effects may affect partitioning between the apoplast and symplast and therefore may be significant for drought sensitivity.     

The Effects of Seed Mass on Germination, Seedling Emergence, and Early Seedling Growth of Eastern White Pine (Pinus strobus L.)

Half-sib seed of several eastern white pine (Pinus strobus L.) families was used to examine seed mass effects on laboratory germination, and seedling emergence and growth under moderate and low light (47 and 13% full sunlight) in a greenhouse. Percent germination and speed of germination under laboratory conditions were not related to seed mass among half-sib families or multi-family seedlots bulked by seed mass. Percent seedling emergence in the greenhouse was not related to seed mass, but families with heavier seeds exhibited faster emergence. Both rate and percent emergence were significantly increased under low light. Family differences in leaf, stem, root, and total seedling dry mass, primary root length, and the number of first-order-lateral-roots were positively related to seed mass in both light environments. Low light diminished the absolute biomass increment per unit seed mass, but the proportional change in biomass with seed mass was similar between light environments. Rate of emergence also influenced seedling size within families, with earlier emergence increasing seedling dry mass from 7 to 58%, dependent on light environment. Biomass partitioning coefficients were influenced by light environment but largely independent of seed mass.