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Global climatic change may cause changes in regional precipitation that have important implications for forest growth in the southern United States. In 1993, a stand-level experiment was initiated on Walker Branch Watershed, Tennessee, to study the sensitivity of forest saplings and large trees to changes in soil water content. Soil water content was manipulated by gravity-driven transfer of precipitation throughfall from a dry treatment plot (-33%) to a wet treatment plot (+33%). A control plot was included. Each plot was 6400 m2. Measurements of stem diameter and observations of mortality were made on large trees and saplings of Acer rubrum L., Cornus florida L., Liriodendron tulipifera L., Nyssa sylvatica Marsh, Quercus alba L. and Quercus prinus L. every 2 weeks during six growing seasons. Saplings of C. florida and A. rubrum grew faster and mortality was less on the wet plot compared with the dry and control plots, through 6 years of soil water manipulation. Conversely, diameter growth of large trees was unaffected by the treatments. However, tree diameter growth averaged across treatments was affected by year-to- year changes in soil water status. Growth in wet years was as much as 2-3 times greater than in dry years. Relationships between tree growth, phenology and soil water potential were consistent among species and quantitative expressions were developed for application in models. These field growth data indicate that differences in seasonal patterns of rainfall within and between years have greater impacts on growth than percentage changes in rainfall applied to all rainfall events.

Direct (i.e., short-term) effects of elevated CO(2) on nocturnal in situ leaf respiration rate were measured in nine deciduous tree species (seven genera) in 20 3.5-4.0-h experiments. During the experiments, CO(2) concentration was alternated between 400 and 800 ppm (approximately 40 and 80 Pa of CO(2)). Data analysis accounted for effects on respiration rate of the normal decline in temperature with time after sunset. The median response to a 40-Pa increase in CO(2) was a 1.5% decrease in respiration rate, with responses ranging from a 5.6% inhibition to a 0.4% stimulation. Direct effects of elevated CO(2) on respiration were similar among the species. Thus, the response of nocturnal leaf respiration rate to a short-term CO(2) increase was small, and of little practical importance to the accuracy of measurements of respiration involving similar changes in CO(2) concentration during measurement. These direct respiratory responses of leaves to elevated CO(2) would translate into only slight, if any, effects on the carbon balance of temperate deciduous forests in a future atmosphere containing as much as 80 Pa CO(2).

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.

Elevated [CO2] and temperature may alter the drought responses of tree seedling growth, photosynthesis, respiration and total non-structural carbohydrate (TNC) status depending on drought intensity and duration. Few studies have addressed these important climatic interactions or their consequences. We grew Eucalyptus globulus Labill. seedlings in two [CO2] concentrations (400 and 640 μl l(-1)) and two temperatures (28/17 and 32/21 °C) (day/night) in a sun-lit glasshouse, and grew them in well-watered conditions or exposed them to two drought treatments having undergone different previous water conditions (i.e., rewatered drought and sustained drought). Progressive drought in both drought treatments led to similar limitations in growth, photosynthesis and respiration, but reductions in TNC concentration were not observed. Elevated [CO2] ameliorated the impact of the drought during the moderate drought phase (i.e., Day 63 to Day 79) by increasing photosynthesis and enhancing leaf and whole-plant TNC content. In contrast, elevated temperature exacerbated the impact of the drought during the moderate drought phase by reducing photosynthesis, increasing leaf respiration and decreasing whole-plant TNC content. Extreme drought (i.e., Day 79 to Day 103) eliminated [CO2] and temperature effects on plant growth, photosynthesis and respiration. The combined effects of elevated [CO2] and elevated temperature on moderate drought stressed seedlings were reduced with progressive drought, with no sustained effects on growth despite greater whole-plant TNC content.

We quantified leaf phenologies of saplings and overstory trees of sugar maple (Acer saccharum Marsh.) and American beech (Fagus grandifolia Ehrh.), and the shrub hobblebush viburnum (Viburnum alnifolium Marsh.) in a 72-year-old northern hardwood forest. Seasonal changes in irradiance in the shrub layer, and in the leaf CO(2) exchange of viburnum, and sugar maple and beech saplings were also measured. Leaf expansion occurred earlier in the spring and green leaves were retained later in the autumn in saplings and shrubs than in overstory trees. During the spring light phase (before overstory closure), large CO(2) gains by all three shrub-layer species occurred as a result of a combination of relatively large leaf area, high photosynthetic capacity, and high irradiance. Throughout the summer shade phase, photosynthetic capacity at a given irradiance remained relatively constant, but CO(2) gain was typically limited by low irradiances. Even though irradiance in the shrub layer increased during the autumn light phase as the overstory opened, CO(2) gains were modest compared to springtime values because of declining leaf area and photosynthetic capacity in all three species. The CO(2) gains during the spring light phase, and to a lesser extent during the autumn light phase, may be important to the carbon balance and long-term persistence of saplings and shrubs in the usually light-limited shrub layer of a northern hardwood forest. Therefore, for some late-successional species, leaf phenology may be an important characteristic that permits their long-term persistence in the shrub layer of mature northern hardwood forests.

Diurnal patterns of leaf conductance, net photosynthesis and water potential of five tree species were measured at the top of the canopy in a tropical lowland rain forest in southwestern Cameroon. Access to the 40 m canopy was by a large canopy-supported raft, the Radeau des Cimes. The measurements were made under ambient conditions, but the raft altered the local energy balance at times, resulting in elevated leaf temperatures. Leaf water potential was equal to or greater than the gravitational potential at 40 m in the early morning, falling to values as low as -3.0 MPa near midday. Net photosynthesis and conductance were typically highest during midmorning, with values of about 10-12 micro mol CO(2) m(-2) s(-1) and 0.2-0.3 mol H(2)O m(-2) s(-1), respectively. Leaf conductance and net photosynthesis commonly declined through midday with occasional recovery late in the day. Photosynthesis was negatively related to leaf temperature above midday air temperature maxima. These patterns were similar to those observed in other seasonally droughted evergreen communities, such as Mediterranean-climate shrubs, and indicate that environmental factors may cause stomatal closure and limit photosynthesis in tropical rain forests during the midday period.