End of season carbon supply status of woody species near the treeline in western China

As trees and shrubs approach the high elevation tree limit, it is often assumed that they fall short in photosynthate (source limitation). Alternatively, low temperature may restrict carbon investment (growth, sink limitation). The content of mobile non-structural carbohydrates (NSC) in tissues is considered a measure of the carbon source–sink balance. To test the source vs. sink limitation hypothesis, we compared late-season NSC concentrations of various woody taxa across altitudinal gradients from the subalpine forest to the treeline at the eastern edge of the Tibetan Plateau. Since we were interested in the generality of trends, we present “community” trends across four taxa, namely Quercus aquifolioides, Abies faxoniana, Rhododendron fabri subsp. prattii and Sorbus rufopilosa. NSC concentrations increased significantly with altitude in branch wood, current-year and last-year leaves, while there were no significant trends in stem sapwood and root xylem. The sugar to starch ratio was roughly 1:1 in branches and evergreen leaves, while stems and roots showed a higher starch fraction. Analyses of total nitrogen in leaves and wood tissues indicated no change in the trees’ nitrogen supply with elevation. The overall altitudinal trends of NSC in this group of woody plant species revealed no depletion of carbon reserves near the tree limit, suggesting that sink limitation predominates woody plant life across this treeline ecotone community.     

Carbon sink limitation and frost tolerance control performance of the tree <Emphasis Type="Italic">Kageneckia angustifolia</Emphasis> D. Don (Rosaceae) at the treeline in central Chile

The decrease in temperature with increasing elevation may determine the altitudinal tree distribution in different ways: affecting survival through freezing temperatures, by a negative carbon balance produced by lower photosynthetic rates, or by limiting growth activity. Here we assessed the relative importance of these direct and indirect effects of altitudinal decrease in temperature in determining the treeline in central Chile (33°S) dominated by Kageneckia angustifolia. We selected two altitudes (2000 and 2200 m a.s.l.) along the treeline ecotone. At each elevation, leaf non-structural carbohydrates (NSC) and gas exchange parameters were measured on five individuals during the growing season. We also determined the cold resistance of K.␣angustifolia, by measuring temperatures that cause 50% seedling mortality (LT₅₀) and ice nucleation (IN). No differences in net photosynthesis were found between altitudes. Although no differences were detected on NSC concentration on a dry matter basis between 2000 and 2200 m, when NSC concentration was expressed on a leaf area basis, higher contents were found at the higher elevation. Thus, carbon sink limitations may occur at the K. angustifolia’s upper altitudinal limit. For seedlings derived from seeds collected at the 2200 m, LT₅₀ of cold-acclimated and non-acclimated plants were −9.5 and −7 °C, respectively. However, temperatures as low as −10 °C can frequently occur at this altitude during the end of winter. Therefore, low temperature injury of seedlings seems also be involved in the treeline formation in this species. Hence, a confluence of global (carbon sink limitation) and regional (freezing tolerance) mechanisms explains the treeline formation in the Mediterranean-type climate zone of central Chile.     

Effects of diffusion limitation on immobilized nitrifying microorganisms at low temperatures

Activation energies of suspended and immobilized nitrifying bacteria were determined and compared to determine if diffusion limitation results in decreased sensitivity for temperature. The activation energy for the respiration activity of suspended Nitrosomonas europaea and Nitrobacter agilis was found to be 86.4 and 58.4 kJ mol(-1), respectively. The activation energy for oxygen diffusion in the support material, kappa-carrageenan, determined from the effect of temperature on the effective diffusion coefficient (D), was 17.2 kJ mol(-1). Consequently, the apparent actvation energy of diffusion limited cells should be lower. It was indeed shown that due to the effect of diffusion limitation and to temperature effects on the Monod constant K(s), the immobilized-cell activity was less sensitive to temperature. The apparent activation energy for immobilized Ns. europaea was between 28.6 and 94.2 kJ mol(-1) and for immobilized Nb. agilis between 1.4 and 72.9 kJ mol(-1), depending on the oxygen concentration and temperature. (c) 1995 John Wiley & Sons, Inc.     

土壤水分状况及环境条件对水稻蒸腾的影响

在南方红壤区用田间测坑试验,研究了高、中、低3种土壤水分条件下的早、晚稻蒸腾速率和水分利用效率的变化,蒸腾日变化结果表明,蒸腾速率受许多田间小气候因子的影响,相关分析及多元逐步回归分析表明,叶片与空气相对湿度差和叶面温度对水稻叶片蒸腾影响最大,处理B（中等土壤水分条件）可以明显提高叶片水利用率和产是,处理C提高了叶片水分利用效率,却导致晚稻减产,早稻中、低土壤水分条件可以减小叶片蒸腾速率,而晚稻上并不能明显减少叶片蒸腾。     

Nitrogen status of conifer needles at the alpine treeline

Background and Aims: High elevation treelines occur worldwide at similar mean growing season temperatures. Does this result from direct impact of low temperature on growth or carbon metabolism, or does nutrient limitation, induced by low soil temperature, play a role? Similar treeline elevations at contrasting soil fertility argue against the latter, but the actual nutritional status of treeline trees (here addressed as foliage nitrogen concentration) has never been assessed systematically. Although needle nitrogen (N) concentration does not necessarily indicate growth limitation by N, the relative abundance of N would indicate obvious depletions at the treeline.

Methods: A central problem with any foliage nutrient assay is that the units for describing the element concentration are dependent on elevation themselves. Here we separate changes in N per unit tissue from changes in reference units.

Results: Needles of Pinus cembra and Picea abies in the Alps do not show elevational differences in N concentration per dry weight, water content, area or volume, thus, there is no N depletion near the elevational tree limit. Hence, nutrient supply is either unaltered, or growth is adjusted so that nutrient depletion in needle tissue does not occur.

Conclusions: Chronic N shortage at needle level is not an explanation for low tree vigour at the treeline.     

Fine-scale distribution of treeline trees and the nurse plant facilitation on the eastern Tibetan Plateau

Above-average climate warming occurred during the 20th century in high altitude regions, and alpine treelines are believed to be an early indicator to respond to these warming-related changes. However, empirical investigations on treeline dynamics showed diverse results. The main objectives of this study are: (1) to investigate if treeline position shifted and if tree recruitment changed along with climate warming, and (2) to test if adult trees have “nursing effect” on tree establishment at treelines. We investigated two Balfour spruce (Picea balfouriana Rehd. et Wils.) treelines in Chang Niang (CNT) and Dang Dui (DDT), Dingqing county, Changdu prefecture, eastern Tibet. At each treeline site, three replicate plots with a size 30 m × 50 m were established. The coordinates of each tree within the plots were recorded and the age of each tree was identified by dendrochronological method. The changes in treeline position and tree recruitment were examined from spatially fine-scale distribution of trees and their age structure. The spatial patterns of individual trees were analyzed to infer the neighborhood effects. Results indicate that plots CNT2, CNT3, DDT1 and DDT2 showed stable treeline position during the last century, whereas plots CNT1 and DDT3 showed treeline advancing movement. Tree recruitments in all the six plots were enhanced during the 20th century, with two peaks occurring in the 1890–1910s and the 1950–1990s. Seedlings and saplings showed a general clustered distribution in all the six plots. The diverse pattern of treeline movement and episodic regeneration suggest that the treeline activity is not merely a result of climate change. “Nursing effects” from adult trees may play an important role in shaping the treeline activities on the eastern Tibetan Plateau. Our findings reveal diverse patterns in treeline dynamics at a local scale and highlight the importance of incorporating biotic interactions into species distribution modeling approaches.     

Effects of early resource limitation and compensatory growth on lifetime fitness in the ladybird beetle (Harmonia axyridis)

Acceleration of growth following a period of diet restriction may result in either complete or partial catch-up in size. The existence of such compensatory growth indicates that organisms commonly grow at rates below their physiological maxima and this implies a cost for accelerated growth. We examined patterns of accelerated growth in response to temporary resource limitation, and assayed both short and long-term costs of this growth in the ladybird beetle Harmonia axyridis. Subsequent to the period of food restriction, accelerated growth resulted in complete compensation for body sizes, although we observed greater larval mortality during the period of compensation. There were no effects on female fecundity or survivorship within 3 months of maturation. Females did not discriminate against males that had undergone compensatory growth, nor did we observe effects on male mating behaviour. However, individuals that underwent compensatory growth died significantly sooner when deprived of food late in adult life, suggesting that longer-term costs of compensatory growth may be quite mild and detectable only under stressful conditions.     

Atmospheric element deposit on tree barks: The opposite effects of rain and transpiration

The elemental composition of the deposit formed on the bark of ash-trees was studied over several months on stems ranging from 3 months to 10 years. For this purpose (1) the total elemental composition of the deposit, (2) the structure and composition of the solid particles and (3) the deposit dry weight per dm² were studied. Concurrently the part of this superficial deposit washed out by rain during 3.5 months was sampled at each rain event and its elemental composition analyzed. This study shows that the deposit was submitted to an intense turnover, with an average leaching-out flux reaching approximately 27 mg dm⁻² month⁻¹ and a very low increase of the deposit weight per dm² during the first five years and almost null afterwards. The origin of this superficial deposit was investigated. The main part (78 ± 10%) was organic matter originating from the atmospheric deposition or from the tree. The inorganic content originated partly from the atmosphere, with geogenic and anthropogenic particles, and also from the tree, in which it was demonstrated by scanning electron microscopy - energy dispersive X-ray that a non-negligible part was obtained from bark transpiration, inducing a superficial deposit, mainly of Ca or K in the area surrounding the lenticels. All those results demonstrate that this bark superficial deposit is a complex matrix which must not be considered as a simple cumulative archive but which seems to be the source of interesting information targeting mostly recent atmospheric pollution pressures, when compared to the bulk of suber integrated particles.     

Nutrient limitation of phytoplankton growth in Waquoit Bay, Massachusetts, USA: a nutrient enrichment study

We conducted nutrient enrichment experiments and field sampling to address three questions: (1) is there nutrient limitation of phytoplankton accumulation within an estuary whose waters are exposed to relatively high nitrogen loading rates, (2) where in the salinity gradient from fresh to seawater (0 to 32‰) is there a shift from phosphorus to nitrogen limitation of phytoplankton accumulation, and (3) is there a seasonal shift in limiting function of phosphorus and nitrogen anywhere in the estuarine gradient. Nitrogen and phosphorus enrichment experiments in the Childs River, an estuary of Waquoit Bay, Massachusetts, USA, showed that the accumulation of phytoplankton biomass in brackish and saline water was limited by supply of nitrate during warm months. The effects of enrichment were less evident in fresh water, with short-lived responses to phosphate enrichment. There was no specific point along the salinity gradient where there was a shift from phosphorus- to nitrogen-limited phytoplankton accumulation; rather, the relative importance of nitrogen and phosphorus changed along the salinity gradient in the estuary and with season of the year. There was no response to nutrient additions during the colder months, suggesting that some seasonally-varying factor, such as light, temperature or a physiological mechanism, restricted phytoplankton accumulation during months other than May-Aug. There was only slight evidence of a seasonal shift between nitrogen- and phosphorus-limitation of chlorophyll accumulation. Phytoplankton populations in nutrient-rich estuaries with short flushing times grow fast, but at the same time the cells may be advected out of the estuaries while still rapidly dividing, thereby providing an important subsidy to production in nearby deeper waters. This revised version was published online in August 2006 with corrections to the Cover Date.     

Below-ground bud banks increase along a precipitation gradient of the North American Great Plains: a test of the meristem limitation hypothesis

In perennial grasslands, the below-ground population of meristems (bud bank) plays a fundamental role in plant population dynamics. Here, we tested the 'meristem limitation hypothesis' prediction - that bud banks increase along an increasing precipitation/productivity gradient in North American grasslands - and assessed the seasonal dynamics of bud banks. We sampled bud and stem populations quarterly at six sites across a 1100 km gradient in central North America. Bud banks increased with average annual precipitation, which explained 80% of the variability between the sites. In addition, seasonal changes in grass bud banks were surprisingly similar across a 2.5-fold range in precipitation and a 4-fold range of productivity: densities peaked in March, decreased in June and increased slightly in September. Increasing meristem limitation may constrain vegetation responses to inter-annual changes in resources. An important consequence of this is that biomes with large bud banks may be the most responsive to environmental change. If meristem limitation represents an important constraint on productivity responses to environmental variability, then bud banks must be considered in developing predictive models for grassland responses to environmental change.     

Phytoplankton growth in the Ohio, Cumberland and Tennessee Rivers, USA: inter-site differences in light and nutrient limitation

Seasonal patterns in resource limitation of phytoplankton growth were assessed monthly within three large rivers with differing extents of water regulation. The Ohio River is regulated by low dams that do not substantially modify discharge, while the Cumberland and Tennessee Rivers are impounded by a series of high dams to enhance water storage for downstream flood control. Laboratory dilution assays with light and nutrient manipulations indicated that light was the main factor limiting phytoplankton growth at irradiances below 7 E m^–2 d^–1. Light limited growth was frequent in the turbid, higher discharge of the Ohio River, but was rare in the heavily regulated Tennessee and Cumberland Rivers. When irradiance exceeded 7 E m^–2 d^–1, phytoplankton were either P-limited (Cumberland River), co-limited by P and N (Tennessee River), or Si limited (Ohio River). Site-specific differences in nutrient limitation were consistent with differences in ambient nutrient levels, with the Tennessee and Cumberland Rivers characterized by lower N and P concentrations, and the Ohio River by lower Si. Downstream nutrient depletion was evident in the Ohio River through comparison of an upstream and a downstream site, with nutrient limitation (Si) occurring more frequently downstream. Phytoplankton growth rates at ambient light and nutrient levels ranged from 0.1 to 1.5 d^–1 in the Ohio River and 0.2 to 0.6 d^–1 in the Tennessee and Cumberland Rivers. Growth rates were greatest at the onset of the summer base pool, as light intensities increased and nutrient levels were maximal. Our findings indicate that multiple factors regulate phytoplankton growth in regulated rivers and that spatial complexity may arise from differences in discharge and water aging.     

A 10-year demographic study of rabbitbrush (Chrysothamnus nauseosus): growth,survival and water limitation

A population of 126 adult rabbitbrush (Chrysothamnus nauseosus var. consimilis) was followed for 10 consecutive years from 1983 through 1992 in a sanddune ecosytem on the north shore of Mono Lake, California. The study area received an average (1981–1992) of 160 mm total annual precipitation of which 55 mm fell from April through October, making this site drier than other known sites in the Great Basin desert where C. nauseosus occurs. Cover of all shrubs was 18.0%, and that of C. nauseosus 7.5%, in the 3077 m² study plot. Water was limiting for this population of C. nauseosus, exerting large effects on new canopy growth and reproductive effort in a given year, on survival, and on establishment of juveniles into the adult population. Growth and survivorship of the adult shrubs were significantly correlated with the depth to ground water, which varied from 3.6 to 6.0 m below the surface during the study, and not with any measure of precipitation in a given year or adjacent years. Shrubs in this population exhibited a net decrease in size between 1983 and 1992. The region experienced a significant drought from 1987 onwards, with some increase in precipitation after 1990. Mortality of shrubs began in 1987 and reached 22% by 1992. Individuals that died were smaller on average initially than individuals surviving throughout the study: 50% or more of the individuals having 4000 cm² of canopy or less at the start of the study died by 1992. Elevation (m msl) of each shrub in the study area, presumably correlated with depth to ground water, had no effect on growth but a weak effect on survival during the study period. Specific microsite had significant effects, with shrubs on steep slopes being smaller, having more negative growth slopes, and being more likely to die during the drought compared to individuals occurring on flatter gradients. Recruiment of juveniles into the population did not occur after 1985. Competition between shrubs could not be definitively demonstrated during the study, but only indirect tests for competition were used. Adult C. nauseosus and adults of all species combined (Sarcobatus vermiculatus with 9.6% cover, Tetradymia tetrameres with 0.8% cover and Chrysothamnus viscidiflorus with 0.02% cover) were randomly distributed, failing to support the hypothesis of competition if competition for water results in a uniform distribution of shrubs. Moreover, comparison of the spatial dispersion of C. nauseosus individuals before (1985) and after (1992) mortality did not reveal competitive thinning toward uniformity. On the other hand, larger individuals were farther from their nearest neighbor than were smaller individuals, an effect postulated to result from competitive thinning.     

Temperature and humidity effects on branchlet gas-exchange in white spruce: an explanation for the increase in transpiration with branchlet temperature

In situ gas-exchange data, for branchlets of white spruce [Picea glauca (Moench) Voss.] in a mature mixed-wood boreal forest in central Canada (53°44′N 105°14′W), were subjected to a multiple regression analysis. Vapor pressure deficit (VPD) and branchlet temperature (t_leaf) were both significant predictors (P<0.0001) of stomatal conductance to water vapor (g_sw) and net photosynthesis (A_n), together explaining 67 and 64% of the variation in g_sw and A_n, respectively. Since VPD and t_leaf were autocorrelated in these field data, but also to further explore the nature of independent effects of temperature and humidity on water and CO₂ exchange in white spruce, steady-state gas-exchange was performed on well-watered greenhouse-grown seedlings of white spruce. Results from laboratory experiments supported the following conclusions: (1) Transpiration (E) increases with VPD to an inflection point that increases linearly with t_leaf. This t_leaf effect on E could not be explained by trends in VPD, RH, A_n or PFD. Rather, our data support a model in which E and g_sw are influenced by the balance between ’supply’ and ’loss’ of water to and from leaf tissue, respectively. The supply of water appears to be in accordance with Darcy’s law, where supply of water is proportional to the driving gradient in pressure/ tension, specific permeability (k), and inverse of water viscosity (n ^–1). Approximately half of the increase in E could be explained by the linear increase in n ^–1 with increasing t_leaf. We propose that increases in k explain the remainder of the increase in E with t_leaf. (2) VPD and t_leaf appear to have independent effects on g_sw. In contrast, RH effects on g_sw or E were subtle and could be explained by a combination of effects of t_leaf and VPD. (3) A_n was affected primarily by t_leaf, being reduced at low (10°C) and high (40°C) temperatures, and only indirectly by humidity parameters via stomatal conductance, viz. intercellular CO₂ concentrations. Our results have implications for the prediction of water fluxes from plants and canopies in areas where plant temperatures vary diurnally or seasonally. Received: 24 September 1998 / Accepted: 20 July 1999     

Carbohydrate partitioning, photosynthesis and growth in Lemna gibba G3. II. Effects of phosphorus limitation

The relationship between CO₂ assimilation rate, growth and partitioning of carbon among starch, sucrose, glucose and fructose were studied in phosphorus (P_i)-limited Lemna gibba L. G3. Two experimental models were used: 1) Cultures were grown at various stable, suboptimal rates regulated by the supply of P_i; 2) cultures growing at optimal rates were transferred to P_i-free medium. The response to a P_i deficiency can be divided into two phases. Phase I is characterized by hyperactivity of the sucrose synthesis pathway, leading to high levels of glucose and fructose. Phase II is characterized by starch accumulation associated with a decrease in the cytoplasmic pools of soluble sugars owing to inhibition of carbon export from the chloroplast. A strong negative correlation was found between the CO₂ assimilation rate and starch levels. No significant correlation was found between assimilation and ATP levels and decrease in relative growth rate did not significantly affect the adenylate energy charge (EC). The regulatory aspects of the partitioning of carbon among soluble sugars and starch as well as the negative correlation between carbohydrate levels and CO₂ assimilation at P_i-limited growth are discussed.     

EarlyZea cultivation in Honduras: Implications for the iltis hypothesis

Heavy deposits of charcoal and associated Zea pollen recovered from the Aguada Petapilla in the Copán Valley of western Honduras strongly support Hugh lltis’s hypothesis concerning the early domestication and use of maize. Evidence of burning and cultivation appear at approximately 2600–2700 b.c., predating the earliest direct archaeological evidence for human occupation of the region by more than 1,000 years. These Copán data are in close agreement with two other sediment cores from Honduras, and also with maize macrofossils recently recovered from dry deposits in El Gigante Cave.     

Effects of light and microcrustacean prey on growth and investment in carnivory in Utricularia vulgaris

SUMMARY 1. In a 5-week enclosure experiment, we studied the effects of light (ambient light, low light) and prey availability (no prey, prey added) on growth and investment in carnivory in Utricularia vulgaris .
2. Investment in carnivory, measured as the proportion of biomass allocated to bladders, was strongly affected by our manipulations of light intensity and prey density. In the treatment with high prey density the light reduction decreased the investment in bladders from 25% to zero. The effect of prey density on investment in bladders was negative. Because prey addition increased the concentration of nutrients, especially phosphorus, we propose that the effect of the prey treatment on investment reflected altered nutrient concentrations.
3. Availability of prey increased growth and apical biomass of Utricularia . As Utricularia had very few bladders in some treatments we suggest that the effect was due to a combination of live prey trapped and increased nutrient availability from dead prey.
4. Abundance of periphyton on Utricularia and on the enclosure walls was highest in the treatments with high prey density where nutrient concentrations were highest. Thus we interpret the response of periphyton as primarily reflecting nutrient availability.     

Environmental and physiological regulation of transpiration in tropical forest gap species: the influence of boundary layer and hydraulic properties

Environmental and physiological regulation of transpiration were examined in several gap-colonizing shrub and tree species during two consecutive dry seasons in a moist, lowland tropical forest on Barro Colorado Island, Panama. Whole plant transpiration, stomatal and total vapor phase (stomatal + boundary layer) conductance, plant water potential and environmental variables were measured concurrently. This allowed control of transpiration (E) to be partitioned quantitatively between stomatal (g _s) and boundary layer (g _b) conductance and permitted the impact of invividual environmental and physiological variables on stomatal behavior and E to be assessed. Wind speed in treefall gap sites was often below the 0.25 m s^–1 stalling speed of the anemometer used and was rarely above 0.5 m s^–1, resulting in uniformly low g _b (c. 200–300 mmol m^–2 s^–1) among all species studied regardless of leaf size. Stomatal conductance was typically equal to or somewhat greater than g _b. This strongly decoupled E from control by stomata, so that in Miconia argentea a 10% change in g _s when g _s was near its mean value was predicted to yield only a 2.5% change in E. Porometric estimates of E, obtained as the product of g _s and the leaf-bulk air vapor pressure difference (VPD) without taking g _b into account, were up to 300% higher than actual E determined from sap flow measurements. Porometry was thus inadequate as a means of assessing the physiological consequences of stomatal behavior in different gap colonizing species. Stomatal responses to humidity strongly limited the increase in E with increasing evaporative demand. Stomata of all species studied appeared to respond to increasing evaporative demand in the same manner when the leaf surface was selected as the reference point for determination of external vapor pressure and when simultaneous variation of light and leaf-air VPD was taken into account. This result suggests that contrasting stomatal responses to similar leaf-bulk air VPD may be governed as much by the external boundary layer as by intrinsic physiological differences among species. Both E and g _s initially increased sharply with increasing leaf area-specific total hydraulic conductance of the soil/root/leaf pathway (G _t), becoming asymptotic at higher values of G _t. For both E and g _s a unique relationship appeared to describe the response of all species to variations in G _t. The relatively weak correlation observed between g _s and midday leaf water potential suggested that stomatal adjustment to variations in water availability coordinated E with water transport efficiency rather than bulk leaf water status.