Ecophysiological studies of Sonoran Desert plants

Summary The gas exchange and water relations of two Sonoran Desert plants are compared during contrasting periods of water and heat stress. Photosynthesis of Acacia greggii, a winter deciduous shrub, and Cercidium microphyllum, a chlorophyllous stemmed tree, show a moderate correlation with dawn plant water potential. For both species a relationship between stomatal conductance and dawn plant water potential was not apparent, although A. greggii demonstrated a greater overall stomatal conductance. This affected a greater daytime decrease in plant water potential at all levels of water stress and suggests A. greggii is less sensitive to water stress. Our results suggest the lower limit for gross photosynthesis occurs when dawn plant water potentials are less than -44 and -31 bars for the shrub and tree species, respectively. During periods of extreme water and heat stress the photosynthetic capacity of both species is regulated more by mesophyll than stomatal conductance. However, partial stomatal closure causes plant water potential to increase during the day and exceed dawn values. During periods of minimal water and heat stress the daily course of photosynthesis parallels the change in stomatal conductance and irradiance. Maximum gross photosynthesis rates are nearly three-fold higher than the rates observed during periods of stress, with those of A. greggii generally greater than the rates observed in plants of C. microphyllum.     

Ecophysiological studies of Sonoran Desert plants

Summary The gas exchange and water relations of two Sonoran Desert plants was measured throughout a 12-month period. Seasonal photosynthesis patterns of both plants followed the seasonal variation in plant water potential. Ambrosia deltoidea, a drought-deciduous shrub, is mainly winter-spring active since maximum photosynthesis rates of 38 mg CO₂ dm^-2 h^-1 were measured at this time. This plant is characterized by marked seasonal variations in plant water potential, and was deciduous for approximately 120 days when plant water potential was below-50 bars. Olneya tesota, a non-riparian microphyllous tree, is evergreen and photosynthetically active throughout the entire year, although demonstrating maximum photosynthesis rates of 12 mg CO₂ dm^-2 h^-1 in spring and summer. The deep-rooted tree species maintains a favorable year-round water balance since minimum plant water potentials were seldom below-33 bars. The two species maintain a relatively high water use efficiency throughout the year, despite the high evaporative gradient characteristic of the Sonoran Desert.The leaves are the major site for carbon assimilation, contributing 87 and 81% of the annual carbon gain for the shrub and tree species, respectively. Above-ground gross primary production throughout the 12-month period was estimated solely from the leaf ¹⁴CO₂ assimilation studies. This production estimate was compared to above-ground net primary production determined by the harvest method. For both plant species gross production was interpreted to exceed net production by nearly a three-fold difference. On a per plant basis gross production was estimated to be 1.14 and 7.42 kg dry wt plant^-1 yr^-1 for A. deltoidea and O. tesota. The large difference between net and gross production is probably related to year-round utilization of carbon.This research was supported by National Science Foundation Grant BMS 74-02671-A04 through the U.S./I.B.P. Desert Biome at Utah State University     

Correlated photosynthetic responses and habitat factors of two successional tree species

Summary Ulmus alata and Diospyros virginiana are components of the shrubearly tree communities of old-field succession in several areas in the deciduous forests of eastern North America. In these habitats, the plants experience high insolation, high temperatures, and low soil moisture during the summer. They exhibit pronounced daily changes in water potential and usually develop more negative water potentials as the season progresses. The species light saturate at 1,150 E m^-2 sec^-1 with photosynthetic rates of 15 mg CO₂ dm^-2 h^-1 for U. alata and 17 mg CO₂ dm^-2 h^-1 for D. virginiana. The optimum temperatures for photosynthesis are 25°C. Ulmus alata maintains maximum photosynthesis to water potentials of-14 bars and recovers from-20 bars to 60% of maximum photosynthesis within 10 hrs after watering. When they are deprived of water, twigs of D. virginiana exhibit faster decline in photosynthesis and leaf conductance than twigs of U. alata. The two species have somewhat different response to the environmental of high insolation and low water supply. Unlike Ulmus, Diospyros virginiana has some adaptations which may explain the persistence of a few individuals in mature forests.     

Increases in Desert Shrub Productivity under Elevated Carbon Dioxide Vary with Water Availability

Productivity of aridland plants is predicted to increase substantially with rising atmospheric carbon dioxide (CO₂) concentrations due to enhancement in plant water-use efficiency (WUE). However, to date, there are few detailed analyses of how intact desert vegetation responds to elevated CO₂. From 1998 to 2001, we examined aboveground production, photosynthesis, and water relations within three species exposed to ambient (around 38 Pa) or elevated (55 Pa) CO₂ concentrations at the Nevada Desert Free-Air CO₂ Enrichment (FACE) Facility in southern Nevada, USA. The functional types sampled—evergreen (Larrea tridentata), drought-deciduous (Ambrosia dumosa), and winter-deciduous shrubs (Krameria erecta)—represent potentially different responses to elevated CO₂ in this ecosystem. We found elevated CO₂ significantly increased aboveground production in all three species during an anomalously wet year (1998), with relative production ratios (elevated:ambient CO₂) ranging from 1.59 (Krameria) to 2.31 (Larrea). In three below-average rainfall years (1999–2001), growth was much reduced in all species, with only Ambrosia in 2001 having significantly higher production under elevated CO₂. Integrated photosynthesis (mol CO₂ m⁻² y⁻¹) in the three species was 1.26–2.03-fold higher under elevated CO₂ in the wet year (1998) and 1.32–1.43-fold higher after the third year of reduced rainfall (2001). Instantaneous WUE was also higher in shrubs grown under elevated CO₂. The timing of peak canopy development did not change under elevated CO₂; for example, there was no observed extension of leaf longevity into the dry season in the deciduous species. Similarly, seasonal patterns in CO₂ assimilation did not change, except for Larrea. Therefore, phenological and physiological patterns that characterize Mojave Desert perennials—early-season lags in canopy development behind peak photosynthetic capacity, coupled with reductions in late-season photosynthetic capacity prior to reductions in leaf area—were not significantly affected by elevated CO₂. Together, these findings suggest that elevated CO₂ can enhance the productivity of Mojave Desert shrubs, but this effect is most pronounced during years with abundant rainfall when soil resources are most available.     

Comparative study of leaf carbon gain in saplings ofThujopsis dolabrata var.hondai andQuercus mongolica var.grosseserrata in a cool-temperate deciduous forest

Seasonal courses of leaf CO₂ gas exchange in a growing season were examined in saplings ofThujopsis dolabrata var.hondai andQuercus mongolica var.grosseserrata in a cool temperate deciduous forest. Between the two tree species there were no large differences in the light compensation point of leaf photosynthesis, except for the season of new leaf expansion. However, light-saturated rates of net photosynthesis were obviously high inT. dolabrata var.hondai. EvergreenT. dolabrata var.hondai saplings had large photosynthetic production in two seasons, before the emergence of new foliage and after foliage fall of the overstory deciduous trees, because of the significantly high solar radiant energy penetrating under the forest canopy during the seasons. Saplings of deciduousQ. mongolica var.grosseserrata were heavily shaded throughout the growing season by foliage of the overstory trees, which resulted in a low daily surplus production. The annual surplus production of leaves in the growing season was estimated to be 2300 mmol CO₂ m⁻² inT. dolabrata var.hondai and −100 mmol CO₂ m⁻², slightly negative, inQ. mongolica var.grosseserrata. These results supported the high survivability ofT. dolabrata var.hondai saplings and the high mortality ofQ. mongolica var.grosseserrata in the deciduous forest.     

Effects of elevated CO2 on fine root dynamics in a Mojave Desert community: a FACE study

Fine roots (≤1 mm diameter) are critical in plant water and nutrient absorption, and it is important to understand how rising atmospheric CO₂ will affect them as part of terrestrial ecosystem responses to global change. This study's objective was to determine the effects of elevated CO₂ on production, mortality, and standing crops of fine root length over 2 years in a free‐air CO₂ enrichment (FACE) facility in the Mojave Desert of southern Nevada, USA. Three replicate 25 m diameter FACE rings were maintained at ambient (～370 μmol mol^?1) and elevated CO₂ (～550 μmol mol^?1) atmospheric concentrations. Twenty‐eight minirhizotron tubes were placed in each ring to sample three microsite locations: evergreen Larrea shrubs, drought‐deciduous Ambrosia shrubs, and along systematic community transects (primarily in shrub interspaces which account for ～85% of the area). Seasonal dynamics were similar for ambient and elevated CO₂: fine root production peaked in April–June, with peak standing crop occurring about 1 month later, and peak mortality occurring during the hot summer months, with higher values for all three measures in a wet year compared with a dry year. Fine root standing crop, production, and mortality were not significantly different between treatments except standing crop along community transects, where fine root length was significantly lower in elevated CO₂. Fine root turnover (annual cumulative mortality/mean standing crop) ranged from 2.33 to 3.17 year^?1, and was not significantly different among CO₂ treatments, except for community transect tubes where it was significantly lower for elevated CO₂. There were no differences in fine root responses to CO₂ between evergreen (Larrea) and drought‐deciduous (Ambrosia) shrubs. Combined with observations of increased leaf‐level water‐use efficiency and lack of soil moisture differences, these results suggest that under elevated CO₂ conditions, reduced root systems (compared with ambient CO₂) appear sufficient to provide resources for modest aboveground production increases across the community, but in more fertile shrub microsites, fine root systems of comparable size with those in ambient CO₂ were required to support the greater aboveground production increases. For community transects, development of the difference in fine root standing crops occurred primarily through lower stimulation of fine root production in the elevated CO₂ treatment during periods of high water availability.     

Comparison of the effect of elevated CO2 on an invasive species (Centaurea solstitialis) in monoculture and community settings

The ongoing increase in atmospheric CO₂ concentration ([CO₂]) is likely to change the species composition of plant communities. To investigate whether growth of a highly invasive plant species, Centaurea solstitialis (yellow starthistle), was affected by elevated [CO₂], and whether the success of this species would increase under CO₂ enrichment, I grew the species in serpentine soil microcosms, both as a monoculture and as a component of a grassland community. Centaurea grown in monoculture responded strongly to [CO₂] enrichment of 350 mol mol^–1, increasing aboveground biomass production by 70%, inflorescence production by 74%, and midday photosynthesis by an average of 132%. When grown in competition with common serpentine grassland species, Centaurea responded to CO₂ enrichment with similar but nonsignificant increases (+69% aboveground biomass, +71% inflorescence production), while total aboveground biomass of the polyculture increased by 28%. Centaurea's positive CO₂ response in monoculture and parallel (but non-significant) response in polyculture provoke questions about possible consequences of increasing CO₂ for more typical California grasslands, where the invader already causes major problems.     

Diurnal and Seasonal Patterns of Photosynthesis and Respiration by Stems of Populus tremuloides Michx

The photosynthetic and respiratory rates of 5- to 7-year-old aspen stems (Populus tremuloides Michx.) were monitored in the field for 1 year to determine the seasonal patterns. The stem was not capable of net photosynthesis, but the respiratory CO₂ loss from the stem was reduced by 0 to 100% depending on the time of year and the level of illumination as a result of bark photosynthesis. The monthly dark respiratory rate ranged from 0.24 mg CO₂/dm²· hr in January to a maximum 7.4 mg CO₂/dm²· hr in June. Individual measurements ranged from 0.02 mg CO₂/dm²· hr in February to 12.3 mg CO₂/dm²· hr in June. Gross photosynthesis followed a pattern similar to the dark respiratory rate. The mean monthly rate was highest in June (1.65 mg CO₂/dm²· hr) and lowest in December (0.02 mg CO₂/dm²· hr). Individual measurements ranged from 0.0 mg CO₂/dm²· hr in winter to 5.5 mg CO₂/dm²· hr in July.     

C4 photosynthetic carbon metabolism in the leaves of aromatic tropical grasses — I. Leaf anatomy,CO2 compensation point and CO2 assimilation

A few species of Cymbopogon and Vetiveria are potentially important tropical grasses producing essential oils. In the present study, we report on the leaf anatomy and photosynthetic carbon assimilation in five species of Cymbopogon and Vetiveria zizanioides. Kranz-type leaf anatomy with a centrifugal distribution of chloroplasts and exclusive localization of starch in the bundle sheath cells were common among the test plants. Besides the Kranz leaf anatomy, these grasses displayed other typical C₄ characteristics including a low (0–5 µl/l) CO₂ compensation point, lack of light saturation of CO₂ uptake at high photon flux densities, high temperature (35°C) optimum of net photosynthesis, high rates of net photosynthesis (55–67 mg CO₂ dm^-2 leaf area h^-1), little or no response of net photosynthesis to atmospheric levels of O₂ and high leaf ¹³C/¹²C ratios. The biochemical studies with ¹⁴CO₂ indicated that the leaves of the above plant species synthesize predominantly malate during short term (5 s) photosynthesis. In pulse-chase experiments it was shown that the synthesis of 3-phosphoglycerate proceeds at the expense of malate, the major first formed product of photosynthesis in these plant species.     

Variation in mistletoe seed deposition: effects of intra- and interspecific host characteristics

We investigated differences in host infection by a desert mistletoe, Phoradendron californicum, and examined one of the processes that contributes to these differences: variation in seed deposition among host individuals and species. In the Sonoran Desert, P. californicum parasitizes the sympatric leguminous trees Olneya tesota, Cercidium microphyllum, Prosopis velutina, Acacia constricta, and Acacia greggii. We hypothesized that seed deposition depends on host height and crown architecture. At a site in Arizona, frequency of infection did not reflect host relative abundance. Olneya tesota was parasitized at a higher frequency than expected from its abundance and maintained the highest mistletoe loads per individual host. In contrast, P. velutina was infected less frequently than expected. Infection frequency increased with host tree height for all hosts. Mistletoe seed deposition by avian dispersers differed among host species and was disproportionately high in O. tesota and P. velutina. Seed deposition was higher in infected than in non‐infected host trees, and increased with tree height in O. tesota but not in C. microphyllum. We suspect that increased seed deposition with height in O. tesota may be due to the preference of seed‐dispersing birds for higher perches. Some host tree species, such as C. microphyllum and A. constricta, probably received fewer mistletoe seeds because birds avoid hosts with dense and spiny crowns. Mistletoe populations are plant metapopulations in which host trees are patches and the frequency of infection in each host species/patch type is the result of interspecific differences in the balance between mistletoe colonization and extinction. From this perspective, our study of host use and seed dispersal is a metapopulation study of patch occupancy and propagule distribution among available patch types. Our seed‐dispersal study demonstrates that the mechanisms that create pattern in patchy plant populations can be investigated in mistletoe systems.     

Carbon dioxide concentrations within forest canopies—variation with time, stand structure, and vegetation type

Vertical CO₂ profiles (between 0.02 and 14.0 m) were studied in forest canopies of Pinus contorta, Populus tremuloides, and in a riparian forest with Acer negundo and Acer grandidentatum during two consecutive growing seasons. Profiles, measured continuously during 1- to 13-day periods in four to five stands differing in overstorey canopy area index (CAI < 4.5; including leaves, branches and stems), were well stratified, with highest [CO₂] just above the forest floor. Canopy [CO₂] profiles were influenced by stand structure (CAI, presence of understorey vegetation), and were highly dependent on vegetation type (deciduous and evergreen). A doubling of CAI in Acer spp. and P. tremuloides stands did not show an effect on upper canopy [CO₂], when turbulent mixing was high. However, increasing understorey biomass in Acer spp. stands had a profound effect on lower canopy [CO₂]. In open stands with a vigorous understorey layer, higher soil respiration rates were offset by increased understorey gas exchange, resulting in [CO₂] below those of the convective boundary layer (CBL). Midday depletions up to 20 ppmv below CBL values could be frequently observed in deciduous canopies. In evergreen canopies, [CO₂] stayed generally above the CBL background values, [CO₂] profiles were more uniform, and gradients were smaller than in deciduous stands with similar CAI. Seasonal changes of canopy [CO₂] reflected changes in soil respiration rates as well as plant phenology and gas exchange of both dominant tree and understorey vegetation. Seasonal patterns were less pronounced in evergreen than in deciduous forests.     

Contribution of stem CO2 fixation to whole-plant carbon balance in nonsucculent species

In many plant species that remain leafless part of the year, CO₂ fixation occurring in green stems represents an important carbon gain. Traditionally, a distinction has been made between stem photosynthesis and corticular photosynthesis. All stem photosynthesis is, sensu stricto, cortical, since it is carried out largely by the stem cortex. We proposed the following nomenclature: stem net photosynthesis (SNP), which includes net CO₂ fixation by stems with stomata in the epidermis and net corticular CO₂ fixation in suberized stems, and stem recycling photosynthesis (SRP), which defines CO₂ ling in suberized stems. The proposed terms should reflect differences in anatomical and physiological traits. SNP takes place in the chlorenchyma below the epidermis with stomata, where the net CO₂ uptake occurs, and it resembles leaf photosynthesis in many characteristics. SRP is found in species where the chlorenchyma is beneath a well-developed stomata-free periderm and where reassimilation of internally respired CO₂ occurs. SNP is common in plants from desert ecosystems, rates reaching up to 60% of the leaf photosynthetic rate. SRP has been demonstrated in trees from temperate forests and it offsets partially a carbon loss by respiration of stem nonphotosynthetic tissues. Reassimilation can vary between 7 and 123% of respired CO₂, the latter figure implying net CO₂ uptake from the atmosphere. Both types of stem photosynthesis contribute positively to the carbon economy of the species, in which they occur; they are advantageous to the plant because they allow the maintenance of physiological activity during stress, an increase of integrated water use efficiency, and they provide the carbon source used in the production of new organs.     

Photosynthesis of Conifers in Relation to Annual Growth Cycles and Dry Matter Production

Observations that deciduous larch species can show annual growth increments equal to or greater than evergreen conifers, and that the saturating light intensity for photosynthesis in needles of Larix leptolepis was almost twice those for several evergreen conifers, led to a study of the photosynthetic mechanism in L. leptolepis. Several features of photosynthesis in L. leptolepis placed this species in an intermediate position between classical C₃ and C₄ plants. Incorporation of ¹⁴C from ¹⁴CO₂ by enzyme preparations of larch needles was eight times greater with PEP as substrate than with ribulose bis phosphate; a chlorophyll a/b ratio of 3.5 was obtained; needles possessed a green starch-containing endodermis but with little orientation of mesophyll cells to this “bundle sheath”; no clear ultrastructural dimorphism was observed between chloroplasts of mesophyll and endodermal cells; a CO₂-compensation point of 20 μl-l^?1 was recorded; and the first measurable product of photosynthesis appeared to be malate rather than phospho-glyceric acid. These results are discussed in relation to the deciduous habit of L. leptolepis and its high productivity in comparison with other conifers.     

Comparative responses of model C3 and C4 plants to drought in low and elevated CO2

Interactive effects of CO₂ and water availability have been predicted to alter the competitive relationships between C3 and C4 species over geological and contemporary time scales. We tested the effects of drought and CO₂ partial pressures (pCO₂) ranging from values of the Pleistocene to those predicted for the future on the physiology and growth of model C3 and C4 species. We grew co-occurring Abutilon theophrasti (C3) and Amaranthus retroflexus (C4) in monoculture at 18 (Pleistocene), 27 (preindustrial), 35 (current), and 70 (future) Pa CO₂ under conditions of high light and nutrient availability. After 27 days of growth, water was withheld from randomly chosen plants of each species until visible wilting occurred. Under well-watered conditions, low pCO₂ that occurred during the Pleistocene was highly limiting to C3 photosynthesis and growth, and C3 plants showed increased photosynthesis and growth with increasing pCO₂ between the Pleistocene and future CO₂ values. Well-watered C4 plants exhibited increased photosynthesis in response to increasing pCO₂, but total mass and leaf area were unaffected by pCO₂. In response to drought, C3 plants dropped a large amount of leaf area and maintained relatively high leaf water potential in remaining leaves, whereas C4 plants retained greater leaf area, but at a lower leaf water potential. Furthermore, drought-treated C3 plants grown at 18 Pa CO₂ retained relatively greater leaf area than C3 plants grown at higher pCO₂ and exhibited a delay in the reduction of stomatal conductance that may have occurred in response to severe carbon limitations. The C4 plants grown at 70 Pa CO₂ showed lower relative reductions in net photosynthesis by the end of the drought compared to plants at lower pCO₂, indicating that CO₂ enrichment may alleviate drought effects in C4 plants. At the Pleistocene pCO₂, C3 and C4 plants showed similar relative recovery from drought for leaf area and biomass production, whereas C4 plants showed higher recovery than C3 plants at current and elevated pCO₂. Based on these model systems, we conclude that C3 species may not have been at a disadvantage relative to C4 species in response to low CO₂ and severe drought during the Pleistocene. Furthermore, C4 species may have an advantage over C3 species in response to increasing atmospheric CO₂ and more frequent and severe droughts.     

Dependence of the aboveground CO2 exchange rate on tree size in field-grown hinoki cypress (Chamaecyparis obtusa)

The CO₂ exchange of the aboveground parts for five different-sized 17-year-old (as of 1991) hinoki cypress (Chamaecyparis obtusa) trees growing in the field was non-destructively measured over one year, using an open CO₂ exchange system. The CO₂ exchange of individual trees decreased with decreasing tree sizes, such as aboveground phytomass, leaf mass and leaf area. However, the CO₂ exchange abruptly decreased near the smallest-suppressed sample tree. The size dependence was well described by a generalized power function. The annual gross photosynthesis of individual trees was proportional to the square root of leaf mass or leaf area. The dependence of CO₂ exchange on annual phytomass increment was described by a simple power function with an exponent value less than unity, suggesting that CO₂ exchange per unit of phytomass increment was lower in larger-sized trees than in smaller-sized trees. The mean photosynthetic activity of a tree, i.e., gross photosynthesis per unit of leaf area, slightly increased to its highest value with decreasing leaf area and then decreased abruptly near the smallest sample tree. The maximum value of mean photosynthetic activity was estimated to be 2.85 kg CO₂ m⁻² year⁻¹ for a leaf area of 1.56 m² tree⁻¹. The ratio of mean photosynthetic activity to the maximum photosynthetic activity was the highest in an intermediate tree and decreased gradually toward larger-sized trees by ca. 60% and also decreased toward the smallest suppressed tree by ca. 35%.     

Elevated CO2 and drought alter tissue water relations of birch (Betula populifolia Marsh.) seedlings

The effect of increasing atmospheric CO₂ concentrations on tissue water relations was examined in Betula populifolia, a common pioneer tree species of the northeastern U.S. deciduous forests. Components of tissue water relations were estimated from pressure volume curves of tree seedlings grown in either ambient (350 l l^–1) or elevated CO₂ (700 l l^–1), and both mesic and xeric water regimes. Both CO₂ and water treatment had significant effects on osmotic potential at full hydration, apoplasmic fractions, and tissue elastic moduli. Under xeric conditions and ambient CO₂ concentrations, plants showed a decrease in osmotic potentials of 0.15 MPa and an increase in tissue elastic moduli at full hydration of 1.5 MPa. The decrease in elasticity may enable plants to improve the soil-plant water potential gradient given a small change in water content, while lower osmotic potentials shift the zero turgor loss point to lower water potentials. Under elevated CO₂, plants in xeric conditions had osmotic potentials 0.2 MPa lower than mesic plants and decreased elastic moduli at full hydration. The increase in tissue elasticity at elevated CO₂ enabled the xeric plants to maintain positive turgor pressures at lower water potentials and tissue water contents. Surprisingly, the elevated CO₂ plants under mesic conditions had the most inelastic tissues. We propose that this inelasticity may enable plants to generate a favorable water potential gradient from the soil to the plant despite the low stomatal conductances observed under elevated CO₂ conditions.     

The response of plants to elevated CO2

Tree saplings, two groups of three species from each of two deciduous tree communities, were grown in competition at three CO₂ concentrations and two light levels. After one growing season, biomass was measured to assess the effect of CO₂ on community structure, and nitrogen and phosphorus concentrations were measured for leaves, stems, and roots of all trees. Gas-exchange measurements were made on the same species grown under the same CO₂ concentrations.Photosynthetic capacity (rate of photosynthesis at saturating CO₂ and light) tended to decline as CO₂ concentration increased, but differences were not statistically significant. Stomatal conductance declined significantly as CO₂ increased. Nitrogen and phosphorus concentrations generally declined as CO₂ increased, but there were some unexpected patterns in roots and stems. CO₂ concentration did not significantly affect the overall growth of either community after one season, but the relative biomass of each species changed in a complex way, depending on CO₂ light level, and community.     

Seasonal soil‐surface carbon fluxes from the root systems of young Pinus radiata trees growing at ambient and elevated CO2 concentration

Elevated atmospheric CO₂ concentration may result in increased below‐ground carbon allocation by trees, thereby altering soil carbon cycling. Seasonal estimates of soil surface carbon flux were made to determine whether carbon losses from Pinus radiata trees growing at elevated CO₂ concentration were higher than those at ambient CO₂ concentration, and whether this was related to increased fine root growth. Monthly soil surface carbon flux density (f) measurements were made on plots with trees growing at ambient (350) and elevated (650 μmol mol^?1) CO₂ concentration in large open‐top chambers. Prior to planting the soil carbon concentration (0.1%) and f (0.28 μmol m^?2 s^?1 at 15 °C) were low. A function describing the radial pattern of f with distance from tree stems was used to estimate the annual carbon flux from tree plots. Seasonal estimates of fine root production were made from minirhizotrons and the radial distribution of roots compared with radial measurements of f. A one‐dimensional gas diffusion model was used to estimate f from soil CO₂ concentrations at four depths. For the second year of growth, the annual carbon flux from the plots was 1671 g y^?1 and 1895 g y^?1 at ambient and elevated CO₂ concentrations, respectively, although this was not a significant difference. Higher f at elevated CO₂ concentration was largely explained by increased fine root biomass. Fine root biomass and stem production were both positively related to f. Both root length density and f declined exponentially with distance from the stem, and had similar length scales. Diurnal changes in f were largely explained by changes in soil temperature at a depth of 0.05 m. Ignoring the change of f with increasing distance from tree stems when scaling to a unit ground area basis from measurements with individual trees could result in under‐ or overestimates of soil‐surface carbon fluxes, especially in young stands when fine roots are unevenly distributed.     

Growth of regenerated tree seedlings associated with microclimatic change in a mature larch plantation after harvesting

In order to further develop methods of sustainable forest management, we evaluated the effects of logging practices during the winter on microclimatic factors and growth of four seral deciduous broad-leaved tree seedlings regenerated in a larch plantation in northern Japan. We found that winter logging practices drastically changed microclimatic factors, especially in terms of light intensity and the vertical distribution pattern of CO₂ concentration. Harvesting overstory larch trees increased photosynthetic photon flux (PPF), which may enhance the photosynthesis of understory plants. We examined the undergrowth for tree seedlings of the following species: two late successional species, Prunus ssiori and Carpinus cordata; one gap phase species, Magnolia hyporeuca; and one mid-late successional species, Quercus mongolica var. crispula. All of the four studied tree species responded well to the practices of winter logging after the second year of harvesting overstoried larch trees. The current shoot diameter and current shoot length increased significantly. Therefore, we conclude that winter logging practices should improve shoot growth of deciduous broad-leaved tree seedlings regenerated in a larch plantation through the change in environmental factors that accompanies larch harvest.     

Greater seed production in elevated CO2 is not accompanied by reduced seed quality in Pinus taeda L.

For herbaceous species, elevated CO₂ often increases seed production but usually leads to decreased seed quality. However, the effects of increased atmospheric CO₂ on tree fecundity remain uncertain, despite the importance of reproduction to the composition of future forests. We determined how seed quantity and quality differed for pine trees grown for 12 years in ambient and elevated (ambient+200 μL L^?1) CO₂, at the Duke Forest free‐air CO₂ enrichment (FACE) site. We also compared annual reproductive effort with yearly measurements of aboveground net primary productivity (ANPP), precipitation (P), potential evapotranspiration (PET) and water availability [precipitation minus potential evapotranspiration (P?PET)] to investigate factors that may drive interannual variation in seed production. The number of mature, viable seeds doubled per unit basal area in high‐CO₂ plots from 1997 to 2008 (P<0.001), but there was no CO₂ effect on mean seed mass, viability, or nutrient content. Interannual variation in seed production was positively related to ANPP, with a similar percentage of ANPP diverted to reproduction across years. Seed production was negatively related to PET (P<0.005) and positively correlated with water availability (P<0.05), but showed no relationship with precipitation (P=0.88). This study adds to the few findings that, unlike herbaceous crops, woody plants may benefit from future atmospheric CO₂ by producing larger numbers of seeds without suffering degraded seed quality. Differential reproductive responses between functional groups and species could facilitate woody invasions or lead to changes in forest community composition as CO₂ rises.