Cold tolerance and water content of current-year red spruce foliage over two winter seasons

Red spruce (Picea rubens Sarg.) in high elevation forests of northeastern North America suffers from frequent and severe winter injury, leading to apical dieback, decreased growth, and high mortality. To examine the role of winter desiccation and freezing injury in winter damage, weekly assessments of cold tolerance and water content were made on current-year foliage collected from native red spruce trees at a high elevation site over two winter seasons. In both years, foliage maintained high water contents and adequate cold tolerance; nonetheless, slight to moderate injury was observed each year on some trees. Despite brief thaw periods each winter, no mid-winter dehardening sufficient to put foliage at risk of freezing injury was evident. These findings suggest that, at least in some years, winter injury to current-year red spruce foliage is produced by a mechanism other than desiccation or absolute low temperatures.     

Hydraulic and stomatal adjustment of Norway spruce trees to environmental stress

A study of how the water conducting systems of 30-50-year-old Norway spruce (Picea abies (L.) Karst.) trees growing at three sites adjust to shade and waterlogging indicated that water relations characteristics varied with the life histories of the trees. Xylem was more efficient at conducting water and stomata were more sensitive to atmospheric evaporative demand in trees subjected to favorable growth conditions (control trees) than in trees growing in shade or waterlogged conditions. At the same soil water availability, shade-grown trees suffered more severely from water deficit than control trees. Under conditions of high atmospheric vapor pressure deficit, foliage of shade-grown trees exhibited low water potentials, as a result of low hydraulic conductance of the vascular system and inefficient stomatal control. Because of the increased internal resistance to water flow, more negative leaf water potentials (Psi(x)) must be reached to provide an adequate water supply to the foliage. It is concluded that dynamic water stress is one of the main causes of the continuing growth retardation in suppressed Norway spruce trees after their release from the overstory. Trees growing in waterlogged soil (bog-grown trees) were characterized by weak stomatal control, resulting in large water losses from the foliage. Although bog-grown trees exhibited uneconomical water use, they possessed mechanisms (e.g., osmotic adjustment) that allowed leaves to tolerate low Psi(x) while stomata remained open. Under conditions of sufficient soil water availability and moderate atmospheric vapor pressure deficit, soil-to-leaf conductance was highest in bog-grown trees (1.45 +/- 0.06 mmol m(-2) s(-1) MPa(-1)), followed by control and shade-grown trees (1.04 +/- 0.04 and 0.77 +/- 0.05 mmol m(-2) s(-1) MPa(-1), respectively). The lowest soil-to-leaf conductance (0.45 +/- 0.04 mmol m(-2) s(-1) MPa(-1)) was recorded in control trees at high atmospheric evaporative demand, and was probably caused by tracheid cavitation.     

Planting stress in newly planted jack pine and white spruce. 2. Changes in tissue water potential components

Water relations of bare-root jack pine (Pinus banksiana Lamb.) and white spruce (Picea glauca (Moench) Voss) planted in a greenhouse and on a boreal cut-over site were examined during the first growing season. In field-planted trees, maximum stomatal conductances (g(wv)) were initially low (< 0.10 cm s(-1)). Base and minimum xylem pressure potentials (Psi(x(base)) and Psi(x(min))) were less than -1.5 and -1.7 MPa for jack pine and -2.0 and -2.6 MPa for white spruce, respectively. During the growing season, maximum g(wv) increased in both species to around 0.2 cm s(-1). Base and minimum xylem pressure potentials also increased in both species to around -0.5 and -1.0 MPa in jack pine and -1.0 and -1.5 MPa in white spruce, respectively. Minimum xylem pressure potentials in white spruce fell below the turgor loss point during the first half of the growing season. Osmotic potential at the turgor loss point Psi(pi(TLP)) decreased after field planting to around -2.7 and -2.3 MPa in jack pine and white spruce, respectively. In the greenhouse, minimum values of Psi(pi(TLP)) were -2.2 and -2.3 MPa in jack pine and white spruce, respectively. Maximum bulk modulus of elasticity was greater in white spruce and underwent greater seasonal change than in jack pine. Relative water content (RWC) at turgor loss ranged between 71 and 74% in jack pine and 80 and 87% in white spruce. Available turgor (T(avail)), defined as the integral of turgor over the range of RWC between Psi(x(base)) and xylem pressure potential at the turgor loss point, was similar in jack pine and white spruce just after field planting. For the rest of the growing season, however, T(avail) in jack pine was two to three times that in white spruce. Diurnal turgor (T(diurnal)), defined as the integral of turgor over the range of RWC between Psi(x(base)) and Psi(x(min)), as a percent of T(avail) was higher in field-planted white spruce than jack pine until the end of the season. Dynamics of tissue water potential components are discussed in relation to plantation establishment.     

Leaf gas exchange of understory spruce-fir saplings in relict cloud forests, southern Appalachian Mountains, USA

The southern Appalachian spruce-fir (Picea rubens Sarg. and Abies fraseri (Pursh) Poir.) forest is found only on high altitude mountain tops that receive copious precipitation ( > 2000 mm year(-1)) and experience frequent cloud immersion. These high-elevation, temperate rain forests are immersed in clouds on approximately 65% of the total growth season days and for 30-40% of a typical summer day, and cloud deposition accounts for up to 50% of their annual water budget. We investigated environmental influences on understory leaf gas exchange and water relations at two sites: Mt. Mitchell, NC (MM; 35 degrees 45'53' N, 82 degrees 15'53' W, 2028 m elevation) and Whitetop Mtn., VA (WT; 36 degrees 38'19' N, 81 degrees 36'19' W, 1685 m elevation). We hypothesized that the cool, moist and cloudy conditions at these sites exert a strong influence on leaf gas exchange. Maximum photosynthesis (A(max)) varied between 1.6 and 4.0 micromol CO(2) m(-2) s(-1) for both spruce and fir and saturated at irradiances between approximately 200 and 400 micromol m(-2) s(-1) at both sites. Leaf conductance (g) ranged between 0.05 and 0.25 mol m(-2) s(-1) at MM and between 0.15 and 0.40 mol m(-2) s(-1) at WT and was strongly associated with leaf-to-air vapor pressure difference (LAVD). At both sites, g decreased exponentially as LAVD increased, with an 80-90% reduction in g between 0 and 0.5 kPa. Predawn leaf water potentials remained between -0.25 and -0.5 MPa for the entire summer, whereas late afternoon values declined to between -1.25 and -1.75 MPa by late summer. Thus, leaf gas exchange appeared tightly coupled to the response of g to LAVD, which maintained high water status, even at the relatively low LAVD of these cloud forests. Moreover, the cloudy, humid environment of these refugial forests appears to exert a strong influence on tree leaf gas exchange and water relations. Because global climate change is predicted to increase regional cloud ceiling levels, more research on cloud impacts on carbon gain and water relations is needed to predict future impacts on these relict forests.     

Shoot water relations of mature black spruce families displaying a genotype x environment interaction in growth rate. I. Family and site effects over three growing seasons

Pressure-volume curves were determined for black spruce (Picea mariana (Mill.) BSP) trees from four full-sib families. During the first two years, trees were measured from a plantation on a dry site. In the third year, trees were sampled from the dry site and a wet site. Diurnal measurements of shoot water potential allowed in situ shoot turgor to be estimated in addition to standard water relations traits. Over all years, Female 59 progeny displayed lower osmotic potentials at saturation (Psi(sat)) than Female 63 progeny. Genetic differences in Psi(sat) were similar on both the dry and wet sites. Modulus of elasticity (epsilon) was greater for Female 59 progeny than for Female 63 progeny, producing a compensatory effect resulting in no genetic or site differences in osmotic potential at turgor loss point (Psi(tlp)) or relative water content at turgor loss point (RWC(tlp)). Mean and predawn shoot turgor pressures (P(x) and P(pd)) were higher for Female 59 progeny than for Female 63 progeny and higher at the wet site than the dry site. Genotype x environment trends were observed; compared to Female 63 progeny, Female 59 progeny displayed 9.8 and 5.1% higher P(pd) on the dry and wet sites, respectively, and 3.4 and 9.8% greater P(pd) values in wet and dry years, respectively. Tree volume growth showed no relationship to Psi(tlp) or RWC(tlp), but was correlated with Psi(sat) and P(x); however, the strongest correlation was with P(pd) (r = 0.90).     

Winter desiccation and injury of subalpine red spruce

Montane red spruce (Picea rubens Sarg.) in the northeastern United States has undergone a decline during the past two decades. One symptom associated with the decline syndrome is the episodic browning of first-year foliage in early spring. To examine the potential role of winter desiccation in this browning, the water relations of red spruce foliage in a subalpine forest on Mt. Moosilauke, New Hampshire, USA, were monitored from January to May, 1989. All sampled trees lost water during the winter and the first-year foliage on some trees turned brown in early spring. The relative water content of first-year shoots during the winter was a significant predictor of spring browning; red spruce trees that showed browning had desiccated faster and reached lower relative water contents. Damaged trees also had more closely packed needles and lower cuticular resistances to water loss. The first-year shoots had a significantly lower average relative water content than older shoots before and after browning. Cuticular resistance to water loss decreased with elevation. Sun-exposed shoots lost more water than shaded shoots because of solar heating of needles. Winter desiccation can occur before the decline-related spring browning of red spruce foliage.     

Dependence of winter water relations of mature high-elevation Picea engelmannii and Abies lasiocarpa on summer climate

Water relations of Engelmann spruce (Picea engelmannii Parry) and subalpine fir (Abies lasiocarpa (Hook.) Nutt.) trees growing at an elevation of 3230 m on Mt. Evans, Colorado, USA, were studied during the winters of 1995-1996 and 1996-1997. During both winters, current-year and 1-year-old shoots were collected weekly and their relative water contents (RWC) determined. Measured meteorological parameters were used in a conifer winter water relations model, WINWAT, to simulate changes in shoot RWC of P. engelmannii and A. lasiocarpa during the winter. The model failed to predict shoot RWCs in 1996-1997 when calibrated with 1995-1996 data. The cold early summer of 1995 inhibited xylem formation, which appears to have caused lower rates of water recharge to the needles during the 1995-1996 winter than during the 1996-1997 winter. We conclude that summer climate strongly affects winter water relations in these subalpine species, and that changes in both summer and winter climate must be considered when predicting future ranges of these species.     

Shoot water relations of mature black spruce families displaying a genotype x environment interaction in growth rate. III. Diurnal patterns as influenced by vapor pressure deficit and internal water status

Pressure-volume curves were constructed and shoot water potentials measured for +20-year-old black spruce (Picea mariana (Mill.) BSP) trees from four full-sib families growing on a moist site and a dry site at the Petawawa Research Forest, Ontario, to determine whether differences in diurnal water relations traits were related to productivity. To assess the basis for the observed diurnal patterns, we analyzed effects of environmental and internal water stress variables on diurnal water relations traits. Among the water relations traits examined, turgor pressure was the most sensitive, responding to site, family and environmental variables and displaying the strongest diurnal responses to varying soil water availability and atmospheric vapor pressure deficit (VPD). Overall, there was an 84% drop in turgor pressure with increasing VPD: turgor pressure fell 46% in response to the first 0.75 kPa increase in VPD, and 9.7% in response to a second 0.75 kPa increase in VPD. The families differed in water relations responses to moderate water stress, but not in responses to minor or more extreme water stresses. Thus, at a VPD of 0.5 kPa, there was an estimated 83% greater family difference in turgor pressure on the dry site compared with the moist site. Soil and atmospheric water stress appeared to exert effects in tandem to elicit these responses (r(2) = 0.728). A comparison of the mechanisms of response to water deficit indicated that osmotic adjustment was more important than change in cell wall elasticity. We used a conceptual water relations model to illustrate the differences between tolerant and intolerant families in their mechanisms of water stress response. We conclude that, because genetic responses to site factors are dynamic, the integrated response over time contributes to the observed genetic x environmental interaction in growth.     

Site preparation: Water relations and growth of newly planted jack pine and white spruce

Bareroot jack pine (Pinus banksiana Lamb.) and white spruce (Picea glauca (Moench) Voss) were planted near Elliot Lake, Ontario, on a boreal reforestation site. Site preparation treatments were mixed, mineral and undisturbed (i.e., control) soil. Seedling water relations and growth were examined during the first field season. During the first 28 days after planting, jack pine base (i.e., predawn) and minimum xylem water potential readings were more negative in the control site preparation treatment. White spruce, during the first 10 days, in all site preparation treatments had base and minimum xylem water potential readings more negative than –1.7 MPa. By day 28 base xylem water potentials of white spruce had increased to approximately –1.0 MPa in all site preparation treatments. As the growing season progressed, white spruce minimum xylem water potential readings ceased exceeding the measured turgor loss point first in the mixed followed by the mineral and then control site preparation treatment. Jack pine minimum xylem water potential readings, in all site preparation treatments, almost never exceeded the measured turgor loss point. Water stress and stomatal optimization integrals, day 28 and 125, for both species showed least water stress and greater stomatal optimization in the mixed, mineral and control site preparation treatments, respectively. Both species had less new root growth in the field during the first 28 days after planting compared to seedlings grown for 28 days in a greenhouse for root growth capacity testing. Root growth at 28 days and both shoot and root development at the end of the growing season, were greatest to least in mixed, mineral, and control site preparation treatments, respectively.     

Site-specific water relations and stomatal response of Quercus ilex in a Mediterranean watershed

Intraspecific variations in the water relations and stomatal response of Quercus ilex L. were analyzed under field conditions by comparing trees at two locations within a Mediterranean watershed (l'Avic, Catalonia, NE Spain). Distinct environmental gradients exist between the two sites (referred to as ridge top at 975 m and valley bottom at 700 m) with greater soil depth for water storage, reduced radiation, reduced wind and higher water vapor pressure deficits at the valley bottom than at the ridge top. Osmotic adjustment and changes in tissue elasticiity did not significantly increase drought resistance in the trees studied. The leaf-to-air vapor pressure difference (Deltaw) threshold for inducing stomatal closure was higher at the ridge top (15.6 kPa MPa(-1) +/- 0.5 SE) than at the valley bottom (9.8 kPa MPa(-1) +/- 1.0 SE). However, increases in Deltaw beyond the threshold were followed by greater reductions in leaf conductance of trees at the ridge top than at the valley bottom. At both sites, maximum leaf conductance was related to predawn shoot water potential which, in turn, was related to watershed stream flow. The effects of water deficits during the dry summer of 1989 were more severe in trees at the valley bottom than at the ridge top. During periods of high evaporative demand, site-specific differences in the control of water loss led to more conservative water use by trees at the ridge top and, thus, to even greater drought avoidance (higher predawn water potentials) in late summer.     

Effects of Filtered Air and Misting Treatments on Cuticles of Red Spruce Needles on Whiteface Mountain,N.Y.

Two sets of large (8-20 m tall) red spruce (Picea rubens Sarg.) trees growing on Whiteface Mountain, N.Y., were used in branch chamber experiments. In the summer of 1988, the treatments were made on four healthy trees and consisted of: (1) open branches (N), and in chambers that were supplied with either: (2) ambient air (A); (3) charcoal-filtered air (T); or (4) charcoalfiltered air and deionized water misting when the forest was immersed in ambient cloud water (M). Each tree had all of the treatments. The treatments for the second year (summer of 1989) were made on four declining trees and consisted of: (1) open branches (NC) and chambers supplied with either (2) ambient air and mist (AA), (3) synthetic acid mist and ambient air (SA), (4) distilled water mist and ambient air (DA), (5) synthetic acid mist with filtered air (SF), and (6) distilled water mist with filtered air (DF). Foliar properties measurcd included: (1) microreflectivity, a measure of surface wax properties; (2) cuticle thickness; (3) cutinized cell wall thickness; (4) total cuticular layer thickness: (5) the mass of surface wax extracted from needle samples; and (6) contact angles, a measure of surface wetting. In addition, needles were collected and measured from healthy trees at lower and higher elevations. In the first experiment, reflectance from the wax in the stomata1 antechambers in treatment M was statistically significantly greater than all the other treatments. The cuticles of needles in treatment M were also statistically significantly thicker than the cuticles measured in all the other treatments (40% thicker than for the open branches). A plot of cuticular layer against elevation showed that the two filtered chambers (M and T) provided environments for cuticle development that mimic lower elevations. In 1989 the weather was cooler and more moist, and although the trees used in 1989 were declining, the results were similar to those of 1988. However, the additional treatments with synthetic acid mist resulted in reduced cuticle thickness. While the cuticles of the chamber trees growing at 1,170 m were, except for treatment M and T, generally thicker in 1989 than in 1988, they were still considerably thinner than those of spruce trees growing at the lower elevations. These results suggest regional scale air and precipitation quality have detrimental effects on red spruce needles at Whiteface Mountain.     

Modeling shoot water contents in high-elevation Picea rubens during winter

During the winter of 1990-1991, a meteorological tower was established at an 880-m elevation site within the spruce-fir zone on Mt. Moosilauke, New Hampshire, USA. Hourly means of air, needle and trunk temperatures, wind velocity, relative humidity and solar radiation were recorded. On a weekly basis, shoots that had elongated during the preceding growing season were collected from four red spruce (Picea rubens Sarg.) trees and their relative water contents (RWC) determined. Cuticular resistances of needles from these shoots were measured four times during the winter.Measured meteorological parameters were used in a previously developed model to simulate changes in red spruce shoot RWC during the winter. The modeled results were compared to measured shoot RWCs. The predictive power of the model was improved when it was modified to include measured values of cuticular resistance and needle and trunk temperatures. The new version of the model accurately predicted RWC from late December 1990 to the beginning of April 1991, after which spring recharge appeared to occur. We conclude that water lost from foliage was easily replaced by stored reserves and that uptake of water by the roots was not required to maintain an adequate foliar water content during the winter.     

Drought sensitivity of Norway spruce is higher than that of silver fir along an altitudinal gradient in southwestern Germany

Context

For Central Europe, climate projections foresee an increase in temperature combined with decreasing summer precipitation, resulting in drier conditions during the growing season. This might negatively affect forest growth, especially at sites that are already water-limited, i.e., at low elevation. At higher altitudes trees might profit from increasing temperatures.

Aims

We analyzed variations in radial growth of silver fir (Abies alba Mill.) and Norway spruce (Picea abies (L.) Karst.) along an altitudinal gradient from 400 until 1,140 m a.s.l. in the Black Forest, to assess climate responses with increasing elevation.

Methods

Climate–growth relationships were analyzed retrospectively using tree-ring and climate data. In total, we sampled stem discs of 135 trees to build 27 species- and site-specific chronologies (n _fir?=?13, n _spruce?=?14).

Results

Our results indicate distinct differences in climate–growth relations between fir and spruce along the gradient. Growth of high-altitude fir was positively related to temperature from January till March. Growth of low-altitude fir and spruce at all elevations was positively related to precipitation and negatively to temperature during the growing season, particularly in July. A self-calibrating Palmer drought severity index underlined summer drought sensitivity of these trees.

Conclusion

Overall, we found that climatic control of tree growth changes over altitude for fir. For spruce, a remarkable synchrony in growth variation and climate response was shown, which indicates that this species is drought sensitive at all studied elevations. In a future warmer climate, the growth of low-altitude fir and spruce along the entire studied gradient may be negatively affected in the Black Forest, if an increased evaporative demand cannot be compensated by increased water supply.     

Evidence that longer needle retention of spruce and pine populations at high elevations and high latitudes is largely a phenotypic response

There is abundant evidence that evergreen conifers living at high elevations or at high latitudes have longer-lived needles than trees of the same species living elsewhere. This pattern is likely caused by the influence of low temperature in combination with related factors such as a short growing season and low nutrient availability. Because it is not known to what degree such patterns result from phenotypic versus genotypic variation, we evaluated needle longevity for common-garden-grown lowland populations of European Scots pine (Pinus sylvestris L.) of wide latitudinal origin and Norway spruce (Picea abies L.) of wide elevational origin. Nine-year-old trees of 16 Scots pine populations ranging in origin from 47 degrees to 60 degrees N were studied in Kórnik, Poland (52 degrees N) and 18-year-old trees of 18 Norway spruce populations ranging in origin from 670 to 1235 m elevation in southwestern Poland were studied near Morawina, Poland (51 degrees N, 180 m elevation). There was no tendency in either species for populations from northern or high elevation origins to retain needles longer than other populations. All of the Scots pine populations had between 2.5 to 3.0 needle age cohorts and all of the Norway spruce populations had between 6.4 and 7.2 needle age cohorts. Thus, extended needle retention in Scots pine and Norway spruce populations in low-temperature habitats at high elevations and high latitudes appears to be largely an environmentally regulated phenotypic acclimation.     

Abiotic factors limiting photosynthesis in Abies lasiocarpa and Picea engelmannii seedlings below and above the alpine timberline

Most research on the occurrence and stability of alpine timberlines has focused on correlations between adult tree growth and mean temperatures rather than on specific mechanisms. Timberline migration to higher altitude is dependent on new seedling establishment in the tree-line ecotone; however, reductions in photosynthetic carbon gain in establishing seedlings have previously been interpreted solely in terms of decreased seedling survival. Our objective was to evaluate the impact of abiotic factors (temperature, light and water) on photosynthetic carbon gain in young seedlings of the two dominant conifer tree species occurring naturally above (tree-line ecotone site, TS) and below (forest site, FS) a Rocky Mountain timberline in southeastern Wyoming, USA. Coincidentally, measurements were made during an unusually dry summer. Mean daily photosynthesis in seedlings of both Abies lasiocarpa (Hook.) Nutt. (subalpine fir) and Picea engelmannii Parry ex Engelm. (Engelmann spruce) was less at TS than at FS (19 and 29%, respectively). Minimum nighttime temperatures below 2 degrees C were more frequent at TS than at FS and were associated with reduced maximum photosynthesis the following day. Low midday water potentials were associated with a reduction in carbon gain at both sites early in the season, prior to snowmelt, as well as late in the season when soils began to dry. However, the lower photosynthetic rates at TS than at FS appeared to be unrelated to seedling water status because seedlings at both sites had similar xylem pressure potentials. Solar irradiance was highly variable at both sites as a result of uneven shading by neighboring trees, although this variation was substantially reduced on cloudy days (44% of all days observed). Compared with sunny days, cloudy days resulted in greater integrated daily carbon gain at both sites (41% increase at TS and 69% increase at FS), based on a simulated photosynthesis model. Photosynthetic responses to temperature, sunlight and water suggest that variable solar irradiance and nighttime temperatures were major abiotic factors limiting photosynthetic carbon acquisition in these young seedlings, especially for seedlings growing in the tree-line ecotone.     

Shoot water relations of mature black spruce families displaying a genotype x environment interaction in growth rate. II. Temporal trends and response to varying soil water conditions

Pressure-volume curves and shoot water potentials were determined for black spruce (Picea mariana (Mill.) BSP) trees from four full-sib families at the Petawawa Research Forest, Ontario, Canada. Trees were sampled from a dry site in 1992 and from the dry site and a wet site in 1993. Modulus of elasticity (epsilon), osmotic potential at turgor loss point (Psi(tlp)) and relative water at turgor loss point (RWC(tlp)) all decreased during the growing season. Osmotic potential at saturation (Psi(sat)) and turgor displayed no general temporal trend. Across a range of environmental conditions, Female 59 progeny had equal or lower Psi(sat), and higher or similar epsilon, mean turgor pressure (P(x)) and predawn turgor pressure (P(pd)) compared with Female 63 progeny. Osmotic potential at saturation decreased as water stress increased from mild to moderate and increased as water stress increased from moderate to severe. Stable genetic differences in Psi(sat) were maintained by the same rate of osmotic adjustment from low to moderate water stress. Modulus of elasticity and RWC(tlp) decreased with decreasing water availability, whereas Psi(tlp) showed no response. The combined effects of Psi(sat) and epsilon resulted in no change in P(pd) as water stress increased from low to moderate values, but turgor declined sharply as water stress increased from moderate to high values. We conclude that drought tolerance traits strongly influence the growth of these black spruce families across sites of varying water availability.     

Reflectance of Alaskan black spruce and white spruce foliage in relation to elevation and latitude

Leaf reflectance at visible and near-infrared wavelengths (400-1000 nm) is related primarily to pigmentation, leaf structure and water content, and is an important tool for studying stress physiology and relationships between plants and their growth environment. We studied reflectance of two co-occurring Alaskan conifers, black spruce (Picea mariana (Mill.) BSP) and white spruce (Picea glauca (Moench) Voss), at elevations from 60 to 930 m a.s.l. along a latitudinal gradient from 61 degrees to 68 degrees N. Black spruce samples were collected from 24 sites and white spruce from 30 sites. Overall, reflectance spectra of the two species were similar, but from 400 to 700 nm, needle reflectance was consistently higher in black spruce than in white spruce (all P 相似文献   

The importance of seedling competition in the segregation of spruce and fir in the southern Appalachians

Fraser fir (Abies fraseri [Pursh] Poir.) and red spruce (Picea rubens Sarg.) are codominants of southern Appalachian spruce-fir forests. Fraser fir generally dominates above 1740 m, while red spruce usually dominates below this elevation. This study was designed to determine whether the present segregation of the two species along elevational gradients is associated with seedling competition or contrasting physiological responses to environmental factors. Seedlings were grown for two years in a replacement series experiment along two elevational transects extending from 1300 m to 1900 m, and harvested for growth analysis.Competition increased with decreasing elevation and Fraser fir was apparently the stronger competitor. Mortality was inversely correlated with elevation and was most evident in red spruce in May, and in fir in late summer. Fir growth increased with elevation and red spruce grew most at the middle elevation (1600 m). Neither competitive interactions nor growth responses completely explained the elevational segregation of red spruce and Fraser fir. We hypothesize that the early-successional Fraser fir quickly dominates at higher elevations following disturbance and that continued disturbance will favor its dominance at higher elevations. However, the late-successional noncompetitive red spruce slowly establishes itself and, while consistently present, may only dominate in the absence of disturbance and where other species are at the margins of their distribution.     

Drought resistance of two hybrid Populus clones grown in a large-scale plantation

Poplar hybrids were grown with irrigation in a large-scale plantation to investigate the mechanisms underlying clonal differences in drought resistance. Beginning in spring 1992, Populus trichocarpa x P. deltoides (TD) and P. deltoides x P. nigra (DN) cuttings received 46, 76, or 137 cm year(-1) of irrigation to supplement the 18-20 cm of annual precipitation, and all trees received the same fertilization regime. Stem volume, assessed as the square of stem diameter at breast height times tree height (D(2)H), and water relations of the trees were studied from the end of their second growing season until the end of their fifth growing season. By the end of the second growing season, stem volume of Clone TD was 40-146% larger than that of Clone DN, but stem volume growth was independent of irrigation in excess of 46 cm year(-1) in both clones. During the third growing season, stem volume growth of both clones was limited by both the 46- and 76-cm irrigation treatments, so that by the end of the third growing season trees in the 46-cm irrigation treatment were only half the size of trees in the 137-cm irrigation treatment. These treatment differences were maintained through the fifth growing season. Although stem volumes of Clone TD trees in the 76- and 137-cm irrigation treatments were larger than the corresponding values for Clone DN trees at the end of the third growing season (1994), these clonal differences gradually decreased in subsequent years and were not detectable after 5 years, because stem volume relative growth rate of Clone DN was greater than that of Clone TD in all treatments. Although both clones exhibited similar predawn leaf water potentials, Clone DN typically maintained higher midday leaf water potentials, suggesting better stomatal control of water loss. Clonal and treatment differences in osmotic potential at full turgor were minimal and could not explain the clonal differences in drought resistance. Root density and root density to stem volume ratio increased more in response to moderate drought in Clone DN than in Clone TD, resulting in enhanced drought resistance (high stem volume growth rate under moderate drought conditions) and an increased capacity to withdraw water from the soil. We conclude that the greater drought resistance of Clone DN compared with Clone TD was the result of the maintenance of a more favorable water balance by stomatal regulation and greater carbon allocation to roots during the early stages of drought. However, the low root density to stem volume ratio in Clone DN growing in the 46-cm irrigation treatment suggests that severe water limitation restricted the preferential allocation of carbon to belowground tissues, so that both root and shoot growth were constrained by severe drought.     

Winter water relations of New England conifers and factors influencing their upper elevational limits. I. Measurements

The upper elevational limits of tree species are thought to be controlled by abiotic factors such as temperature and the soil and atmospheric conditions affecting plant water status. We measured relative water contents (RWC), water potentials (Psi) and cuticular conductances (g(c)) of shoots of four conifer species-eastern hemlock (Tsuga canadensis (L.) Carr.), eastern white pine (Pinus strobus L.), red pine (P. resinosa Ait.) and red spruce (Picea rubens Sarg.)-during two winters on Mt. Ascutney, Vermont, USA. Some micrometeorological measurements are also reported. Eastern hemlock and white pine were studied near their upper elevational limits at a 640-m site, and red pine was studied near its upper elevational limit at 715 m. Red spruce was also studied at the 715-m site, which is in the middle of its elevational range on this mountain. There was no evidence of winter desiccation stress in any species. The observed distribution of seedlings suggested that the upper elevational limits on shade-intolerant eastern white pine and red pine are set by the absence of suitable seed beds after 100 years without fire. Eastern hemlock is able to reproduce in deep shade on organic substrates, but germination at high elevations may be restricted by low temperatures.