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Tissue-specific expression conferred by the S-adenosyl-L-methionine synthetase promoter of Arabidopsis thaliana in transgenic poplar.

In Arabidopsis the promoter of the gene encoding S-adenosyl-L-methionine synthetase (SAM-S) Psam-1 confers expression preferentially in the vascular tissue. In search for promoters that drive expression in particular cells of the lignifying tissues in trees, we have analyzed the expression pattern conferred by the Psam-1 promoter in transgenic poplar. Histochemical analyses demonstrated beta-glucuronidase (GUS) activity mainly in phloem and cortex tissue throughout the plant, and in root tips. Fluorimetric assays showed high GUS activity in the tissues outside (phloem, cortex and cork) compared to those inside (xylem and pith) of the cambial layer. In contrast, the endogenous SAM-S activity was high in tissues inside and low in tissues outside of the cambial layer. RNA gel blot analysis demonstrated a high transcript level of the endogenous sam-s gene(s) in tissues both outside and inside the cambial layer. This indicates that the low SAM-S activity in the bark was at least partially due to translational and/or post-translational regulation of the endogenous sam-s gene(s). In dormant transgenics, the tissue specificity was conserved, but the activity levels were up to 10-fold reduced.

Arabidopsis↗

Leaf growth characteristics of fast-growing poplar hybrids Populus trichocarpa x P. deltoides.

Fast-growing hybrid poplar trees (Populus trichocarpa Torr. & A. Gray x P. deltoides Bartr. ex Marsh.) were compared with slower-growing parental types in both field and laboratory experiments to determine physiological components of leaf growth that could be closely related to biomass production. Stem volume was correlated with individual leaf area (r = 0.81) and leaf growth rate (r = 0.82). Hybrids had a greater total leaf area, not because they produced more leaves, but because they had larger leaves than either parental type. The greater leaf size of the hybrids may be explained by inheritance of larger cell number from P. deltoides and larger cell size from P. trichocarpa. Rates of enlargement of isolated leaf discs in liquid culture were approximately 50% of those observed in intact leaves of field-grown plants.

Journal Article↗

Comparison of yellow poplar growth models on the basis of derived growth analysis variables.

Quadratic and cubic polynomials, and Gompertz and Richards asymptotic models were fitted to yellow poplar growth data. These data included height, leaf area, leaf weight and new shoot height for 23 weeks. Seven growth analysis variables were estimated from each function. The Gompertz and Richards models fitted the data best and provided the most accurate derived variables. However, the Richards model was more complex to fit.

Journal Article↗

Enhancement of the endogenous cytokinin concentration in poplar by transformation with Agrobacterium T-DNA gene ipt.

The agrobacterial isopentenyltransferase (ipt) gene, under the control of its native promoter, was transferred to poplar (Populus tremula x P. alba) by an Agrobacterium co-cultivation method. The ipt-transformed stem explants developed calli that regenerated many buds in the absence of exogenous cytokinins. Microcuttings of the ipt transformants exhibited frequently branching shoots with short internodes that were unable to root. In this material, the concentrations of zeatin, zeatin riboside and isopentenyladenosine, determined by an indirect enzyme-linked immunosorbent assay (ELISA), were 4.8-. 17.1- and 14.6-fold higher, respectively, than in non-transformed shoots. Results are discussed with regard to cytokinin metabolism.

Journal Article↗

Effects of near-lethal heat stress on bud break, heat-shock proteins and ubiquitin in dormant poplar (Populus nigra Charkowiensis x P. nigra incrassata).

We assessed the effects of near-lethal heat stress on bud break, heat-shock proteins (HSPs) and ubiquitin in hybrid poplar (Populus nigra (L.) Charkowiensis x P. nigra (L.) incrassata). Shoots, with 10-15 buds each, were collected from September to March and exposed to temperatures between 20 and 60 degrees C for 2 h. Shoots were then placed in a greenhouse at 18-22 degrees C with supplemental light and cumulative bud break was recorded over a 4-week period. Samples of bud tissues were collected during and up to 96 h after heat treatment for protein analysis. De novo synthesis of proteins was monitored by exposing excised buds to [(35)S]-methionine for 3 h before, during, or after heat treatment. Heat treatments of 40-45 degrees C resulted in both a release from endodormancy and a decrease in thermal units needed for bud break during ecodormancy. The response to near-lethal heat stress was complex and was affected by intrinsic thermal sensitivity. Heat treatments were least effective during August and became progressively more effective as endodormancy progressed. In the later stages of ecodormancy, a heat treatment of 45 degrees C either inhibited bud break or killed the buds. Although temperatures of 42.5 to 45 degrees C inhibited incorporation of [(35)S]-methionine into proteins for at least 48 h, several HSPs were synthesized in response to temperatures of 40-45 degrees C. Immunoblots indicated that one of the heat-induced proteins was immunologically related to HSP70. Increases in free and conjugated forms of ubiquitin were also observed in response to heat treatment. Production of HSPs and ubiquitin, however, was not consistently associated with the heat treatments that induced the highest percentage of bud break. The roles of heat-induced protein degradation, HSPs, and ubiquitin in overcoming dormancy by near-lethal heat stress are discussed.

Journal Article↗

Clonal and seasonal differences in leaf osmotic potential and organic solutes of five hybrid poplar clones grown under field conditions.

Leaf osmotic potential at full turgor (Psi(pio)) and the major solutes that contribute to osmotic potential were characterized in five hybrid poplar clones of Populus trichocarpa Torr. & Gray x P. deltoides Bartr. (TD) and P. deltoides x P. nigra L. (DN), growing under field conditions at two sites in eastern Washington and Oregon, USA. Trees were drip irrigated with 46, 76 or 137 cm of supplemental irrigation during each growing season. Trees at Wallula, WA, which were in their third growing season in 1994, were sampled twice a year for two years (1994 and 1995), and trees at Boardman, OR, which were in their second growing season in 1994, were sampled once a year for three years (1994-1996). At Wallula, the TD and DN clones exhibited lower predawn leaf water potentials in the 46-cm treatment than in the 137-cm treatment (-1.2 versus -0.7 MPa) during a hot, dry period in July 1994. Clone TD had a lower Psi(pio) than Clone DN (-1.67 versus -1.56 MPa) during the same period and the difference was also evident in 1995 (-1.81 versus -1.72 MPa) when trees were in their fourth growing season. There was also a significant treatment effect on Psi(pio) in Clone TD, with trees in the 46-cm treatment having lower Psi(pio) than trees in the 137-cm treatment in July 1994. At Boardman, Psi(pio) was generally high with no treatment differences during the 1994-96 samplings. The TD clones had significantly lower Psi(pio) than the DN clones in 1994 (-1.44 versus -1.36 MPa) and 1996 (-1.72 versus -1.54 MPa), but there was no difference between clones in 1995 (-1.40 versus -1.43 MPa). In 1995, at Wallula, osmotic adjustment in Clone TD was largely accounted for by an increase in sucrose, which constituted 70% of total organic solutes. Although the total concentration of free primary amino acids in this clone was 28% higher in trees in the 46-cm treatment than in trees in the 137-cm treatment, amino acids constituted only a small fraction of the total solute pool. Sixty-two percent of total solutes were inorganic ions in Clone TD compared to 52% in Clone DN, and potassium was the main ion constituting about 30% of total solutes and 50% of total ions. However, the clonal difference in Psi(pio) was not fully accounted for by the difference in solute concentration. Osmotic potential at full turgor declined over the growing season and with age. We conclude that, because the extent of osmotic adjustment exhibited by these clones was small, other drought resistance mechanisms contributed to the clonal differences in field performance.

Journal Article↗

Growth, productivity and water use in three hybrid poplar clones.

Three drought treatments (none, medium and severe) were applied to three container-grown, hybrid poplar clones (Beaupré, Trichobel and Ghoy) as a series of drying cycles over two years-in a greenhouse in 1994 and outside in 1995. Leaf area development, annual biomass production and annual water use were recorded. Clone Beaupré had greater rates of leaf area development (up to 11.8 x 10(-4) m(2) degrees C day(-1)) and subsequently greater dry matter production (up to 816 g tree(-1) year(-1)) and water use (up to 168 l tree(-1) year(-1)) than the other two clones. In all clones, rates of leaf area development, annual biomass production and water use were less under drought conditions than under well-watered conditions. On a proportional basis, the highest reduction in biomass by drought was in Trichobel (up to 74%), whereas, in absolute terms, the reduction was greatest in Beaupré (up to 500 g tree(-1)). However, under drought conditions, annual biomass production was greater in Clone Beaupré than in the other clones. The relationship between water use and biomass production was constant over both years, and was similar for Beaupré and Trichobel at 4.4 g l(-1), but was less for Ghoy at 3.5 g l(-1).

Journal Article↗

Interactive effects of nitrogen and water availabilities on gas exchange and whole-plant carbon allocation in poplar.

Cuttings of balsam spire hybrid poplar (Populus trichocarpa var. Hastata Henry x Populus balsamifera var. Michauxii (Dode) Farwell) were grown in sand culture and irrigated every 2 (W) or 10 (w) days with a solution containing either 3.0 (N) or 0.5 (n) mol nitrogen m(-3) for 90 days. Trees in the WN (control) and wn treatments had stable leaf nitrogen concentrations averaging 19.4 and 8.4 mg g(-1), respectively, over the course of the experiment. Trees in the Wn and wN treatments had a similar leaf nitrogen concentration, which increased from 12.0 to 15.8 mg g(-1) during the experiment. By the final harvest, mean stomatal conductances of trees in the wN and wn treatments were less than those of trees in the Wn and WN treatments (1.8 versus 4.6 mm s(-1)). Compared to the WN treatment, biomass at the final harvest was reduced by 61, 72 and 75% in the Wn, wN and wn treatments, respectively. At the final harvest, WN trees had a mean total leaf area of 4750 +/- 380 cm(2) tree(-1) and carried 164 +/- 8 leaves tree(-1) with a specific leaf area of 181 +/- 16 cm(2) g(-1), whereas Wn trees had a smaller mean total leaf area (1310 +/- 30 cm(2) tree(-1)), because of the production of fewer leaves (41 +/- 6) with a smaller specific leaf area (154 +/- 2 cm(2) g(-1)). A greater proportion of biomass was allocated to roots in Wn trees than in WN trees, but component nitrogen concentrations adjusted such that there was no Wn treatment effect on nitrogen allocation. Compared with WN trees, rates of photosynthesis and respiration per unit weight of tissue of Wn trees decreased by 28 and 31%, respectively, but the rate of photosynthesis per unit leaf nitrogen remained unaltered. The wN and Wn trees had similar leaf nitrogen concentrations; however, compared with the Wn treatment, the wN treatment decreased mean total leaf area (750 +/- 50 cm(2) tree(-1)), number of leaves per tree (29 +/- 2) and specific leaf area (140 +/- 6 cm(2) g(-1)), but increased the allocation of biomass and nitrogen to roots. Net photosynthetic rate per unit leaf nitrogen was 45% lower in the wN treatment than in the other treatments. Rates of net photosynthesis and respiration per unit weight of tissue were 48 and 33% less, respectively, in wN trees than in Wn trees.

Journal Article↗

Nitrogen and potassium effects on xylem cavitation and water-use efficiency in poplars.

Effects of N and K nutrition on drought and cavitation resistance were examined in six greenhouse-grown poplar clones: Populus trichocarpa (Torr. & Gray) and its hybrids with P. deltoides Bartr. and P. euramericana (Dole) Guinier, before and after preconditioning to water stress. Both tendency to cavitate and water-use efficiency (WUE) increased when N supply was increased, whereas K supply had little impact on cavitation. Mean xylem vessel diameters increased from 36.6 &mgr;m at low-N supply to 45.2 &mgr;m at high-N supply. Drought-hardy clones, which were relatively resistant to cavitation, had the smallest mean vessel diameters. Vulnerability to cavitation had a weakly positive relationship with vessel diameter, and a negative correlation with transpiration. Drought hardening offered no protection against cavitation in a subsequent drought. Under drought conditions, increasing N supply increased leaf loss and decreased water potentials, whereas increasing K supply decreased leaf loss. Drought-resistant clones exhibited similar WUE to drought-susceptible clones, but had smaller, more numerous stomata and greater leaf retention under drought conditions.

Journal Article↗

Transpiration by two poplar varieties grown as coppice for biomass production.

Fast-growing tree clones selected for biomass plantations are highly productive and therefore likely to use more water than the agricultural crops they replace. We report field measurements of transpiration through the summer of 1994 from two poplar clones, Beaupré (Populus trichocarpa Torr. & A. Gray x P. deltoides Bartr. ex Marsh.) and Dorschkamp (P. deltoides x P. nigra L.), grown as unirrigated short-rotation coppice in southern England. Stand transpiration was quantified by scaling up from sap flow measurements made with the heat balance method in a sample of stems. Leaf conductances, leaf area development, meteorological variables and soil water deficit were also measured to investigate the response of the trees to the environment. High rates of transpiration were found for Beaupré. In June, when soil water was plentiful, the mean (+/- SD) transpiration rate over an 18-day period was 5.0 +/- 1.8 mm day(-1), reaching a maximum of 7.9 mm day(-1). Transpiration rates from Dorschkamp were lower, as a result of its lower leaf area index. High total leaf conductances were measured for both Beaupré (0.34 +/- 0.17 mol m(-2) s(-1)) and Dorschkamp (0.39 +/- 0.16 mol m(-2) s(-1)). Leaf conductance declined slightly with increasing atmospheric vapor pressure deficit in both clones, but only in Beaupré did leaf conductance decrease as soil water deficit increased.

Journal Article↗

Quenching analysis in poplar clones exposed to ozone.

Effects of ozone (O(3)) on photo-inhibition of photosynthesis were investigated in two poplar clones (Populus deltoides Bartr. ex Marsh. x maximowiczii A. Henry clone Eridano, O(3)-sensitive, and P. x euramericana (Dode) Guinier clone I-214, O(3)-resistant) by using pulse amplitude modulation chlorophyll fluorescence. After a 15-day exposure to O(3) at 60 nl l(-1) for 5 h day(-1), the effective photosynthetic quantum yield of photosystem II (PSII), as indicated by the ratio of variable to maximum chlorophyll fluorescence, significantly declined in both young and mature leaves of the two clones. Compared with control samples, mature leaves of both clones showed differences in fluorescence quenching and quantum yield of PSII when exposed to ozone fumigation. We conclude that photo-inhibition is caused by transformation of active reaction centers to photochemically inactive centers that dissipate excitation energy into heat, thus causing non-photochemical fluorescence quenching. The changes observed in chlorophyll a fluorescence of O(3)-treated young leaves in both clones are consistent with O(3) having a negligible effect on photosynthetic performance, but inducing a premature decrease in chlorophyll a concentration. A possible role of O(3) pollution on chlorophyll fluorescence yield is discussed.

Journal Article↗

Radial variation in sap velocity as a function of stem diameter and sapwood thickness in yellow-poplar trees.

Canopy transpiration and forest water use are frequently estimated as the product of sap velocity and cross-sectional sapwood area. Few studies, however, have considered whether radial variation in sap velocity and the proportion of sapwood active in water transport are significant sources of uncertainty in the extrapolation process. Therefore, radial profiles of sap velocity were examined as a function of stem diameter and sapwood thickness for yellow-poplar (Liriodendron tulipifera L.) trees growing on two adjacent watersheds in eastern Tennessee. The compensation heat pulse velocity technique was used to quantify sap velocity at four equal-area depths in 20 trees that ranged in stem diameter from 15 to 69 cm, and in sapwood thickness from 2.1 to 14.8 cm. Sap velocity was highly dependent on the depth of probe insertion into the sapwood. Rates of sap velocity were greatest for probes located in the two outer sapwood annuli (P1 and P2) and lowest for probes in closest proximity to the heartwood (P3 and P4). Relative sap velocities averaged 0.98 at P1, 0.66 at P2, 0.41 at P3 and 0.35 at P4. Tree-specific sap velocities measured at each of the four probe positions, divided by the maximum sap velocity measured (usually at P1 or P2), indicated that the fraction of sapwood functional in water transport (f(S)) varied between 0.49 and 0.96. There was no relationship between f(S) and sapwood thickness, or between f(S) and stem diameter. The fraction of functional sapwood averaged 0.66 +/- 0.13 for trees on which radial profiles were determined. No significant depth-related differences were observed for sapwood density, which averaged 469 kg m(-3) across all four probe positions. There was, however, a significant decline in sapwood water content between the two outer probe positions (1.04 versus 0.89 kg kg(-1)). This difference was not sufficient to account for the observed radial variation in sap velocity. A Monte-Carlo analysis indicated that the standard error in estimated mean f(S) declined rapidly with increasing sample size. At n = 10, the coefficient of variation in mean f(S) was 7% and at n = 15 it was slightly less than 5%. These observations indicate that radial variation in sap velocity is an important, albeit often overlooked, source of uncertainty in the scaling process. Failure to recognize that not all sapwood is functional in water transport will introduce systematic bias into estimates of both tree and stand water use. Future studies should devise sampling strategies for assessing radial variation in sap velocity and such strategies should be used to identify the magnitude of this variation in a range of non-, diffuse- and ring-porous trees.

Journal Article↗

Free-air CO2 enrichment (FACE) enhances biomass production in a short-rotation poplar plantation.

This paper investigates the possible contribution of Short Rotation Cultures (SRC) to carbon sequestration in both current and elevated atmospheric CO2 concentrations ([CO2]). A dense poplar plantation (1 x 1 m) was exposed to a [CO2] of 550 ppm in Central Italy using the free-air CO2 enrichment (FACE) technique. Three species of Populus were examined, namely P. alba L., P. nigra L. and P. x euramericana Dode (Guinier). Aboveground woody biomass of trees exposed to elevated [CO2] for three growing seasons increased by 15 to 27%, depending on species. As a result, light-use efficiency increased. Aboveground biomass allocation was unaffected, and belowground biomass also increased under elevated [CO2] conditions, by 22 to 38%. Populus nigra, with total biomass equal to 62.02 and 72.03 Mg ha-1 in ambient and elevated [CO2], respectively, was the most productive species, although its productivity was stimulated least by atmospheric CO2 enrichment. There was greater depletion of inorganic nitrogen from the soil after three growing seasons in elevated [CO2], but no effect of [CO2] on stem wood density, which differed significantly only among species.

Biomass↗

Diurnal and seasonal changes in stem increment and water use by yellow poplar trees in response to environmental stress.

To evaluate indicators of whole-tree physiological responses to climate stress, we determined seasonal, daily and diurnal patterns of growth and water use in 10 yellow poplar (Liriodendron tulipifera L.) trees in a stand recently released from competition. Precise measurements of stem increment and sap flow made with automated electronic dendrometers and thermal dissipation probes, respectively, indicated close temporal linkages between water use and patterns of stem shrinkage and swelling during daily cycles of water depletion and recharge of extensible outer-stem tissues. These cycles also determined net daily basal area increment. Multivariate regression models based on a 123-day data series showed that daily diameter increments were related negatively to vapor pressure deficit (VPD), but positively to precipitation and temperature. The same model form with slight changes in coefficients yielded coefficients of determination of about 0.62 (0.57-0.66) across data subsets that included widely variable growth rates and VPDs. Model R2 was improved to 0.75 by using 3-day running mean daily growth data. Rapid recovery of stem diameter growth following short-term, diurnal reductions in VPD indicated that water stored in extensible stem tissues was part of a fast recharge system that limited hydration changes in the cambial zone during periods of water stress. There were substantial differences in the seasonal dynamics of growth among individual trees, and analyses indicated that faster-growing trees were more positively affected by precipitation, solar irradiance and temperature and more negatively affected by high VPD than slower-growing trees. There were no negative effects of ozone on daily growth rates in a year of low ozone concentrations.

Circadian Rhythm↗

Gas exchange and foliage characteristics of two poplar clones grown in soil amended with industrial waste.

Plants can remove or immobilize various environmental contaminants; however, little is known about the physiological mechanisms underlying responses to soil amendment with biosolids contaminated with heavy metals. We investigated the responses of cuttings of hybrid poplar clones Eridano and I-214 grown for a season in soil amended with nutrient-rich organic material from tanneries, which contains potentially toxic amounts of heavy elements. Plant growth traits, gas exchange parameters, stomatal density and leaf layer thickness of frozen-hydrated leaves, and foliar concentrations of heavy metals and nitrogen were determined. Overall, soil amendment increased net assimilation rate and growth, but the cuttings accumulated only small quantities of heavy metal soil contaminants.

Industrial Waste↗

Phytohormones and shoot growth in a three-generation hybrid poplar family.

Hybrid vigor for secondary growth in poplar has been linked with increased gibberellin (GA) concentration in cambial tissue, but the relationship between concentrations of GAs and hybrid vigor of primary growth has not been investigated. We explored associations between concentrations of GAs, abscisic acid (ABA) and indoleacetic acid (IAA) and shoot extension in a hybrid family of Populus deltoides Bartr. ex Marsh. (Clone ILL-129) and P. trichocarpa Torr. & Gray (Clone 93-968) and two F1 and 67 F2 genotypes. Rapidly elongating subapical internodes from shoots of 4-year-old clonal saplings were selected for hormone analysis. The F1 hybrids displayed heterosis (hybrid vigor) for primary growth as a result of the complementation of dominance for increased internode length from the P. trichocarpa parent and dominance for increased diameter from the P. deltoides parent. Internodes from the faster-elongating shoots of the P. trichocarpa parent had a fourfold higher concentration of bioactive GA1 and higher concentrations of GA20, GA44, GA29 and GA8 than the P. deltoides parent. However, the two fast-growing F1 hybrids had low concentrations of all five GAs, with concentrations similar to those of the slower-elongating P. deltoides parent. Concentrations of ABA and IAA were correlated with GA concentrations and there was thus no evidence of a consistent promoting or inhibiting effect of ABA or IAA within the F1 family. These results indicate that heterosis for internode growth was not primarily regulated by hormone concentration. The segregating population of F2 hybrids was analyzed to assess the possible utility of hormone screening as a selection tool. The internodes of most of the F2 hybrids were smaller than those of their F1 parents and the larger P. trichocarpa parent. Among the F2 hybrids, mature internode length varied fourfold, and was correlated with lengths of young, subapical internodes from which hormones were analyzed. In these internodes, GA1 concentrations were negatively correlated with length (r = 0.41), diameter (0.33) and mass (0.50). Indoleacetic acid concentration was negatively correlated only with diameter (r = 0.37) and positively correlated with GA1 (r = 0.46), whereas ABA concentration was not correlated with any parameter. Thus, in the F2 population, variation in hormone concentration was not primarily responsible for the variation in shoot growth, indicating that selection for individuals with high GA concentration may not be an effective strategy for identifying vigorous hybrid genotypes.

Abscisic Acid↗

Physiological responses of wild type and putrescine-overproducing transgenic cells of poplar to variations in the form and concentration of nitrogen in the medium.

We determined: (a) the physiological consequences of overproduction of putrescine in transgenic poplar (Populus nigra x maximoviczii) cells expressing an ornithine decarboxylase transgene; and (b) effects of variation in nitrogen (N) concentration of the medium on cellular polyamine concentration in transgenic and non-transgenic cells. Cells grown in the presence of supplemental (to the normal concentrations of N sources in the growth medium) and reduced amounts of NH4NO3 and KNO3 were used to study effects on membrane permeability, mitochondrial respiratory activity, protein accumulation, growth rates and changes in cellular polyamine concentration. The N concentration of the MS medium was not a limiting factor for continued overproduction of putrescine in transgenic cells. However, continued supplies of NH4+ and NO3- were required to maintain homeostatic amounts of putrescine in both cell lines. The presence of high amounts of putrescine in transgenic cells had significant effects on the physiological parameters measured. Compared with non-transgenic cells, transgenic cells had greater plasma membrane permeability, less tolerance to NH4NO3, more tolerance to KNO3, and accumulated higher amounts of soluble protein.

Cells, Cultured↗

Transgenic poplar characterized by ectopic expression of a pine cytosolic glutamine synthetase gene exhibits enhanced tolerance to water stress.

Physiological responses to water stress in hybrid poplar (INRA 7171-B4, Populus tremula L. x P. alba L.) lines transformed to overexpress a pine cytosolic glutamine synthetase (GS1) gene were compared with those of non-transgenic plants. Before, during and after a drought treatment, net photosynthetic rates (Anet) were higher in transgenic than in non-transgenic plants. Stomatal conductance (gs) was higher in transgenic than in non-transgenic plants before, but not after exposure to drought. Before drought treatment, a sudden reduction in photosynthetic photon flux caused a greater burst of CO2 efflux in transgenic than non-transgenic plants, indicating greater photorespiratory activity. Drought caused greater reductions in photochemical quenching, photosystem II (PSII) antennae transfer efficiency (Fv'/Fm') and light-adapted PSII yield (PhiPSII) in non-transgenic than in transgenic plants, especially at low irradiances. Antennae-based thermal dissipation was higher in transgenic plants than in non-transgenic plants both during the imposition of drought and 1 or 3 days after the relief of drought. Under severe water stress and subsequently, transgenic plants maintained a higher expression of glutamine synthetase, glutamate synthase and Rubisco and higher concentrations of chlorophyll and glycine than non-transgenic plants. These findings indicate that overexpression of pine cytosolic GS1 enhanced sustained photosynthetic electron transport capacity during severe stomatal limitation. The data also suggest that ectopic expression of cytosolic GS increases photorespiratory activity, and that this serves as a protective sink for electrons from photosynthetic reaction centers.

Chlorophyll↗