Effects of atmospheric CO(2) on longleaf pine: productivity and allocation as influenced by nitrogen and water

Longleaf pine (Pinus palustris Mill.) seedlings were exposed to two concentrations of atmospheric CO(2) (365 or 720 micro mol mol(-1)) in combination with two N treatments (40 or 400 kg N ha(-1) year(-1)) and two irrigation treatments (target values of -0.5 or -1.5 MPa xylem pressure potential) in open-top chambers from March 1993 through November 1994. Irrigation treatments were imposed after seedling establishment (i.e., 19 weeks after planting). Seedlings were harvested at 4, 8, 12, and 20 months. Elevated CO(2) increased biomass production only in the high-N treatment, and the relative growth enhancement was greater for the root system than for the shoot system. In water-stressed trees, elevated CO(2) increased root biomass only at the final harvest. Root:shoot ratios were usually increased by both the elevated CO(2) and low-N treatments. In the elevated CO(2) treatment, water-stressed trees had a higher root:shoot ratio than well-watered trees as a result of a drought-induced increase in the proportion of plant biomass in roots. Well-watered seedlings consistently grew larger than water-stressed seedlings only in the high-N treatment. We conclude that available soil N was the controlling resource for the growth response to elevated CO(2) in this study. Although some growth enhancement was observed in water-stressed trees in the elevated CO(2) treatment, this response was contingent on available soil N.     

Growth and thinning effects during a rotation period of hybrid aspen in southern Sweden

Hybrid aspen is an interesting tree species for wood production in northern Europe. In this study we examined growth dynamics over the whole rotation period. Height and diameter development, as well as annual growth of stem volume and stem biomass, were repeatedly recorded in 14 planted and 2 root sucker stands, aged up to 26 years, in southern Sweden. A main aim was to study the productivity level for hybrid aspen forestry with an expected rotation period of about 25 years. The study verified earlier prognoses, showing a mean annual increment (MAI) of 19.5 m³ of stem wood ha^?1 yr^?1 after 25 years. This corresponds to a dry weight of stem biomass of 7.2 tons dry matter (DM) ha^?1 yr^?1. Maximum MAI was still not reached after 25 years, although the growth curve was flattening out. If branch biomass is included, MAI is estimated to about 9 tons DM ha^?1 yr^?1 and further improvements in growth is expected by using the best genotypically selected clones available. Results from three different thinning regimens showed that thinning intensity provided significantly larger diameter growth, while no significant yield effects were seen among the thinning regimens.     

Short-day treatment during the growing period limits shoot growth and increases frost hardiness of hybrid aspen plants in the nursery

In Finland, under nursery conditions hybrid aspen may continue their shoot growth until early September. Thus, frost hardening is usually delayed. To solve this problem, we used a three-week period of short-day (SD) treatment between late July and mid-August. During autumn after frost exposure, frost hardiness (FH) was assessed three times with a stem-browning test. The re-sults showed that after SD treatment shoot growth ceased and FH increased when compared to untreated hybrid aspen. Furthermore, the height of SD-treated hybrid aspen varied much less than that of the control plants. We conclude that SD treatment in the nursery during the growing period can be used as a supplementary method for producing well-hardened and uniform hybrid aspen plants.     

Histology of magnesium-deficient Norway spruce needles influenced by nitrogen source

Effects of magnesium deficiency and variation in nitrate to ammonium ratio on needle histology and chlorophyll concentration were investigated in current-year and one-year-old needles of clonal Norway spruce trees (Picea abies (L.) Karst.). Six-year-old trees were grown for one year in sand culture with circulating nutrient solutions containing a sufficient (0.2 mM) or a limiting (0.04 mM) concentration of Mg. The nitrogen concentration was not varied (5 mM), but the NO(3) (-)/NH(4) (+)-ratio was adjusted to 0.76 in Mg-sufficient and to 1.86, 0.76 or 0.035 in Mg-limited plants. Visible symptoms of Mg deficiency occurred only in current-year needles, indicating adequate Mg nutrition before the experiment. Under conditions of Mg limitation, chlorophyll and Mg concentrations were lowest in needles of trees supplied with NH(4) (+) as the major nitrogen source and highest in needles of trees supplied with NO(3) (-) as the major nitrogen source. In current-year and one-year-old needles, starch accumulation induced by Mg deficiency was increased when NH(4) (+) was the major nitrogen source. The accumulation of tannin spherules in current-year needles, which occurred in response to Mg deficiency, also increased with decreasing NO(3) (-)/NH(4) (+)-ratios. Deficient Mg supply caused premature aging in tissues of the vascular bundle, as indicated by modifications of the cambium and increased amounts of collapsed sieve cells. The number of collapsed sieve cells was slightly lower in needles grown in a NH(4) (+)-dominated nutrient regime than in needles grown in a NO(3) (-)-dominated nutrient regime. We conclude that was not directly toxic to Norway spruce trees at the applied concentrations. However, effects of Mg deficiency were considerably greater in an NH(4) (+)-dominated nutrient regime than in a NO(3) (-)-dominated nutrient regime.     

Uptake and distribution of calcium and phosphorus in beech (Fagus sylvatica) as influenced by aluminum and nitrogen

We studied the effects of excess nitrogen added as nitrate (NO(3) (-)) or ammonium (NH(4) (+)), or both, on mineral nutrition and growth of beech (Fagus sylvatica L.) plants grown at pH 4.2 in Al-free nutrient solution or in solutions containing 0.1 or 1.0 mM AlCl(3). A high external concentration of NH(4) (+) increased the concentration of nitrogen in roots, stems and leaves. The root/shoot dry weight ratio was less in plants grown in the presence of NH(4) (+) than in plants grown in the presence of NO(3) (-). The concentration of phosphorus in the roots was increased and the concentration of potassium in all parts of the plant was decreased by NH(4) (+). A high external concentration of NO(3) (-) caused a decrease in phosphorus concentrations of the root, stem and leaf. Uptake of (45)Ca(2+) by roots was reduced in the presence of high concentrations of NH(4) (+) or NO(3) (-), and a combination of high concentrations of nitrogen and aluminum further reduced the uptake of (45)Ca(2+). Uptake of phosphate ((32)P) and concentrations of phosphorus in root and shoot were increased when plants were grown in the presence of 0.1 mM Al. Exposure to 1.0 mM Al, however, reduced the concentration of phosphorus in roots and shoots and the reduction was greater when plants were grown in the presence of a high external NO(3) (-) concentration. Aluminum binds to roots, and plants grown in the presence of 1.0 mM Al had a slightly higher concentration of aluminum in roots than plants grown in the presence of 0.1 mM Al, whereas the concentration of Al in the shoot was increased 2 to 3 times in plants exposed to 1.0 mm Al. Furthermore, the effects of 1.0 mM Al on uptake of other macronutrients were quite different from the effects of 0.1 mM Al. We conclude that 0.1 mM Al facilitates uptake and transport of phosphorus in beech and that between 0.1 and 1.0 mM Al there is a dramatic change in the effects of Al on uptake and transport of divalent cations and phosphorus.     

Growth,productivity and water use in three hybrid poplar clones

Three drought treatments (none, medium and severe) were applied to three container-grown, hybrid poplar clones (Beaupré, Trichobel and Ghoy) as a series of drying cycles over two years-in a greenhouse in 1994 and outside in 1995. Leaf area development, annual biomass production and annual water use were recorded. Clone Beaupré had greater rates of leaf area development (up to 11.8 x 10(-4) m(2) degrees C day(-1)) and subsequently greater dry matter production (up to 816 g tree(-1) year(-1)) and water use (up to 168 l tree(-1) year(-1)) than the other two clones. In all clones, rates of leaf area development, annual biomass production and water use were less under drought conditions than under well-watered conditions. On a proportional basis, the highest reduction in biomass by drought was in Trichobel (up to 74%), whereas, in absolute terms, the reduction was greatest in Beaupré (up to 500 g tree(-1)). However, under drought conditions, annual biomass production was greater in Clone Beaupré than in the other clones. The relationship between water use and biomass production was constant over both years, and was similar for Beaupré and Trichobel at 4.4 g l(-1), but was less for Ghoy at 3.5 g l(-1).     

Effect of oil impregnation on water repellency,dimensional stability and mold susceptibility of thermally modified European aspen and downy birch wood

Conventional chemical wood preservatives have been banned or restricted in some applications due to human and animal toxicity and their adverse impact on the surrounding environment. New, low-environmental-impact wood treatments that still provide effective protection systems are needed to protect wood. Thermal modification of wood could reduce hygroscopicity, improve dimensional stability and enhance resistance to mold attack. The aim of this study was to investigate if these properties enhanced in thermally modified (TM) wood through treatments with oils. In this study, TM European aspen (Populus tremula) and downy birch (Betula pubescens) wood were impregnated with three different types of oil: water-miscible commercial Elit Träskydd (Beckers oil with propiconazole and 3-iodo-2-propynyl butylcarbamate, IPBC), a pine tar formulation and 100% tung oil. The properties of oil-impregnated wood investigated were water repellency, dimensional stability and mold susceptibility. The treated wood, especially with pine tar and tung oil, showed an increase in water repellency and dimensional stability. However, Beckers oil which contains biocides like propiconazole and IPBC showed better protection against mold compared with pine tar and tung oil. To enhance the dimensional stability of the wood, pine tar and tung oil can be used, but these oil treatments did not significantly improve mold resistance rather sometimes enhanced the mold growth, whereas a significant anti-mold effect was observed on Beckers oil treated samples.     

Stem dimensional fluctuation in Jeffrey pine from variation in water storage as influenced by thinning and prescribed fire

Forest thinning utilizing cut-to-length and whole-tree harvesting systems with subsequent underburning were assessed for their impacts on water storage in the extensible tissues of dominant and codominant trees in an uneven-aged Jeffrey pine (Pinus jeffreyi Grev. & Balf.) stand on the east slope of the Sierra Nevada. Prior to the onset of the third growing season following thinning and the second season after burning, manual band dendrometers were installed at breast height on the selected trees and readings of diurnal fluctuation in stem circumference, an indication of bole water status, were taken monthly for one year. Diameter and relative diameter fluctuation were calculated from the circumference measurements. Overall, thinning had a positive influence on stem water recharge capacity, with the most pronounced effects evident in the latter part of the growing season. During this period, bole contraction in thinned stand portions was 49 to 55% greater than in the unthinned control, suggesting that both a greater volume of stored water was available for transpiration and was transpired in trees of the former treatment. There was no clear evidence that harvesting method affected stem water storage and influences of underburning were also absent entirely. Seasonal effects on diurnal changes in stem diameter were prominent, as the extent to which boles contracted generally increased over the course of the growing season, whereas fluctuations were at a minimum during the colder months. The magnitude of stem dimensional flux was found to be negatively correlated with initial tree DBH in one instance, while negative relationships between the former and live crown length as well as percentage were also revealed, albeit infrequently. Changes in bole size were positively correlated with residual basal area in some cases. These results suggest that improvement in water relations can be realized from density management in a dry site forest type with no apparent compromise of this benefit by broadcast underburning.     

Growth and nitrogen status of young walnuts as affected by intercropped legumes in a Mediterranean climate

Walnut trees (Juglans nigra L. and Juglans nigra × regia NG23) were intercropped with alfalfa (Medicago sativa L.) or sainfoin (Onobrychis sativa L.) forage legumes compared to a grass (Festuca arundinacea Schr., fescue), or to spontaneous weeds as a control in two Mediterranean sites (Castries and Notre-Dame de Londres) near Montpellier (France). Tree growth, soil water depletion and nitrogen content of the tree leaves were monitored to assess the impact of both water competition and possible facilitation resulting from fixed nitrogen transfer from the leguminous crops to the trees. At Castries, where alfalfa and fescue were compared, they were found to have the same impact on tree growth. At Notre-Dame de Londres where sainfoin was compared with a spontaneous grassing treatment, the sainfoin crop was more competitive to the walnuts. The nitrogen content of walnut leaves was enhanced when intercropped with nitrogen fixing species at both sites. Perennial leguminous intercrops were more competitive for soil water resources than fescue or weeds, but in the long term this may be compensated by the improved nitrogen status of the trees. This compensating effect was observed during a rather rainy year. The overall impact of leguminous intercrops on tree growth may depend on the frequency of dry (competitive) and wet (non-competitive) years. This revised version was published online in June 2006 with corrections to the Cover Date.     

Fungal decomposition of sapwood and heartwood of European aspen,Populus tremula L.

In contrast to the woodrotting fungi which can only attack Populus tremula after felling, Phellinus tremulae successfully invades the central part of the living tree. Possible causes for this difference were investigated. It was found that sterile wood samples were decomposed more rapidly by saprophytic fungi than by parasitic fungi. With both types of fungi, heartwood was more resistant to decay than sapwood. It is postulated that living sapwood inhibits infection by most woodrotting fungi, but that P. tremulae can successfully overcome this barrier. This postulate cannot be confirmed by experiments using sterilized wood samples.     

Weed emergence as influenced by soil moisture and air temperature

Emergence of weed seedlings depends on soil environmental conditions; mainly temperature and moisture, with the latter being fundamental and particularly important in environments which are characterised by irregular amounts and distribution of rainfall throughout the year. Thus, this study looks at the influence of soil moisture and air temperature on the emergence of weed seedlings. The experiment was carried out under controlled environmental conditions, using rings filled with samples of undisturbed topsoil (0–2.5 cm and 0–5 cm deep) of a Luvisol. There were four moisture levels that were maintained constant, with several repetitions (16–20). The results indicate that the maximum population density of weed plants was obtained with soil moisture near field capacity. With regard to the depth of the soil containing the seeds which contributed towards optimising population density, it was noted that the first 2–3 cm of undisturbed soil were critical for maximising the population. The emergence of seedlings was modelled using data from the 5 cm topsoil with soil moisture close to field capacity. This model indicates that a mean daily temperature sum of 446°C, which under Mediterranean autumn conditions represent a period of approximately one month, is needed, in order to obtain 80% of potential autumn-winter plant density, relative to the observed potential.     

Growth and herbage productivity of Gliricidia in Panicum maximum pasture as influenced by seedbed preparation,planting and weed control techniques

Interplanting Gliricidia (Gliricidia sepium [Jacq.] Walp.) into poor quality native grass can improve the quality of pasture. But information on methods to establish the tree legume into existing pasture is scarce. This study was designed to compare the effects of disc-ploughing, hoeing, or rotary tillage operations, and direct seeding or planting out potted seedlings on growth and dry matter yields of Gliricidia in grass pasture. The Guinea grass (Panicum maximum) pasture was first mowed to a height of 10 cm. Then the seedbed was prepared by either hoeing 0.5 m strip or rototilling 0.5 m strip with a 2-wheel tractor or disc-ploughing 1.0 m strip with a 4-wheel tractor. Eight weeks old potted seedlings were planted or seeds were sown direct. At 6 months after planting (MAP), trees in disc-ploughed strips averaged 81 cm in height, about 35% taller than trees in hoed- or rotary-tilled strips. Potted seedlings (average height 77 cm) grew 37% taller than direct-seeded trees. At 12 MAP trees in disc-ploughed strips yielded 95% more leaf DM (1170 vs. 600 kg/ha) than trees in hoed- or rotary-tilled strips. In order to intercrop Gliricidia into existing Guinea grass pasture, it is necessary to plant potted seedlings previously raised in nursery into 1.0 m wide strips disc-ploughed with 4-wheel tractor. Weed control is necessary.     

Effects of browsing on recruitment and mortality of European aspen (Populus tremula L.)

Abstract

High browsing pressure may reduce or halt regeneration of browsing-prone tree species. In this study, exclosures were used to quantify the effects of browsing on vital demographic parameters in European aspen (Populus tremula L.) stands. The experiment focused on stands located in managed forest landscape with densities of 0.85 moose (Alces alces L.)km^?2. Other herbivores such as mountain hare (Lepus timidus L.) were present at densities common for this region and roe deer (Capreolus capreolus L.) at low densities. Recruitment and mortality of aspen ramets were recorded before and after the growing season, and over four consecutive years. Seven to 19% of the ramets within browsing reach were browsed annually. No differences were found in recruitment or mortality attributable to browsing per se. Instead, there were large among-stand variability in vital population parameters unrelated to browsing incidence. In general, aspen stands with high ramet density showed higher recruitment rates than stands with low ramet density. There was substantial variability in self-replacement capability unrelated to ecological factors. The results suggest that more attention should be given to finding and managing stands with high capacity for suckering, as these stands may have a disproportionate impact on aspen recruitment at the landscape scale.     

Growth and photosynthesis of tropical forest tree seedlings (Bischofia javanica Blume) as influenced by a change in light availability

Acclimation in seedlings of Bischofia javanica Blume, which are commonly found in canopy gaps in the moist forests of tropical Asia, to a change in light availability was examined in a controlled environment simulating forest shade and daylight. Seedlings were grown in a high (1000 micro mol m(-2) s(-1); red/far-red, 1.45) or low (40 micro mol m(-2) s(-1); red/far-red, 0.10) light regime and then transferred to the contrasting light environment for nine weeks. Control seedlings were maintained in the same light regime throughout the study. The availability of light influenced relative growth rate through morphological and physiological adjustments. Transferred seedlings retained the leaves that had been developed before transfer, and no leaf-shedding was observed till the end of the experiment. Leaves formed in the new light regime were physiologically and morphologically identical to those of the corresponding controls. High-light seedlings transferred to low light displayed significantly lower relative growth rate than the low-light controls because of a lower leaf area ratio carried over from the previous high-light environment. A reverse pattern of response with respect to relative growth rate was observed for the low-light seedlings transferred to high light compared to the high-light controls. The higher relative growth rate in the low-light seedlings transferred to high light was the result of higher net assimilation rate and higher leaf area ratio. The higher leaf area ratio in the low-light seedlings transferred to high light was the consequence of the effects of previous environment, and the relatively lower net assimilation rate in the high-light control seedlings was, at least partly, due to the effects of self-shading rather than to the photosynthetic capacity of the leaves. The results suggest that the species has a wide acclimation potential to a change in light availability that might occur in nature following gap creation or canopy closure.     

The impact of nitrogen deposition on carbon sequestration by European forests and heathlands

In this study, we present estimated ranges in carbon (C) sequestration per kg nitrogen (N) addition in above-ground biomass and in soil organic matter for forests and heathlands, based on: (i) empirical relations between spatial patterns of carbon uptake and influencing environmental factors including nitrogen deposition (forests only), (ii) ¹⁵N field experiments, (iii) long-term low-dose N fertilizer experiments and (iv) results from ecosystem models. The results of the various studies are in close agreement and show that above-ground accumulation of carbon in forests is generally within the range 15–40 kg C/kg N. For heathlands, a range of 5–15 kg C/kg N has been observed based on low-dose N fertilizer experiments. The uncertainty in C sequestration per kg N addition in soils is larger than for above-ground biomass and varies on average between 5 and 35 kg C/kg N for both forests and heathlands. All together these data indicate a total carbon sequestration range of 5–75 kg C/kg N deposition for forest and heathlands, with a most common range of 20–40 kg C/kg N. Results cannot be extrapolated to systems with very high N inputs, nor to other ecosystems, such as peatlands, where the impact of N is much more variable, and may range from C sequestration to C losses.     

Initial sprouting, growth and mortality of European aspen and birch after selective coppicing in central Sweden

For reasons of aesthetics, sheltering, biodiversity, and children’s need of playgrounds, selective coppicing could be a useful and effective silvicultural method in urban areas. A randomised block experiment was carried out in three broad-leaved coppices situated within the fenced area of Arlanda Airport to compare a selective coppice regime (SCR), where all trees above a certain height limit were cut, with a traditional coppice regime, where all trees were felled. The effects of the treatments of the dominating tree species European aspen (Populus tremula L.) and birch (Betula pendula Roth and B. pubescens Ehrh.) were studied. During three growing seasons after performed treatment, the proportion of sprouting stools and root shooting of European aspen were reduced in SCR. Birch stools had fewer sprouts stool⁻¹, and sprouts of both birch and European aspen tended to be shorter in SCR. There was a tendency that the sprout mortality was lower in SCR for both birch and European aspen. Residual stands of SCR consisted of proportionally less birch, and more rowan, bird cherry, oak and other species. Coppicing with single tree selection from above suppressed light demanding tree species and promoted the more shade-tolerant species compared to traditional coppicing.     

Seed and seedling characteristics of hybrid chestnuts (Castanea spp.) derived from a backcross blight-resistance breeding program

New Forests - American chestnut (Castanea dentata) was a foundation species in the eastern United States until chestnut blight (Cryphonectria parasitica) infestation resulted in range-wide...     

水生植物对富营养化水体去除氮磷研究

选取5种典型水生植物菖蒲、千屈菜、旱伞草、水葫芦、黑藻,研究各单种植物单元对模拟富营养化水体生态系统的净化效果.结果表明:5种水生植物均能在富营养化水体中正常生长,对水体中的氮、磷均有一定的净化效果.5种植物去氮能力的大小依次为水葫芦>黑藻>旱伞草>菖蒲>千屈菜,5种植物去磷能力的大小依次为水葫芦>旱伞草>黑藻>菖蒲>千屈菜;5种植物对氮的累积率为10.49％—60.73％,对磷的累积率为13.45％—43.55％;5种植物对氮、磷的去除贡献率分别为61.65％—75.93％,45.66％—68.81％.试验证明,5种植物均可有效应用于城市富营养化水体中氮、磷的净化.     

Growth, nutrition, photosynthesis and transpiration responses of longleaf pine seedlings to light, water and nitrogen

Early growth and physiology of longleaf pine (Pinus palustris Mill.) seedlings were studied in response to light, water and nitrogen under greenhouse conditions. The experiment was conducted with 1-year-old seedlings grown in 11.3 l pots. The experimental design was a split-plot factorial with two levels (low and high) of each of the factors, replicated in three blocks. The four factorial combinations of water and nitrogen were randomly applied to 15 pots (sub-plots) in each of the light treatment (main plot). Data were collected on survival, root collar diameter (RCD), and height on a monthly basis. Biomass (shoot, root and needle), leaf area index, specific needle area, and needle nutrient (N, P, K, Ca, and Mg) concentrations were determined following final harvest after 16 months. Physiological data (net photosynthesis and transpiration) were collected monthly from March to July during the second growing season.

Height and RCD were significantly influenced by nitrogen and water and by the interaction between them with no apparent effect of light. Seedlings grew 93% taller in the high nitrogen and well watered (HNWW) treatment compared to the low nitrogen and water stressed (LNWS) treatment. Similarly, a significant increase (78%) in RCD was observed for seedlings in the HNWW treatment over the LNWS treatment. Light, along with water and nitrogen, played an important role in seedling biomass growth, especially when water was not limiting. Biomass partitioning (as measured by root:shoot ratio) was affected only by nitrogen and water. Nutrient stress had a greater influence on carbon allocation (69% increase in root:shoot ratio) than water stress (19% increase). Net photosynthesis (P_net) was significantly higher for seedlings in the high resource than in the low resource treatments with significant light×water and nitrogen×water interactions. Transpiration rate was higher (75%) under the WW treatment compared to the WS treatment. Longleaf pine seedlings grown under the LNWW treatment had the lowest foliar nitrogen (0.71%) whereas seedlings in the HNWS treatment had the highest (1.46%). Increasing the availability of light (through larger canopy openings or controlling midstory density) and soil nitrogen (through fertilization) may not result in greater P_net and improved seedling growth unless soil water is not limiting.     

Root water flow and growth of aspen (Populus tremuloides) at low root temperatures

Effects of root zone temperature on growth, shoot water relations, and root water flow were studied in 1-year-old aspen (Populus tremuloides Michx.) seedlings. Seedlings were grown in solution culture and exposed to day/night air temperatures of 22/16 degrees C and solution culture temperatures of 5, 10, or 20 degrees C for 28 days after bud flush. Compared with root growth at 20 degrees C, root growth was completely inhibited at 5 degrees C and inhibited by 97% at 10 degrees C. The 5 and 10 degrees C treatments severely reduced shoot growth, leaf size, and total leaf area. Root water flow was inhibited by the 5 and 10 degrees C treatments. However, when seedlings were grown for 28 days at 5 degrees C and root water flow was measured at 20 degrees C, there was an increase in flow rate. This increase in root water flow was similar in magnitude to the decrease in root water flow observed when seedlings were grown for 28 days at 20 degrees C and root water flow was measured at 5 degrees C. Reduced root water flow of seedlings grown at 5 and 10 degrees C resulted in decreased stomatal conductance, net assimilation, and shoot water potentials. Root water flow was positively correlated with leaf size, total leaf area, shoot length, and new root growth. Transferring seedlings from 5 to 20 degrees C for 24 h significantly increased root water flow, shoot water potential, and net photosynthesis, whereas transferring seedlings from 10 to 20 degrees C resulted in only a slightly increased shoot water potential. Transferring seedlings from 20 to 5 degrees C greatly reduced root water flow, stomatal conductance, and net photosynthesis, whereas shoot water potential decreased only slightly.