Influence of Human Pressure on Forest Resources and Productivity at Stand and Tree Scales: The Case Study of Yunnan Pine in SW China.

This paper examines human impact on stands and individual trees of Pinus yunnanensis growing near the small mountain villages of Pianshui and Yangjuan in southwestern Sichuan Province, China. In an effort to assess whether use of these forests was sustainable, we examined the effects of human use in two ways. First, we directly measured the effect of cutting branches, for fuel and fodder, on tree growth. We hypothesized that branch cutting would negatively impact tree growth. We established 12 plots on four hills and compared 14 pairs of trees, one tree in each pair with an apparently full crown and the other with a considerable portion of the crown removed. Second, we assessed stand and tree properties over a 500 m elevation gradient above the villages where we hypothesized that as elevation increases, stand and tree properties should show fewer human impacts. Although extensive branch cutting reduced the live crown, tree height and diameter, compensatory processes likely enabled trees to recover and to add basal area increments (BAIs) similar to those added by trees with full crowns. Trees and stands close to villages showed less growth and lower basal areas, respectively, than stands and trees at intermediate or distant elevations from villages. Areas relatively close to the villages showed considerable effects of human-related disturbances such as branch cutting, grazing, tree and shrub removal, losses of litter, and human and animal trails. Such areas had increased soil erosion and often loss of the 'A' horizon. Stands close to villages had younger trees, lower stand basal areas, smaller basal area increments, and more stumps. Our results suggest an increasingly vulnerable interface between occupants of these two villages and their surrounding forests.

Is the Returning Farmland to Forest Program a Success? Three Case Studies from Sichuan.

China's tuigeng huanlin or "Returning Farmland to Forest" (RFFP) program has been widely praised as the world's largest and most successful payment for ecosystem services program, as well as a major contributor to China's dramatic increase in forest cover from perhaps as low as 8% in 1960 to about 21% today. By compensating rural households for the conversion of marginal farmland to forestland and financing the afforestation of barren mountainsides, the program, in addition to expanding forestland, aims to reduce soil erosion and alleviate poverty. This paper presents qualitative and quantitative studies conducted on the local implementation of RFFP in three diverse townships in Sichuan. We find the actual results to be more mixed than the official figures would indicate. Though there have been some positive results, we identify problems with site and species selection, compensation for land taken out of cultivation, shift of labor to off-farm activities, and monitoring of replanted sites, which challenge the ecological and economic impacts of these programs and reveal much of the effort of the program has been misdirected. We suggest that efforts are misplaced because of the top-down, panacea nature of the program, which in turn is a feature of Chinese bureaucratic management.

Defining how aging Pseudotsuga and Abies compensate for multiple stresses through multi-criteria assessment of a functional-structural model.

Many hypotheses have been advanced about factors that control tree longevity. We use a simulation model with multi-criteria optimization and Pareto optimality to determine branch morphologies in the Pinaceae that minimize the effect of growth limitations due to water stress while simultaneously maximizing carbohydrate gain. Two distinct branch morphologies in the Pareto optimal space resemble Pseudotsuga menziesii (Mirb.) Franco and Abies grandis (Dougl. ex D. Don) Lindl., respectively. These morphologies are distinguished by their performance with respect to two pathways of compensation for hydraulic limitation: minimizing the mean path length to terminal foliage (Pseudotsuga) and minimizing the mean number of junction constrictions to terminal foliage (Abies). Within these two groups, we find trade-offs between the criteria for foliage display and the criteria for hydraulic functioning, which shows that an appropriate framework for considering tree longevity is how trees compensate, simultaneously, for multiple stresses. The diverse morphologies that are found in a typical old-growth conifer forest may achieve compensation in different ways. The method of Pareto optimization that we employ preserves all solutions that are successful in achieving different combinations of criteria. The model for branch development that we use simulates the process of delayed adaptive reiteration (DAR), whereby new foliage grows from suppressed buds within the established branch structure. We propose a theoretical synthesis for the role of morphology in the persistence of old Pseudotsuga based on the characteristics of branch morphogenesis found in branches simulated from the optimal set. (i) The primary constraint on branch growth for Pseudotsuga is the mean path length; (ii) as has been previously noted, DAR is an opportunistic architecture; and (iii) DAR is limited by the number of successive reiterations that can form. We show that Pseudotsuga morphology is not the only solution to old-growth constraints, and we suggest how the model results should be used to guide future empirical investigation based on the two contrasting morphologies and how the morphological contrast may relate to physiological processes. Our results show that multi-criteria optimization with Pareto optimality has promise to advance the use of models in theory development and in exploration of functional-structural trade-offs, particularly in complex biological systems with multiple limiting factors.

Actual and potential transpiration and carbon assimilation in an irrigated poplar plantation.

We examined the tradeoffs between stand-level water use and carbon uptake that result when biomass production of trees in plantations is maximized by removing nutrient and water limitations. A Populus trichocarpa Torr. x P. deltoides Bartr. & Marsh. plantation was irrigated and received frequent additions of nutrients to optimize biomass production. Sap flux density was measured continuously over four of the six growing-season months, supplemented with periodic measurements of leaf gas exchange and water potential. Measurements of tree diameter and height were used to estimate leaf area and biomass production based on allometric relationships. Sap flux was converted to canopy conductance and analyzed with an empirical model to isolate the effects of water limitation. Actual and soil-water-unlimited potential CO(2) uptakes were estimated with a canopy conductance constrained carbon assimilation (4C-A) scheme, which couples actual or potential canopy conductance with vertical gradients of light distribution, leaf-level conductance, maximum Rubisco capacity and maximum electron transport. Net primary production (NPP) was about 43% of gross primary production (GPP); when estimated for individual trees, this ratio was independent of tree size. Based on the NPP/GPP ratio, we found that current irrigation reduced growth by about 18% compared with growth with no water limitation. To achieve maximum growth, however, would require 70% more water for transpiration, and would reduce water-use efficiency by 27%, from 1.57 to 1.15 g stem wood C kg(-1) water. Given the economic and social values of water, plantation managers appear to have optimized water use.

Tree water storage and its diurnal dynamics related to sap flow and changes in stem volume in old-growth Douglas-fir trees.

Diurnal and seasonal tree water storage was studied in three large Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco) trees at the Wind River Canopy Crane Research site. Changes in water storage were based on measurements of sap flow and changes in stem volume and tissue water content at different heights in the stem and branches. We measured sap flow by two variants of the heat balance method (with internal heating in stems and external heating in branches), stem volume with electronic dendrometers, and tissue water content gravimetrically. Water storage was calculated from the differences in diurnal courses of sap flow at different heights and their integration. Old-growth Douglas-fir trees contained large amounts of free water: stem sapwood was the most important storage site, followed by stem phloem, branch sapwood, branch phloem and needles. There were significant time shifts (minutes to hours) between sap flow measured at different positions within the transport system (i.e., stem base to shoot tip), suggesting a highly elastic transport system. On selected fine days between late July and early October, when daily transpiration ranged from 150 to 300 liters, the quantity of stored water used daily ranged from 25 to 55 liters, i.e., about 20% of daily total sap flow. The greatest amount of this stored water came from the lower stem; however, proportionally more water was removed from the upper parts of the tree relative to their water storage capacity. In addition to lags in sap flow from one point in the hydrolic pathway to another, the withdrawal and replacement of stored water was reflected in changes in stem volume. When point-to-point lags in sap flow (minutes to hours near the top and stem base, respectively) were considered, there was a strong linear relationship between stem volume changes and transpiration. Volume changes of the whole tree were small (equivalent to 14% of the total daily use of stored water) indicating that most stored water came from the stem and from its inelastic (sapwood) tissues. Whole tree transpiration can be maintained with stored water for about a week, but it can be maintained with stored water from the upper crown alone for no more than a few hours.

Variation in specific needle area of old-growth Douglas-fir in relation to needle age, within-crown position and epicormic shoot production.

Variation in specific needle area (SNA; cm2 projected fresh needle area g-1 oven-dried needle weight) was investigated in relation to needle age, within-crown position and epicormic shoot production in 450-year-old Douglas-fir (Pseudotsuga menziesii Mirb. (Franco) var. menziesii) trees. Specific needle area decreased with increasing needle age. The magnitude and rate of change in SNA with needle age were greatest for lower-crown branches, and decreased toward the middle- and upper-crown branches. For all branches, there was no difference between regular and epicormic shoots in the relationship between SNA and needle age. Specific needle area decreased with increasing distance from branch base, and this relationship was significant for the majority of needle age classes of the upper- and middle-crown branches. In the lowercrown branches, SNA did not vary with distance from branch base for the majority of needle age classes. For all branches, there was no difference between regular and epicormic shoots in the relationship between SNA and distance from branch base for the majority of needle age classes. These results indicate that renewal of foliage by epicormic shoot production maintains needle quality. Branch SNA increased linearly with decreasing height in the crown at a mean rate of 0.951 +/- 0.110 cm2 g-1 per vertical meter. Total needle area of branches was estimated from total needle dry weight taking into account within-branch variation in SNA. Analyses of allometric relationships between branch size and foliage amount (needle area and needle dry weight) showed that branch length was a better predictor of foliage amount than branch diameter for old Douglas-fir trees. Total needle dry weight and needle area of the sample trees, estimated from branch length and branch height and taking into account vertical within-crown variation in branch SNA, ranged from 42.4 to 154.2 kg and from 246.2 to 816.0 m2 per tree, respectively.

Boundary layer conductance, leaf temperature and transpiration of Abies amabilis branches.

We used three methods to measure boundary layer conductance to heat transfer (g(bH)) and water vapor transfer (g(bV)) in foliated branches of Abies amabilis Dougl. ex J. Forbes, a subalpine forest tree that produces clumped shoot morphology on sun-formed branches. Boundary layer conductances estimated in the field from energy balance measurements increased linearly from approximately 10 mm s(-1) at low wind speeds (< 0.1 m s(-1)) to over 150 mm s(-1) at wind speeds of 2.0 m s(-1). Boundary layer conductances measured on shoot models in a wind tunnel were consistently higher than field measurements. The difference between wind tunnel values and field measurements was attributable to variation in path length between the two experimental environments. Boundary layer conductance estimated by subtracting stomatal resistance (r(sV)) measured with a porometer from the total branch vapor phase resistance were unusually small. Sensitivity analysis demonstrated that this method is not suitable for coniferous foliage or when stomatal conductance (g(sV)) is small compared with g(bV). Analysis of the relative magnitudes of g(sV) and g(bV) revealed that, under most conditions, A. amabilis branches are well coupled (i.e., g(sV) is the dominant controller of transpiration). The boundary layer conductance to heat transfer is small enough that leaf temperature can become substantially higher than air temperature when radiation is high and wind speed is low. Over a two-month period, the maximum difference between leaf and air temperatures exceeded 6 degrees C. Leaf temperature exceeded air temperature by more than 2 degrees C on 10% of the daylight hours during this period. Consideration of both the photosynthetic temperature response of A. amabilis foliage as well as the summer air temperature conditions in its habitat suggests that these elevated leaf temperatures do not have a significant impact on carbon gain during the growing season.

The effects of light acclimation during and after foliage expansion on photosynthesis ofAbies amabilis foliage within the canopy.

Variation in the photosynthetic function ofAbies amabilis foliage within a canopy was examined and related to three different processes that affect foliage function: foliage aging, sun-shade acclimation that occurred while foliage was expanding, and reacclimation after expansion was complete. Foliage produced in the sun had higher photosynthesis at light saturation (A max, µmol·m-2·s-1), dark respiration (µmol·m-2·s-1), nitrogen content (g·m-2), chlorophyll content (g·m-2), and chlorophylla:b ratio, and a lower chlorophyll to nitrogen ratio (chl:N), than foliage produced in the shade. As sun foliage becomes shaded, it becomes physiologically similar to shade foliage, even though it still retains a sun morphology. Shaded sun foliage exhibited lowerA max, dark respiration, nitrogen content, and chlorophylla:b ratio, and a higher chl:N ratio than sun foliage of the same age remaining in the open. However, shaded sun foliage had a higher chlorophyll content than sun foliage remaining in the open, even though true shade foliage had a lower chlorophyll content than sun foliage. This anomaly arises because as sun foliage becomes shaded, it retains a higher nitrogen content than shade foliage in a similar light environment, but the two forms have similar chl:N ratios. Within the canopy, most physiological indicators were more strongly correlated with the current light environment than with foliage age or leaf thickness, with the exception of chlorophyll content.A max decreased significantly with both decreasing current light environment of the foliage and increasing foliage age. The same trend with current light and age was found for the chlorophylla:b ratio. Foliage nitrogen content also decreased with a decrease in current light environment, but no distinct pattern was found with foliage age. Leaf thickness was also important for predicting leaf nitrogen content: thicker leaves had more nitrogen than thinner leaves regardless of light environment or age. The chl:N ratio had a strong negative correlation with the current light environment, and, as with nitrogen content, no distinct pattern was found with foliage age. Chlorophyll content of the foliage was not well correlated with any of the three predictor variables: current light environment, foliage age or leaf thickness. On the other hand, chlorophyll content was positively correlated with the amount of nitrogen in a leaf, and once nitrogen was considered, the current light environment was also highly significant in explaining the variation in chlorophyll content. It has been suggested that the redistribution of nitrogen both within and between leaves is a mechanism for photosynthetic acclimation to the current light environment. Within theseA. amabilis canopies, both leaf nitrogen and the chl:N ratio were strongly correlated with the current light environment, but only weakly with leaf age, supporting the idea that changing light is the driving force for the redistribution of nitrogen both within and between leaves. Thus, our results support previous theories on nitrogen distribution and partitioning. However,A max was significantly affected by both foliage age and the current light environment, indicating that changes in light alone are not enough to explain changes inA max with time.

Models of water flux through forest stands: critical leaf and stand parameters.

The basis for estimating water flux through forest stands is described using models. By means of the Penman-Monteith equation, transpiration is described in relation to the degree of coupling between the canopy and the surrounding air. Models of canopy and aerodynamic conductance are available, but more mechanistic models of stomatal conductance, and further development of turbulence theory, are needed along with improved estimates of leaf area index, leaf area distribution, and seasonal dynamics. Three models are presented to show current capabilities in estimating water uptake and flux through tree components, including the effects of capacitance. Defining conductance to water movement through tree components in terms of the properties of the pathway (sapwood area, sapwood relative conductivity, leaf area) is a useful functional approach that can be tested for a range of species at different sites. Further research is required to relate water conducting properties of tree components to architectural arrangement, especially for roots, and improved methods for measuring water potentials and partial flows at different points within the system are necessary. The role of water potential and the significance of its variability within a canopy are discussed. Relating growth processes to an integral of water potential (an accumulated product of water potential and time) is recommended. The need for scaling and integrating information about processes from one level to higher levels is recognized. The importance of understanding the roles that temporal, spatial, and developmental levels have on the ability to scale or integrate individual leaf measurements of, for example, leaf conductance to the stand level is emphasized.