Interacting effects of warming and drought on regeneration and early growth of Acer pseudoplatanus and A. platanoides.

Climate change is acting on several aspects of plant life cycles, including the sexual reproductive stage, which is considered amongst the most sensitive life-cycle phases. In temperate forests, it is expected that climate change will lead to a compositional change in community structure due to changes in the dominance of currently more abundant forest tree species. Increasing our understanding of the effects of climate change on currently secondary tree species recruitment is therefore important to better understand and forecast population and community dynamics in forests. Here, we analyse the interactive effects of rising temperatures and soil moisture reduction on germination, seedling survival and early growth of two important secondary European tree species, Acer pseudoplatanus and A. platanoides. Additionally, we analyse the effect of the temperature experienced by the mother tree during seed production by collecting seeds of both species along a 2200-km long latitudinal gradient. For most of the responses, A. platanoides showed higher sensitivity to the treatments applied, and especially to its joint manipulation, which for some variables resulted in additive effects while for others only partial compensation. In both species, germination and survival decreased with rising temperatures and/or soil moisture reduction while early growth decreased with declining soil moisture content. We conclude that although A. platanoides germination and survival were more affected after the applied treatments, its initial higher germination and larger seedlings might allow this species to be relatively more successful than A. pseudoplatanus in the face of climate change.

Rapid freezing injury in red spruce: seasonal changes in sensitivity and effects of temperature range.

On calm, cold days in winter, sun-exposed needles of red spruce (Picea rubens Sarg.) may warm 10 to 20 degrees C above ambient air temperature, and undergo rapid (>/= 1 degrees C min(-1)) fluctuations in temperature as light breezes or passing clouds alter the energy balance of the foliage. It has been proposed that the resulting rapid freeze-thaw cycles (freezing stress) cause a type of winter injury in montane red spruce that is characterized by necrosis of sun-exposed foliage. In autumn and winter, we monitored rapid freezing stress response of needle sections from 10 montane red spruce trees by subjecting needles to rapid freezing over the temperature span typically recorded in the field. In autumn, experimental rapid freezing stress produced severe injury only at temperatures considerably lower than expected for that time of year. In winter, rapid freezing caused occasional, moderate injury in fully hardened foliage of trees susceptible to both slow and rapid freezing. Seasonal changes in sensitivity to rapid and slow freezing were correlated, suggesting that environmental factors that are known to affect sensitivity to slow freezing may also affect sensitivity to rapid freezing. Experimental manipulation of the start and end temperatures of rapid freezing stress events showed that moderate to severe needle injury can occur in susceptible trees at temperature spans slightly more extreme than those typically recorded in the field. The extent of injury was similar at different starting temperatures if rapid freezing occurred over the same temperature span. Year-old foliage was consistently less sensitive to rapid freezing stress than current-year foliage, but some year-old foliage was damaged when the rapid freezing stress regime caused severe injury in current-year foliage. We conclude that rapid freeze-thaw cycles can explain light to moderate injury of current-year foliage, but they do not explain the more severe and widespread pattern of foliar damage that has occurred intermittently over at least the last 18 years.

Acid mist and soil Ca and Al alter the mineral nutrition and physiology of red spruce.

We examined the effects and potential interactions of acid mist and soil solution Ca and Al treatments on foliar cation concentrations, membrane-associated Ca (mCa), ion leaching, growth, carbon exchange, and cold tolerance of red spruce (Picea rubens Sarg.) saplings. Soil solution Ca additions increased foliar Ca and Zn concentrations, and increased rates of respiration early in the growing season (July). Soil Al treatment had a broad impact, reducing foliar concentrations of Ca, Mg, Mn, P and Zn, and resulting in smaller stem diameters, sapling heights and shoot lengths compared with soil treatments with no added Al. Aluminum treatment also reduced respiration when shoots were elongating in July and decreased net photosynthesis at the end of the growing season (September). Three lines of evidence suggest that Al-induced alterations in growth and physiology were independent of foliar Ca status: (1) Ca concentrations in foliage of Al-treated saplings were within the range of sufficiency established for red spruce; (2) mCa concentrations were unaffected by Al treatment; and (3) no Al x Ca interactions were detected. Acid mist treatment increased foliar Fe and K concentrations and increased leaching of Ca, Mg, Mn, Zn, Fe, and Al from foliage. Leaching losses of Ca were more than twice those of the element with the next highest amount of leaching (Zn), and probably led to the reductions in mCa concentration and membrane stability of acid-treated saplings. Acidic mist resulted in enhanced shoot growth, and consistent reductions in foliar cold tolerance in the fall and winter. Of the few significant interactions among treatments, most involved the influence of mist pH and Al treatment on foliar nutrition. In general, reductions in cation concentration associated with Al addition were greater for pH 5.0-treated saplings than for pH 3.0-treated saplings. We propose that H(+)-induced leaching of mCa from mesophyll cells is the mechanism underlying acid-induced reductions in foliar cold tolerance of red spruce.

Photosynthetic capacity of red spruce during winter.

We measured the photosynthetic capacity (P(max)) of plantation-grown red spruce (Picea rubens Sarg.) during two winter seasons (1993-94 and 1994-95) and monitored field photosynthesis of these trees during one winter (1993-94). We also measured P(max) for mature montane trees from January through May 1995. Changes in P(max) and field photosynthesis closely paralleled seasonal changes in outdoor air temperature. However, during thaw periods, field photosynthesis was closely correlated with multiple-day temperature regimes, whereas P(max) was closely correlated with single-day fluctuations in temperature. There was a strong association between short-term changes in ambient temperature and P(max) during the extended thaw of January 1995. Significant increases in P(max) occurred within two days of the start of this thaw. Repeated measurements of cut shoots kept indoors indicated that temperature-induced increases in P(max) can occur within 3 h. Although significant correlations between P(max) and stomatal conductance (g(s)) or intracellular CO(2) concentration (C(i)) raised the possibility that increases in P(max) resulted from increases in stomatal aperture, fluctuations in g(s) or C(i) explained little of the overall variation in P(max). Following both natural and simulated thaws, P(max) increased considerably but plateaued at only 37% of the mean photosynthetic rate reported for red spruce during the growing season. Thus, even though shoots were provided with near-optimal environmental conditions, and despite thaw-induced changes in physiology, significant limitations to winter photosynthesis remained.

Physiological implications of seasonal variation in membrane-associated calcium in red spruce mesophyll cells.

We examined the pattern of seasonal variation in total foliar calcium (Ca) pools and plasma membrane-associated Ca (mCa) in mesophyll cells of current-year and 1-year-old needles of red spruce (Picea rubens Sarg.) and the relationship between mCa and total foliar Ca on an individual plant and seasonal basis. Foliar samples were collected from seedlings and analyzed on 16 dated at 2- to 3-week intervals between June 1994 and March 1995. Concentrations of mCa in current-year needles were more seasonally dynamic and responsive to temporal environmental changes than either mCa concentrations of 1-year-old needles, which were largely stable, or total foliar Ca concentrations in both tissues. In current-year needles, mCa was barely evident in early summer, increased steadily through summer, and then increased dramatically in early fall and surpassed the concentration in 1-year-old needles. Coincident with the first severe frost, mCa concentrations in current-year needles declined significantly and subsequently maintained concentrations comparable to those of 1-year-old needles. Following an extended January thaw, which included 5 days of minimum temperatures > 5 degrees C, mCa concentrations of current-year needles temporarily, but significantly, declined. However, there was no change in mCa concentrations of 1-year-old needles or total Ca concentrations of either tissue. Total Ca concentrations were stable through midsummer in both tissues, doubled in late summer, and then were stable in both tissues throughout fall and winter. Total Ca concentrations were consistently higher in 1-year-old than in current-year needles. Correlations between concentrations of mCa and total foliar Ca were consistently low and mostly nonsignificant. Thus, the dominant, but insoluble, extracellular Ca pool reflected in commonly measured total foliar Ca concentrations is not a meaningful surrogate for the physiologically important and labile pool associated with the plasma membrane-cell wall compartment of red spruce mesophyll cells. It is likely that shifts in the critical mCa compartment would not be detected by analysis of total foliar Ca pools. Seasonal changes in mCa concentration seemed to parallel seasonal changes in membrane structure, and possibly the important role of extracellular Ca in transducing messages associated with environmental signals.

Physiological changes in red spruce seedlings during a simulated winter thaw.

We evaluated net photosynthesis, respiration, leaf conductance, xylem pressure potential (XPP) and cold hardiness in red spruce (Picea rubens Sarg.) seedlings exposed to either a continuous thaw (CT) or a daytime thaw with freezing nights (FN) for 8 days during mid-winter. Physiological differences between CT and FN seedlings were evident for all measured parameters. However, the temporal expression of treatment differences varied among parameters. When compared to FN seedlings, CT seedlings had higher rates of respiration following 24 h of treatment, and a higher net photosynthetic rate, leaf conductance and XPP after 48 h of treatment. The CT seedlings were significantly less cold tolerant than the FN seedlings following 4 days of thaw, whereas FN seedlings did not deharden over the 8 days of treatment. Examination of temporal trends among thaw-associated changes in physiology suggested that, although greater carbon exchange occurred as stomatal conductance increased, the transition from negative to positive net photosynthesis was not the result of increases in conductance, but may have been associated with thaw-induced increases in XPP. Because thaw-associated changes in gas exchange and cold hardiness were offset in time, we conclude that, if changes in these processes are physiologically linked, the linkage is indirect.

Effects of mist acidity and ambient ozone removal on montane red spruce.

Recent forest studies have established that high-elevation (> 900 m) populations of red spruce (Picea rubens Sarg.) in the northeastern USA are declining. Because it has been suggested that changes in air quality are responsible for the decline, we examined the effects of acidic mists and ozone on several biochemical and growth parameters in mature montane red spruce. We used branch-sized environmental chambers to introduce mists of controlled composition and exclude ambient clouds and ozone from individual branches within a tree. Mists consisting of distilled water increased the end-of-season pigment concentration and shoot length of enclosed branches relative to ambient or artificial mists. Needle and twig weights and starch concentrations were not significantly altered by the acidic mist treatments. Removal of ambient ozone had no apparent effect on the variables measured.

Midwinter needle temperature and winter injury of montane red spruce.

To assess the role of solar warming and associated temperature fluctuations in the winter injury of sun-exposed red spruce foliage, we used fine wire thermocouples to monitor midwinter needle temperature in the upper canopy of mature red spruce trees over two winters. In 1989-1990, 15-min mean temperatures were recorded for six needles in a single tree. In 1990-1991, 10-min mean temperatures of six needles in one tree, and 1-min mean temperatures of seven needles in a second tree were recorded during rapid temperature changes. Warming was more frequent and greatest on terminal shoots of branches with a south to southwest aspect. The maximum rise above ambient air temperature exceeded 20 degrees C, and the maximum one minute decrease in temperature was 9 degrees C, with maximum rates of 0.8 and 0.6 degrees C min(-1) sustained over 10- and 15-min intervals, respectively. These data demonstrate that red spruce is subject to rapid temperature fluctuations similar to those known to produce visible injury in American aborvitae, a much hardier species. We concluded that solar warming to temperatures above the freezing point was unlikely to result in dehardening and subsequent freezing injury, because warming was infrequent, of short duration, and did not always raise needle temperature above the freezing point. Parts of branches and some individual shoots were frequently covered by snow or rime that may have prevented injury by reducing the frequency or intensity of needle temperature fluctuations. Radiation load on exposed shoots may have been increased by reflection of short wave radiation from snow and rime deposits on surrounding surfaces, which would exacerbate temperature fluctuations.

In situ experimental freezing produces symptoms of winter injury in red spruce foliage.

Two mechanisms have been proposed to explain winter injury to needles of red spruce (Picea rubens Sarg.): (1) desiccation, which is characterized by net loss of foliar water from the needle to the environment, with cell injury resulting from dehydration; and (2) freezing, which is characterized by direct injury to cells resulting from intracellular or extracellular ice formation during exposure to low temperature. To compare the separate and combined effects of freezing and desiccation, branches of a mature red spruce at 1160 m were (a) experimentally frozen in situ to -50 degrees C; (b) cut and tied in their original orientation and allowed to desiccate passively; or (c) both frozen in situ and cut and tied in their original orientation. Needle water content, electrolyte leakage (an index of cell injury), and needle color were monitored for 60 days after treatment. Freezing resulted in immediate increases in electrolyte leakage, rapid water loss, and reddening necrosis of needles similar to that of naturally injured needles. Cutting resulted in more gradual water loss, no significant changes in electrolyte loss until severe desiccation had occurred, and a change in the color of the needles to a dull green. Because freezing produced reddening necrosis, a key symptom of winter injury, whereas desiccation did not, we conclude that freezing is probably the primary cause of winter injury in red spruce, and that desiccation is a secondary effect.