Environmental influences on the productivity of three desert succulents in the south-western United States

Abstract Net CO₂ uptake over 24 h periods for shoots of Agave deserti, Ferocactus acanthodes, and Opuntia ficus-indica was measured under the ranges of water status, air temperature, and photo-synthetically active radiation (PAR) that occur in the south-western U.S.A. An environmental productivity index (EPI) was constructed indicating the overall influence of these three factors on net CO₂ uptake. Using growth chambers whose conditions were changed monthly to simulate the environmental conditions at a field site, the observed shoot dry weight increases per unit surface area changed in concert with monthly changes in EPI. The observed dry weight gain of the shoot was 17–19% lower than the predicted shoot net CO₂ uptake, which could be accounted for by carbon diversion to the roots. Mean monthly EPI was also predicted for 21 sites in the south-western U.S.A. All three species had low EPIs in the Colorado River basin, which has low annual rainfall and high summer temperatures, and in the north-central part of the region, which has low temperatures and low PAR during winter when water is available. The two native species, A. deserti and F. acanthodes, had high EPIs beyond their range in coastal southern California, where competition by other vegetation for PAR may limit net CO₂ uptake. Such regions as well as south-central California and south-central Arizona had high EPIs for all three species, indicating that these areas would be appropriate for the cultivation of O. ficus-indica.     

LEAF ORIENTATION,RADIATION INTERCEPTION,AND NOCTURNAL ACIDITY INCREASES BY THE CAM PLANT AGAVE DESERTI (AGAVACEAE)

Both field measurements and a computer model were used to study the interception of photosynthetically active radiation (PAR) by Agave deserti (Engelm.), a desert CAM plant with a basal rosette of massive opaque leaves. PAR interception was determined in the winter and the summer for upper and lower leaf surfaces on a plant with about 60 leaves. Total daily PAR on the leaf surfaces was approximately 10 mol m^-2 for a winter day and 20 mol m^-2 for a summer day. For a PAR of 15 mol m^-2, the nocturnal increase in acidity was about 0.6 mol m^-2 for both leaf surfaces and various leaf orientations, except for the oldest most horizontal leaves where the increase was less than half as large. The acidity increase measured in the field was 90% saturated at 25 mol m^-2. Thus, daytime PAR in the desert is often limiting for the nocturnal acidity increase, especially for the lower leaf surfaces. Simulated tilting of the plant by 55° so that the vertical axis pointed to the sun at solar noon on a winter day increased the PAR incident on the upper surfaces of the leaves, but did not affect the total nocturnal increase in acidity by the whole plant. Although simulated removal of alternate leaves increased the PAR per unit leaf area for the remaining leaves, it reduced the total increase in nocturnal acidity of the whole plant by 31%. PAR interception by plants on slopes facing steeply north, east, or west was substantially reduced compared to the horizontal. Thus, the model proved to be quite useful for quantifying the relation between leaf orientation, PAR interception, and nocturnal increases in acidity by A. deserti, and it indicated that the lower frequency of plants on north- compared to south-facing slopes was due to PAR limitations.     

Productivity of Agave deserti: measurement by dry weight and monthly prediction using physiological responses to environmental parameters

Summary An environmental productivity index based on physiological responses to three environmental variables was used to predict the net productivity of a common succulent perennial of the Sonoran Desert, Agave deserti, on a monthly basis. Productivity was also independently measured in the field from dry weight changes. The index was based on soil water availability, day/night air temperatures, and photosynthetically active radiation (PAR), which were individually varied in the laboratory and the effect on net CO₂ uptake by the leaves determined. From monthly precipitation, temperature, and PAR at the field site together with the responses measured in the laboratory, an index (maximum value of unity) was assigned to each of these three environmental variables and their product was termed the environmental productivity index. This index indicates the fraction of maximal CO₂ uptake expected in the field for each month (well-watered A. deserti assimilated 285 mmol CO₂ m^-2 leaf area day^-1 at PAR saturation and optimal day/night temperatures of 25° C/15° C). The dry weight analysis was based on the monthly unfolding of new leaves from the central spike of the rosette and their seasonal increase in dry weight, which were determined in the field. The production of new leaves was highly correlated with the environmental productivity index (r²=0.93), which in turn was highly correlated with the water status index (r²=0.97). After correction for respiration by folded leaves, stem, and roots, plant productivity predicted by the average environmental productivity index (0.36) over a wet June-to-October period agreed within 4% with the productivity based on the conventional dry weight analysis. The net productivity of A. deserti over this 5-month period was 0.57 kg m^-2 ground area (5.7 Mg ha^-1), a large value for a desert CAM plant. The environmental productivity index proposed here may provide a reliable means for predicting net productivity on a monthly basis, which may be particularly useful for species in relatively variable environments such as deserts.     

Temperature influences on root growth for Encelia farinosa (Asteraceae), Pleuraphis rigida (Poaceae), and Agave deserti (Agavaceae) under current and doubled CO2 concentrations

To help evaluate root distribution patterns, elongation rates of individual roots were measured as a function of soil temperature for Encelia farinosa (a C₃ species), Pleuraphis rigida (C₄), and Agave deserti (CAM), sympatric codominants in the northwestern Sonoran Desert. Measurements were made at current and doubled CO₂ concentrations under winter and summer conditions of air temperature (day/night temperatures of 17 C/10 C and 33 C/22 C, respectively). The three species had different optimal temperatures for root elongation (T_opt) under winter conditions (25 C for E. farinosa, 35 C for P. rigida, and 30 C for A. deserti); T_opt increased by 2-3 C under summer conditions for all three species. The limiting temperatures for elongation also acclimated from winter to summer conditions. The rate of root elongation at T_opt was higher under summer than winter conditions for E. farinosa (9 vs. 6 mm d⁻¹) and P. rigida (20 vs. 14 mm d⁻¹), reflecting conditions for maximum photosynthesis; no difference occurred for A. deserti (9 vs. 10 mm d⁻¹). Decreased elongation rates at extreme temperatures were associated with less cell division and reduced cell extension. The doubled CO₂ concentration increased average daily root elongation rates for A. deserti under both winter (7%) and summer (12%) conditions, reflecting increased cell extension, but had no effect for the other two species. Simulations of root elongation as a function of soil temperatures showed that maximum elongation would occur at different depths (16-20 cm for E. farinosa, 4-8 cm for P. rigida, and 0-4 cm for A. deserti) and during different seasons (winter to spring for E. farinosa, spring to summer for P. rigida, and all year for A. deserti), contributing to their niche separation. Shading of the soil surface moderated daily variations in soil temperature, reducing seasonal root elongation for winter and spring and increasing elongation for summer. Shading also altered root distribution patterns, e.g., optimal rooting depth for A. deserti and especially P. rigida increased for a hot summer day.     

Spring phenology at different altitudes is becoming more uniform under global warming in Europe

Under current global warming, high‐elevation regions are expected to experience faster warming than low‐elevation regions. However, due to the lack of studies based on long‐term large‐scale data, the relationship between tree spring phenology and the elevation‐dependent warming is unclear. Using 652k records of leaf unfolding of five temperate tree species monitored during 1951–2013 in situ in Europe, we discovered a nonlinear trend in the altitudinal sensitivity (S_A, shifted days per 100 m in altitude) in spring phenology. A delayed leaf unfolding (2.7 ± 0.6 days per decade) was observed at high elevations possibly due to decreased spring forcing between 1951 and 1980. The delayed leaf unfolding at high‐elevation regions was companied by a simultaneous advancing of leaf unfolding at low elevations. These divergent trends contributed to a significant increase in the S_A (0.36 ± 0.07 days 100/m per decade) during 1951–1980. Since 1980, the S_A started to decline with a rate of ?0.32 ± 0.07 days 100/m per decade, possibly due to reduced chilling at low elevations and improved efficiency of spring forcing in advancing the leaf unfolding at high elevations, the latter being caused by increased chilling. Our results suggest that due to both different temperature changes at the different altitudes, and the different tree responses to these changes, the tree phenology has shifted at different rates leading to a more uniform phenology at different altitudes during recent decades.     

Water Relations and Photosynthesis of a Desert CAM Plant, Agave deserti

The water relations and photosynthesis of Agave deserti Engelm., a plant exhibiting Crassulacean acid metabolism, were measured in the Colorado desert. Although no natural stomatal opening of A. deserti occurred in the summer of 1975, it could be induced by watering. The resistance for water vapor diffusion from a leaf (R_WV) became less than 20 sec cm⁻¹ when the soil water potential at 10 cm became greater than −3 bars, as would occur after a 7-mm rainfall. As a consequence of its shallow root system (mean depth of 8 cm), A. deserti responded rapidly to the infrequent rains, and the succulent nature of its leaves allowed stomatal opening to continue for up to 8 days after the soil became drier than the plant. When the leaf temperature at night was increased from 5 to 20 C, R_WV increased 5-fold, emphasizing the importance of cool nighttime temperatures for gas exchange by this plant. Although most CO₂ uptake occurred at night, a secondary light-dependent rise in CO₂ influx generally occurred after dawn. The transpiration ratio (mass of water transpired/mass of CO₂ fixed) had extremely low values of 18 for a winter day, and approximately 25 for an entire year.     

Frequencies, Microclimate and Root Properties for Three Codominant Perennials in the Northwestern Sonoran Desert on North-vs.South-facing Slopes

At a site in the northwestern Sonoran Desert the percent groundcover for the C₃subshrubEncelia farinosawas eight-times higheron more arid 20° south-facing slopes than on 20° north-facingslopes at 820 m elevation, and was six-times higher on north-facingslopes at a 300-m-lower elevation, also the more arid condition.The ground cover of the C₄bunchgrassPleuraphis rigidadecreasedover 50% from 20° north-facing slopes to the more arid conditionsof a 36° north-facing slope, a 20° south-facing slopeand a 20° north-facing slope at a 300-m-lower elevation.The CAM leaf succulentAgave desertialso had greater ground coverfor the 20° north-facing slopes at 820 m compared with 520m. For these three codominants that averaged 58% of the totalground cover, the key for the relative frequency ofE. farinosawasapparently its greater root growth on the warmer slopes duringthe winter. The key for the other two species was most likelysoil water availability, especially during the seedling stageforA. deserti. The wetter soil conditions on 20° north-facingslopes at 820 m apparently led to individual plants ofP. rigidathatwere twice as large as on south-facing slopes. Thus root propertiesmay exert the primary influence on relative plant frequencyin this desert ecosystem for which soil temperature and wateravailability are crucial.Copyright 1997 Annals of Botany Company Agave deserti; Encelia farinosa; Pleuraphis rigida; rooting patterns; soil temperature; Sonoran Desert; water availability     

Field productivity of a CAM plant, Agave salmiana, estimated using daily acidity changes under various environmental conditions

Nocturnal acidity increases of the Crassulaceae acid metabolism succulent Agave salmiana Otto ex Salm. ssp. crassispina (Trel.) Gentry were determined for various photosynthetically active radiation (PAR) levels in the field and various day/night temperatures and soil water potentials in the laboratory. When a particular factor was optimal for nocturnal acid accumulation, it was assigned an index value of unity. The product of the three indices was termed the environmental productivity index (EPI), which summarizes the effect of PAR, temperature, and water status on nocturnal acid accumulation. A monthly value for EPI was determined using microclimatic conditions at the field site near Salinas de Hidalgo, San Luis Potosf, Mexico. EPI was highly correlated with the number of leaves unfolding monthly from plants in the field (r²= 0.95), where monitoring of such leaf unfolding is a non-destructive technique for assessing plant productivity. By using the measured leaf area index for four groups of A. salmiana and a relationship between acid accumulation and net CO₂ up-take, the EPI acidity data were converted to dry weight gain per unit ground area. This measure of productivity closely agreed with the 1.05 kg m⁻² year⁻¹ determined by conventional harvesting techniques. The productivity is also in agreement with studies on other desert agaves, but about 10-fold higher than that generally considered for desert ecosystems.     

Long‐term physiological and growth responses of Himalayan fir to environmental change are mediated by mean climate

High‐elevation forests are experiencing high rates of warming, in combination with CO₂ rise and (sometimes) drying trends. In these montane systems, the effects of environmental changes on tree growth are also modified by elevation itself, thus complicating our ability to predict effects of future climate change. Tree‐ring analysis along an elevation gradient allows quantifying effects of gradual and annual environmental changes. Here, we study long‐term physiological (ratio of internal to ambient CO₂, i.e., C_i/C_a and intrinsic water‐use efficiency, iWUE) and growth responses (tree‐ring width) of Himalayan fir (Abies spectabilis) trees in response to warming, drying, and CO₂ rise. Our study was conducted along elevational gradients in a dry and a wet region in the central Himalaya. We combined dendrochronology and stable carbon isotopes (δ¹³C) to quantify long‐term trends in C_i/C_a ratio and iWUE (δ¹³C‐derived), growth (mixed‐effects models), and evaluate climate sensitivity (correlations). We found that iWUE increased over time at all elevations, with stronger increase in the dry region. Climate–growth relations showed growth‐limiting effects of spring moisture (dry region) and summer temperature (wet region), and negative effects of temperature (dry region). We found negative growth trends at lower elevations (dry and wet regions), suggesting that continental‐scale warming and regional drying reduced tree growth. This interpretation is supported by δ¹³C‐derived long‐term physiological responses, which are consistent with responses to reduced moisture and increased vapor pressure deficit. At high elevations (wet region), we found positive growth trends, suggesting that warming has favored tree growth in regions where temperature most strongly limits growth. At lower elevations (dry and wet regions), the positive effects of CO₂ rise did not mitigate the negative effects of warming and drying on tree growth. Our results raise concerns on the productivity of Himalayan fir forests at low and middle (<3,300 m) elevations as climate change progresses.     

Influences of root distribution and growth on predicted water uptake and interspecific competition

Summary Root distribution and growth measured in the field were incorporated into a water uptake model for the CAM succulent Agave deserti and its nurse plant Hilaria rigida, a common desert bunchgrass. Agave deserti responds to the infrequent rainfalls of the Sonoran Desert by extending its existing established roots and by producing new roots. Most of such root growth was completed within one month after soil rewetting, total root length of A. deserti increasing by 84% for a seedling and by 58% for a mediumsized plant in the summer. Root growth in the winter with its lower soil temperatures was approximately half as much as in the summer. For a 15-year period, predicted annual root growth of A. deserti varied more than 18-fold because of annual variations in rainfall amount and pattern as well as seasonal variation in soil temperature. Predicted annual water uptake varied 47-fold over the same period. The nurse plant, which is crucial for establishment of A. deserti seedlings, reduced seedling water uptake by 38% during an average rainfall year. Lowering the location of the root system of a medium-sized A. deserti by 0.24 m reduced its simulated annual water uptake by about 25%, reflecting the importance of shallow roots for this desert succulent. Lowering the root system of a medium-sized H. rigida by 0.28 m increased the simulated annual water uptake of an associated A. deserti seedling by 17%, further indicating the influence of root overlap on competition for water.     

Seasonal variability in the effect of elevated CO2 on ecosystem leaf area index in a scrub-oak ecosystem

We report effects of elevated atmospheric CO₂ concentration (C_a) on leaf area index (LAI) of a Florida scrub‐oak ecosystem, which had regenerated after fire for between three and five years in open‐top chambers (OTCs) and was yet to reach canopy closure. LAI was measured using four nondestructive methods, calibrated and tested in experiments performed in calibration plots near the OTCs. The four methods were: PAR transmission through the canopy, normalized difference vegetation index (NDVI), hemispherical photography, and allometric relationships between plant stem diameter and plant leaf area. Calibration experiments showed: (1) Leaf area index could be accurately determined from either PAR transmission through the canopy or hemispherical photography. For LAI determined from PAR transmission through the canopy, ecosystem light extinction coefficient (k) varied with season and was best described as a function of PAR transmission through the canopy. (2) A negative exponential function described the relationship between NDVI and LAI; (3) Allometric relationships overestimated LAI. Throughout the two years of this study, LAI was always higher in elevated C_a, rising from, 20% during winter, to 55% during summer. This seasonality was driven by a more rapid development of leaf area during the spring and a relatively greater loss of leaf area during the winter, in elevated C_a. For this scrub‐oak ecosystem prior to canopy closure, increased leaf area was an indirect mechanism by which ecosystem C uptake and canopy N content were increased in elevated C_a. In addition, increased LAI decreased potential reductions in canopy transpiration from decreases in stomatal conductance in elevated C_a. These findings have important implications for biogeochemical cycles of C, N and H₂O in woody ecosystems regenerating from disturbance in elevated C_a.     

Parent-ramet connections in Agave deserti: influences of carbohydrates on growth

Summary Agave deserti, a monocarpic perennial occurring in the northwestern Sonoran Desert, produces ramets on rhizomes that extend from the base of a parent plant. Shading ramtes to light compensation for two years did not decrease their relative growth rate (RGR) compared with unshaded ramets. However, the parents experienced a 30% decrease in total nonstructural carbohydrate (TNC) level, suggesting that carbohydrates were translocated from parents to ramets. Shaded parents had RGR's similar to unshaded parents, due in large part to consumption of 50% of the TNC reserves of shaded parents, but about 10% of the growth of the shaded parents was attributed to TNC received from their attached ramets. Estimates of parent and ramet growth separately, based on changes in TNC levels (converted to dry weight using a measured production value), net CO₂ uptake of unfolded leaves, and respiration of roots, stems, and folded leaves, were similar to measured growth of parents and ramets combined, suggesting that parents and ramets are physiologically integrated and grow as a unit. Large TNC reserves, which were also shown to support growth during conditions of water limitation in the field, enhance the growth of ramets in an environment where seedling establishment is rare.Abbreviations DW dry weight - EPI environmental productivity index - PAR photosynthetically active radiation - RGR relative growth rate - TNC total nonstructural carbohydrate     

RELATION BETWEEN MONTHLY GROWTH OF FEROCACTUS ACANTHODES AND AN ENVIRONMENTAL PRODUCTIVITY INDEX

Most productivity studies use destructive harvesting methods, prohibiting continuous plant monitoring under various environmental conditions. Here a nondestructive procedure involving the displacement of areoles from the apex was utilized to study the growth of Ferocactus acanthodes, a common barrel cactus of the Sonoran Desert. Net CO₂ uptake measured under controlled conditions of temperature, water, and photosynthetically active radiation in the laboratory was used to indicate how F. acanthodes would respond to field values of these parameters. For example, net CO₂ uptake over 24 hr was maximal at day/night air temperatures of 23 C/14 C, the mean annual values in the field, and was approximately halved at 11 C/5 C and 32 C/23 C, the monthly extreme values in the field. An environmental productivity index (EPI), constructed as the product of indices for the three environmental variables, indicated the fraction of maximal CO₂ uptake expected. The monthly production of areoles on 33 plants was highly correlated with EPI (r² = 0.81). Areole production for individual plants, however, tended to be in pulses represented by Fibonacci numbers. EPI predicted an annual stem growth of 8% compared with 9 ± 3% measured previously in the field. Thus, morphological and physiological studies can be usefully combined and applied to indicate field productivity of F. acanthodes and, by extension, of other plants.     

Plant transpiration at high elevations: Theory,field measurements,and comparisons with desert plants

Summary The influence of elevational changes on plant transpiration was evaluated using leaf energy balance equations and well-known elevational changes in the physical parameters that influence water vapor diffusion. Simulated transpirational fluxes for large leaves with low and high stomatal resistances to water vapor diffusion were compared to small leaves with identical stomatal resistances at elevations ranging from sea level to 4 km. The specific influence of various air temperature lapse rates was also tested. Validation of the simulated results was accomplished by comparing actual field measurements taken at a low elevation (300 m) desert site with similar measurements for a high elevation (2,560 m) mountain research site. Close agreement was observed between predicted and measured values of transpiration for the environmental and leaf parameters tested.Substantial increases in solar irradiation and the diffusion coefficient for water vapor in air (D _wv) occurred with increasing elevation, while air and leaf temperatures, the water vapor concentration difference between the leaf and air, longwave irradiation, and the thermal conductivity coefficient for heat in air decreased with increasing elevation. These changes resulted in temperatures for sunlit leaves that were further above air temperature at higher elevations, especially for large leaves. For large leaves with low stomatal resistances, transpirational fluxes for low-elevation desert plants were close to those predicted for high-elevation plants even though the sunlit leaf temperatures of these mountain plants were over 10°C cooler. Simulating conditions with a low air temperature lapse rate (0.003° C m^-1 and 0.004° C m^-1) resulted in predicted transpirational fluxes that were greater than those calculated for the desert site. Transpiration for smaller leaves decreased with elevation for all lapse rates tested (0.003° C m^-1 to 0.010° C m^-1). However, transpirational fluxes at higher elevations were considerably greater than expected for all leaves, especially larger leaves, due to the strong influence of increased solar heating and a greater D _wv. These results are discussed in terms of similarities in leaf structure and plant habit observed among low-elevation desert plants and high-elevation alpine and subalpine plants.     

Leaf dynamics of a deciduous forest canopy: no response to elevated CO<Subscript>2</Subscript>

Leaf area index (LAI) and its seasonal dynamics are key determinants of terrestrial productivity and, therefore, of the response of ecosystems to a rising atmospheric CO₂ concentration. Despite the central importance of LAI, there is very little evidence from which to assess how forest LAI will respond to increasing [CO₂]. We assessed LAI and related leaf indices of a closed-canopy deciduous forest for 4 years in 25-m-diameter plots that were exposed to ambient or elevated CO₂ (542 ppm) in a free-air CO₂ enrichment (FACE) experiment. LAI of this Liquidambar styraciflua (sweetgum) stand was about 6 and was relatively constant year-to-year, including the 2 years prior to the onset of CO₂ treatment. LAI throughout the 1999–2002 growing seasons was assessed through a combination of data on photosynthetically active radiation (PAR) transmittance, mass of litter collected in traps, and leaf mass per unit area (LMA). There was no effect of [CO₂] on any expression of leaf area, including peak LAI, average LAI, or leaf area duration. Canopy mass and LMA, however, were significantly increased by CO₂ enrichment. The hypothesized connection between light compensation point (LCP) and LAI was rejected because LCP was reduced by [CO₂] enrichment only in leaves under full sun, but not in shaded leaves. Data on PAR interception also permitted calculation of absorbed PAR (APAR) and light use efficiency (LUE), which are key parameters connecting satellite assessments of terrestrial productivity with ecosystem models of future productivity. There was no effect of [CO₂] on APAR, and the observed increase in net primary productivity in elevated [CO₂] was ascribed to an increase in LUE, which ranged from 1.4 to 2.4 g MJ^–1. The current evidence seems convincing that LAI of non-expanding forest stands will not be different in a future CO₂-enriched atmosphere and that increases in LUE and productivity in elevated [CO₂] are driven primarily by functional responses rather than by structural changes. Ecosystem or regional models that incorporate feedbacks on resource use through LAI should not assume that LAI will increase with CO₂ enrichment of the atmosphere.     

Altitudinal variation in photosynthetic capacity, diffusional conductance and δ13C of butterfly bush (Buddleja davidii) plants growing at high elevations

In this study, we have examined several physiological, biochemical and morphological features of Buddleja davidii plants growing at 1300 m above sea level (a.s.l.) and 3400 m a.s.l., respectively, to identify coordinated changes in leaf properties in response to reduced CO₂ partial pressure (P_a). Our results confirmed previous findings that foliar δ¹³C, photosynthetic capacity and foliar N concentration on a leaf area basis increased, whereas stomatal conductance (g_s) decreased with elevation. The net CO₂ assimilation rate (A_max), maximum rate of electron transport (J_max) and respiration increased significantly with elevation, although no differences were found in carboxylation efficiency of Rubisco (V_cmax). Consequently, also the J_max to V_cmax ratio was significantly increased by elevation, indicating that the functional balance between Ribulose‐1,5‐biphosphate (RuBP) consumption and RuBP regeneration changes as elevation increases. Our results also indicated a homeostatic response of CO₂ transfer conductance inside the leaf (mesophyll conductance, g_m) to increasing elevation. In fact, with elevation, g_m also increased compensating for the strong decrease in g_s and, thus, in the P_i (intercellular partial pressure of CO₂) to P_a ratio, leading to similar chloroplast partial pressure of CO₂ (P_c) to P_a ratio at different elevations. Because there were no differences in V_cmax, also A measured at similar PPFD and leaf temperature did not differ statistically with elevation. As a consequence, a clear relationship was found between A and g_m, and between A and the sum of g_s and g_m. These data suggest that the higher dry mass δ¹³C of leaves at the higher elevation, indicative of lower long‐term P_c/P_a ratio, cannot be attributed to changes either in diffusional resistances or in carboxylation efficiency. We speculate that because temperature significantly decreases as the elevation increases, it dramatically affects CO₂ diffusion and hence P_c/P_a and, consequently, is the primary factor influencing ¹³C discrimination at high elevation.     

Seasonal and diurnal changes in photosynthetic limitation of young sweet orange trees

This study tests the hypothesis that potted sweet orange plants show a significant variation in photosynthesis over seasonal and diurnal cycles, even in well-hydrated conditions. This hypothesis was tested by measuring diurnal variations in leaf gas exchange, chlorophyll fluorescence, leaf water potential, and the responses of CO₂ assimilation to increasing air CO₂ concentrations in 1-year-old ‘Valência’ sweet orange scions grafted onto ‘Cleopatra’ mandarin rootstocks during the winter and summer seasons in a subtropical climate. In addition, diurnal leaf gas exchange was evaluated under controlled conditions, with constant environmental conditions during both winter and summer. In relation to our hypothesis, a greater rate of photosynthesis is found during the summer compared to the winter. Reduced photosynthesis during winter was induced by cool night conditions, as the diurnal fluctuation of environmental conditions was not limiting. Low air and soil temperatures caused decreases in the stomatal conductance and in the rates of the biochemical reactions underlying photosynthesis (ribulose-1,5-bisphosphate (RuBP) carboxylation and RuBP regeneration) during the winter compared to the values obtained for those markers in the summer. Citrus photosynthesis during the summer was not impaired by biochemical or photochemical reactions, as CO₂ assimilation was only limited by stomatal conductance due to high leaf-to-air vapor pressure difference (VPD) during the afternoon. During the winter, the reduction in photosynthesis during the afternoon was caused by decreases in RuBP regeneration and stomatal conductance, which are both precipitated by low night temperature.     

Spatial and temporal variation of photosynthesis in intertidal Mazzaella laminarioides (Bory)Fredericq (Rhodophyta, Gigartinales)

The red alga Mazzaella laminarioides is an economically important species with an extended latitudinal distribution along the Chilean coast. Its populations form mid-intertidal stands, several meters wide, and therefore are differentially exposed to environmental variables that result in temporal and spatial variability in productivity. We evaluated the effect of latitude and intertidal height on productivity by in situ measurement of photosynthetic performance. Daily and seasonal variations of O₂-evolution rate and maximal quantum yield (F _v/F _m) were determined in plants from the upper and lower intertidal zone at two localities 1500,km apart. Results suggest that plant responses were mainly affected by irradiation, temperature and desiccation. At local level, upper intertidal plants showed a reduced photosynthetic rate and quantum efficiency as compared to those displayed by plants from the lower intertidal, indicating their higher level of excitation energy acclimation. Stronger acclimation differences between upper and lower intertidal plants were observed in spring and summer. Differences in photosynthetic parameters between reproductive phases were recorded in autumn and winter, regardless of the position of the individuals in the intertidal zone. The effects of tidal elevation on seasonal patterns of photosynthesis were also influenced by latitude. Seasonal variation in photosynthetic efficiency was observed in plants from the northern population at both intertidal elevations, but only at the upper intertidal level in the southern population. This study shows that production variability in M. laminarioides results from differences in the intensity of environmental factors observed seasonally at local (intertidal) and latitudinal scales.     

Hydrochory,seed banks,and regeneration dynamics along the landscape boundaries of a forested wetland

Following the environmental sieve concept, the setting in which the recruitment of Taxodium distichum occurs in, becomes increasingly restrictive from the seed to seedling stage in an impounded forested wetland. Although a wide elevational band of dispersing seed moves across the boundary of a swamp-field in the water sheet, the zone of germination is relegated to that portion of the forested wetland that draws down during the growing season. Seedling recruitment is further restricted to the uppermost zone of the winter water sheet. These patterns are likely applicable to other species of dominant swamp species, e.g., Cephalanthus occidentalis crossed the boundary of a forested wetland and abandonded field in winter flooding (November–December and November–March, respectively) in Buttonland Swamp. The elevation of the boundary was 101.3 m NGVD. While the seeds of at least 40 swamp species were dispersed across the boundary, few viable seeds were dispersed after the winter season. Kriged maps showed seeds of T. distichum and C. occidentalis dispersed in patches in the water depending on the position of the water sheet. Most species of both water- and gravity-dispersed species had a localized pattern of seed distribution (either spherical or exponential) and this indicated that seeds may not be dispersed for great distances in the swamp. Water-dispersed T. distichum and C. occidentalis had larger dispersal ranges (A ₀=225 and 195 m, respectively) than Bidens frondosa and B. discoidea (A ₀=14 and 16 m, respectively). Seed dispersal varied with season depending on the availability of seeds. In Buttonland Swamp, viable seeds typically were dispersed for T. distichum in November–June, and for C. occidentalis in November-July. Low water occurred in August 1993 and high in February 1994 (99.8 and 101.6 m NGVD, respectively). The seed banks along the landscape boundary varied in species composition according to elevation (r ² = 0.996). While the similarity of species richness between water-dispersed seeds and the seed bank at elevations that flooded (during June 1993 through May 1995) was high (10–17%), it was low between water-dispersed seeds and the seed bank at elevations that did not flood (5%). T. distichum seeds had a short germination window in that seeds germinated within a year following their production in zones that were flooded in the winter followed by drawdown during the next growing season. After 1 year, less than 5% of the T. distichum seeds remained viable on the surface of the soil. Germination of T. distichum was confined to specific elevations (above 99.3 but below 101.6 m NGVD) during this study with 4.1% of the seedlings surviving for more than 2 years at a mean of 101.4 m NGVD. All seedlings below this elevation died. To maximize natural regeneration along the boundaries of swamps in abandoned farm fields targeted for restoration, this study suggests a flood pulse regime consisting of high water in the winter to maximize dispersal of live seeds followed by low water in the summer to facilitate seed germination and seedling recruitment. Hydrologic restoration could assist in the natural recovery of damaged wetlands if a seed source exists nearby.     

Seasonal patterns of daily net photosynthesis,transpiration and net primary productivity of Juncus roemerianus and Spartina alterniflora in a Georgia salt marsh

Summary Studies of the seasonal CO₂ and water vapor exchange patterns of Juncus roemerianus and Spartina alterniflora were conducted in an undisturbed marsh community on Sapelo Island, Georgia. Daily patterns of net photosynthesis, transpiration, leaf diffusive conductance and water-use efficiency in response to ambient conditions were monitored on intact, in situ plants. Net primary productivity was calculated from the daytime CO₂ fixation totals, nighttime CO₂ loss, leaf standing stock and aboveground to belowground biomass ratios for each plant type.The tall form of S. alterniflora had higher rates of photosynthesis and higher water-use efficiency values which, in conjunction with low respiratory losses and large leaf standing crop, results in high values of net primary productivity. The environmental factors in the marsh which permit these physiological responses occur in less than 10% of the marsh. Overall, the physiological capabilities of the short form of S. alterniflora were reduced in comparison to the tall form, but the combination of environmental factors which determine the physiological responses of this form occur in a much greater portion of the marsh, and the short form of S. alterniflora dominates the Sapelo Island marshes.The response patterns of the C₃ species, J. roemerianus, differed somewhat from the height forms of S. alterniflora. A large, seasonally constant leaf standing crop coupled with moderate rates of photosynthesis resulted in a net primary productivity value which was between the tall and short height forms of S. alterniflora. However, as with the tall S. alterniflora, the environmental conditions under which this high productivity and high water loss rate can be sustained are restricted to specific regions of the environmental gradient in the marsh.Contribution No. 435 from the University of Georgia Marine Institute