Gas exchange and leaf water relations in two peanut cultivars of different drought tolerance

In summer seasons of 1991 and 1992 the gas exchange and leaf water relations were analysed in two peanut cultivars: drought tolerant cv. GG 2 (DT) and drought sensitive cv. JL 24 (DS). Soil moisture stress was imposed by withholding irrigation at pod development phase. The decrease in photosynthesis (P_N) under stress was associated with a decrease in stomatal conductance (g_s) and relative water content (RWC). The P_N and RWC were significantly higher under stress in DT than DS. On relief of stress the g_s and RWC recovered more quickly in DT than DS. The maintenance of higher RWC (>80 %), g_s and P_N under stress appears to be imparting drought tolerance in peanut.     

Ribulose-1, 5-bisphosphate carboxylase activity,chlorophyll and protein concentrations in differentPopulus clones

Ribulose-1,5-bisphosphate carboxylase activity (RuBPC), chlorophyll (chl) and protein (prot) concentrations and chlorophyll/protein (chl/prot) ratios were determined in five differentPopulus clones together with their maximal net CO₂ uptake rates (P_max). A classic reference clone (Populus ×euramericana “Robusta” (Dode) Guinier) was compared with four recently selected euramerican and interamerican crossings. Chl/prot ratio and RuBPC activity varied among the different clones, while chl a/chlb ratio showed only a very low coefficient of variation (1.7%) for the five clones. Poplar clone “Robusta” could be distinguished from the recent faster growing clones based on the different biochemical characteristics. A significant correlation was found between both total chl concentration and chl/prot ratio with P_max for the five clones.     

Comparison of the Drought Stress Responses of Tolerant and Sensitive Wheat Cultivars During Grain Filling: Changes in Flag Leaf Photosynthetic Activity,ABA Levels,and Grain Yield

Water status parameters, flag leaf photosynthetic activity, abscisic acid (ABA) levels, grain yield, and storage protein contents were investigated in two drought-tolerant (Triticum aestivum L. cv. MV Emese and cv. Plainsman V) and two drought-sensitive (cvs. GK élet and Cappelle Desprez) wheat genotypes subjected to soil water deficit during grain filling to characterize physiological traits related to yield. The leaf water potential decreased earlier and at a higher rate in the sensitive than in the tolerant cultivars. The net CO₂ assimilation rate (P _N) in flag leaves during water deficit did not display a strict correlation with the drought sensitivity of the genotypes. The photosynthetic activity terminated earliest in the tolerant cv. Emese, and the senescence of flag leaves lasted 7 days longer in the sensitive Cappelle Desprez. Soil drought did not induce characteristic differences between sensitive and tolerant cultivars in chlorophyll a fluorescence parameters of flag leaves during post-anthesis. Changes in the effective quantum yield of PSII (Φ_PSII) and the photochemical quenching (qP) depended on the genotypes and not on the sensitivity of cultivars. In contrast, the levels of ABA in the kernels displayed typical fluctuations in the tolerant and in the sensitive cultivars. Tolerant genotypes exhibited an early maximum in the grain ABA content during drought and the sensitive cultivars maintained high ABA levels in the later stages of grain filling. In contrast with other genotypes, the grain number per ear did not decrease in Plainsman and the gliadin/glutenin ratio was higher than in the control in Emese during drought stress. A possible causal relationship between high ABA levels in the kernels during late stages of grain filling and a decreased grain yield was found in the sensitive cultivars during drought stress.     

The Cause of the Difference in Leaf Net Photosynthetic Rate between Two Soybean Cultivars

To explore the cause of difference in photosynthetic performance between different cultivars of crops, leaf net photosynt rate (P _N) and photosystem 2 (PS2) photochemical efficiency (F_v/F_m), apparent quantum yield of carbon assimilation (_c), electron transport rate, photophosphorylation activity, etc. were measured in two soybean cultivars, Heinong 42 and Heinong 37. At pod setting and filling, significant differences in P _N between them were observed. The former with a higher P _N (from 7 to 38 %) had a significantly higher leaf thickness, leaf dry mass/area (LMA), chlorophyll content, soluble protein content, apparent quantum yield of electron transport through PS2 (_e), carboxylation efficiency (CE), and ribulose-1,5-bisphosphate carboxylase (RuBPC) activity. The significantly higher P _N of Heinong 42 is mainly due to its higher content and activity of RuBPC.     

氮素水平对茶树叶片氮代谢关键酶活性及非结构性碳水化合物的影响

以福鼎大白茶(FD)、保靖黄金茶1号(HJ1)、白毫早(BHZ)为材料,设置不施氮N_0(0 g)、低氮N1(11 g)、中氮N_2(22 g)和高氮N_3(33 g)4个氮素水平的盆栽实验,研究了铵态氮对3个品种茶树的根系活力、氮代谢关键酶及非结构性碳水化合物(NSC)的影响。结果表明:随着施氮水平的提高,N_2、N_3处理的茶树根系活力较对照N0显著增加(P0.05),但二者间无显著差异;叶片谷氨酰胺合成酶(GS)、谷氨酸合成酶(GOGAT)活性总体呈上升趋势;与对照比较,茶树叶片全氮和可溶性蛋白含量增加,其中HJ1在N_2和N_3处理后显著增加(P0.05);在3个茶树品种中,非结构性碳水化合物中可溶性总糖含量均呈上升趋势,淀粉含量具有品种特异性,施氮处理后3个茶树品种氮代谢关键酶活性及NSC含量变化存在差异,以HJ1的氮同化关键酶GS、GOGAT酶活性较高、根系活力较强,氮代谢产物显著增加,表明其具有较高的氮同化速率。施氮后HJ1的总NSC的含量及碳氮比的变化幅度较另外2个品种小,能够更好的保持碳氮平衡,游离氨基酸含量增幅较高,品质更优。因此,通过茶树氮代谢关键酶活性及非结构性化合物的研究能为茶树品种的品质评价以及提高茶树的品质和氮素利用效率提供依据。     

Drought-inhibited ribulose-1,5-bisphosphate carboxylase activity is mediated through increased release of ethylene and changes in the ratio of polyamines in pakchoi

To study the mechanisms of drought inhibiting photosynthesis and the role of PAs and ethylene, the photosynthetic rate (P_n), the maximal photochemical efficiency of PSII (F_v/F_m), the intercellular CO₂ concentration (C_i), photorespiratory rate (P_r), the amount of chlorophyll (chl), antioxidant enzyme activity, ethylene levels, RuBPC (ribulose-1,5-bisphosphate carboxylase) activity and endogenous polyamine levels of pakchoi were examined, and an inhibitor of S-adenosylmethionine decarboxylase (SAMDC) and an inhibitor of ethylene synthesis and spermidine (Spd) were used to induce the change of endogenous polyamine levels. The results show that drought induced a decrease in P_n and RuBPC activity, an increase in the intercellular CO₂ concentration (C_i), but no change in the actual photochemical efficiency of PSII (Φ_PSII), and chlorophyll content. In addition, drought caused an increase in the free putrescine (fPut), the ethylene levels, a decrease in the Spd and spermine (Spm) levels, and the PAs/fPut ratio in the leaves. The exogenous application of Spd and amino oxiacetic acid (AOAA, an inhibitor of ethylene synthesis) markedly reversed these drought-induced effects on polyamine, ethylene, P_n, the PAs/fPut ratio and RuBPCase activity in leaves. Methylglyoxal-bis(guanylhydrazone) (MGBG), an inhibitor of SAMDC resulting in the inability of activated cells to synthesize Spd and Spm, exacerbates the negative effects induced by drought. These results suggest that the decrease in P_n is at least partially attributed to the decrease of RuBPC activity under drought stress and that drought inhibits RuBPC activity by decreasing the ratio of PAs/fPut and increasing the release of ethylene.     

Leaf Water Diffusion Resistance in Clonal Tea (Camellia sinensis L.): Effects of Water Stress, Leaf Age and Clones

Leaf diffusion resistance was influenced by leaf age in well-wateredand water-stressed clonal tea plants. In well-watered plantsand in two of the three clones studied, young leaves showeda significantly lower diffusion resistance than old leaves.In water-stressed plants young leaves always exhibited a higherdiffusion resistance than old leaves. The highest diffusionresistance, irrespective of leaf age and water stress, occurredin clone DN which is known to be the most drought tolerant ofthe three clones studied, suggesting that drought tolerancein clone DN is caused, at least in part, by a stomatal regulationmechanism. Water release characteristic curves for the threeclones indicated differences in the water content-water potentialrelationship between young and old leaves as well as betweenclones. The drought tolerant clones had a higher relative watercontent for a given water potential compared with the drought-susceptibleclone. Camellia sinensis L., tea, diffussion resistance, water stress     

Photosynthetic acclimation to elevated CO2 in wheat cultivars

Wheat (T. aestivum) cvs. Kalyansona and Kundan grown under atmospheric (CA) and elevated CO₂ concentrations (650±50 cm³ m^-3 - CE) in open top chambers were examined for net photosynthetic rate (PN), stomatal limitation (l _s) of P _N, ribulose-1,5-bisphosphate carboxylase (RuBPC) activity, and saccharide content of the leaves. The P _N values of both CA- and CE-grown plants compared at the same CO₂ concentration showed a down regulation under CE at the post-anthesis stage. The negative acclimation of P _N appeared to be due to both stomatal and mesophyll components, and the RuBPC activity got also adjusted. There was a decrease in activation state of RuBPC under CE. In connection with this, an increased accumulation of saccharides in wheat leaf under CE was observed. Kalyansona, owing to its larger sink potential in terms of the number of grains, showed a greater enhancement under CE in both post-ear emergence dry matter production and grain yield. Under CE, this cultivar also showed a lower down regulation of P _N than Kundan. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effect of Heat Stress During Grain Filling on Phosphoenolpyruvate Carboxylase and Ribulose-1,5-bisphosphate Carboxylase/Oxygenase Activities of Various Green Organs in Winter Wheat

In two winter wheat (Triticum aestivum L.) cultivars differing in their response to high temperature, JD8 (tolerant) and J411 (sensitive) we studied the effect of heat stress on the activities of phosphoenolpyruvate carboxylase (PEPC) and ribulose-1,5-bisphosphate carboxylase (RuBPC) in green organs during grain-filling. There were significantly higher PEPC activities and lower RuBPC activities in each of the non-leaf organs (awn, glume, lemma, peduncle, and sheath) than in the flag leaf blade. Under heat stress for 12 d, the activity of RuBPC quickly declined and the activity of PEPC first increased and later declined in all organs, resulting in a great increase of the PEPC/RuBPC ratios in the organs, particularly in non-leaf organs which had a higher PEPC/RuBPC than the flag leaf blade in all times. The PEPC activity and PEPC/RuBPC ratio in every organ of JD8 were higher than those in the same organ of J411. Thus the differences in PEPC activities and PEPC/RuBPC may be associated with the differences in photosynthetic heat tolerance among the organs of the same plant or between the two cultivars.     

Effect of drought on ear and flag leaf photosynthesis of two wheat cultivars differing in drought resistance

We investigated net photosynthetic rate (P_N) of ear and two uppermost (flag and penultimate) leaves of wheat cultivars Hongmangmai (drought resistant) and Haruhikari (drought sensitive) during post-anthesis under irrigated and non-irrigated field conditions. The P_Nof ear and flag leaf were significantly higher and less affected by drought in Hongmangmai than in Haruhikari. The rate of reduction in stomatal conductance (g_s) was similar for the two cultivars, but intercellular CO₂concentration (C_i) in the flag leaf of Hongmangmai was lower than that of Haruhikari in non-irrigated treatment. No differences were observed in leaf water potential (₁) and osmotic adjustment of the flag leaf of the cultivars. These results imply that differences in photosynthetic inhibition on the flag leaf at low leaf ₁between the cultivars were primarily due to non-stomatal effects. Hence the main physiological factor associated with yield stability of Hongmangmai under drought stress may be attributed to the capacity for chloroplast activity in the flag leaf, which apparently allows sustained P_Nof flag leaf during grain filling under drought stress. The higher P_Nof ear in Hongmangmai under drought could also be related to its drought resistance.This revised version was published online in March 2005 with corrections to the page numbers.     

Effect of fruiting and drought or flooding on carbon balance of apple trees

The response of fruiting or deblossomed trees to water stress such as drought or flooding was investigated in six semi open-top cuvettes each containing one apple (Malus domestica Borkh. cv. Golden Delicious) tree. Xylem water potentials of leaves dropped from -1.2 to -4.1 MPa within 7 d of drought, the effect being enhanced by fruiting. Apple trees without fruits showed smaller reductions in net photosynthetic rate (P _N ) and dark respiration rate (R _D ) after 2 d of drought and hence more positive carbon balances relative to fruiting trees. Flooding for 4 d had a more pronounced effect on P _N than on transpiration, resulting in a reduced water use efficiency (WUE). This reduction in WUE was greater in the non-fruiting trees. Flooding reduced P _N of the whole apple canopies irrespective of fruiting; aple trees without fruits increased R _D resulting in a less positive carbon balance relative to fruiting trees. Fruiting increased the sensitivity to drought of apple trees (R _D and P _N ), but decreased their sensitivity to flooding (R _D and WUE), suggesting different adaptation mechanisms for the two forms of water stress. This revised version was published online in August 2006 with corrections to the Cover Date.     

Influence of drought stress on pistil physiology and reproductive success of two Gossypium hirsutum cultivars differing in drought tolerance

The causes of reproductive failure under drought stress (DS) are poorly understood. We hypothesized that reproductive failure was related to drought-induced changes in pistil biochemistry. To address this hypothesis, a water deficit-induced experiment was conducted with two cotton cultivars (Dexiamian 1, drought tolerant; Yuzaomian 9110, drought sensitive). Results showed that DS decreased the photosynthesis of subtending leaf and downregulated sucrose transporter gene (GhSUT-1) expression in pistil for both cultivars, resulting in lower pistil carbon accumulation which was reflected in the decreased starch accumulation. Lower starch, as potential energy, and adenosine triphosphate (ATP), as direct energy, in droughted pistils suggested less energy for pollen tube entrance into ovules, reducing the fertilized ovule number and fertilization efficiency. Further, although pistil peroxidase activity increased under DS, a higher hydrogen peroxide (H₂O₂) level still was measured in droughted pistils than well-watered pistils, damaging reproductive activities. Moreover, larger decreases in photosynthesis, pistil GhSUT-1 expression, carbon accumulation, starch and ATP contents caused by DS for Yuzaomian 9110 than Dexiamian 1, and different responses of superoxide dismutase and catalase activities, and ascorbic acid and H₂O₂ contents to DS between the two cultivars might be the reasons causing a greater decrease in fertilization efficiency for Yuzaomian 9110 than Dexiamian 1 under DS. Thus, we suggest that decreased ovule fertilization under DS was related to the disorganized carbohydrate metabolism and inefficient antioxidant defense in droughted pistils, and the effects of DS on pistil carbohydrate metabolism and antioxidant defense were more significant for drought-sensitive cultivars than drought-tolerant cultivars.     

Adaptation to high temperature mitigates the impact of water deficit during combined heat and drought stress in C3 sunflower and C4 maize varieties with contrasting drought tolerance

Heat and drought stress frequently occur together, however, their impact on plant growth and photosynthesis (P_N) is unclear. The frequency, duration and severity of heat and drought stress events are predicted to increase in the future, having severe implications for agricultural productivity and food security. To assess the impact on plant gas exchange, physiology and morphology we grew drought tolerant and sensitive varieties of C3 sunflower (Helianthus annuus) and C4 maize (Zea mays) under conditions of elevated temperature for 4 weeks prior to the imposition of water deficit. The negative impact of temperature on P_N was most apparent in sunflower. The drought tolerant sunflower retained ribulose‐1,5‐bisphosphate carboxylase/oxygenase (RubisCO) activity under heat stress to a greater extent than its drought sensitive counterpart. Maize exhibited no varietal difference in response to increased temperature. In contrast to previous studies, where a sudden rise in temperature induced an increase in stomatal conductance (G_s), we observed no change or a reduction in G_s with elevated temperature, which alongside lower leaf area mitigated the impact of drought at the higher temperature. The drought tolerant sunflower and maize varieties exhibited greater investment in root‐systems, allowing greater uptake of the available soil water. Elevated temperatures associated with heat‐waves will have profound negative impacts on crop growth in both sunflower and maize, but the deleterious effect on P_N was less apparent in the drought tolerant sunflower and both maize varieties. As C4 plants generally exhibit water use efficiency (WUE) and resistance to heat stress, selection on the basis of tolerance to heat and drought stress would be more beneficial to the yields of C3 crops cultivated in drought prone semi‐arid regions.     

Photosynthesis in Drought-Adapted Cassava

After 45 d of limited water supply, cassava (Manihot esculenta Crantz) exhibited pronounced reduction in shoot growth, high leaf fall, and decreased stomatal conductance. However, the water status of the remaining leaves was unaffected. This was combined with an amplified heliotropic response and drooping which minimises radiant energy interception at mid-day, suggesting that leaves are sensitive to high irradiance (I). In well-irrigated plants, CO₂-saturated oxygen evolution and net photosynthetic rate (P _N) in air were markedly higher (5-fold) in young (expanding) leaves than in mature leaves. Water limitation did not strongly modify CO₂-saturated oxygen evolution but it altered P _N in air for both types of leaves, although differently. The mature leaves of drought-adapted plants displayed residual rate of P _N and deteriorated photosystem 2 (PS2) photochemistry estimated from chlorophyll (Chl) a fluorescence measurements. In young leaves at moderate I, P _N was depressed by only 66 % in stressed plants. Moreover, the photochemical quenching of Chl a fluorescence and the quantum efficiency of PS2 photochemistry in young leaves were comparable in both control and stressed plants. In contrast at high I, P _N was almost null and marked decreases in the two fluorescence parameters were apparent. Hence the strong heliotropic response and drooping displayed by young leaves under water limitation is an important strategy for avoiding inactivation of P _N by high I and therefore for cassava tolerance to drought.     

Effect of NaCl,water stress or both on gas exchange and growth of wheat

Responses of wheat (Triticum aestivum L.) to various concentrations of NaCl and levels of drought were followed. With the rise of NaCl or drought, or NaCl and drought together, growth was retarded. The water content of shoots and roots was mostly unchanged. The chlorophyll and carotenoid contents were increased in plants subjected to salinity or drought or both. Only high salinity level induced a considerable decrease in net photosynthetic rate (P_N) and dark respiration rate (R_D). P_N and R_D were decreased with the decrease of soil moisture content. The content of Na⁺ in the shoots and roots of wheat plants increased with increasing salinity or decreasing soil moisture content or both treatments. Considerable variations in the content of K⁺, Ca²⁺ or Mg²⁺ were induced by the NaCl, drought or both treatments.     

Responses to flooding and drought stress by two citrus rootstock seedlings with different water-use efficiency

Leaf water relations, net gas exchange and leaf and root constituent responses to 9 days of drought stress (DS) or soil flooding were studied in 6‐month‐old seedlings of Carrizo citrange [Citrus sinensis (L.) Osb. ×Poncirus trifoliata L.; Carr] and Cleopatra mandarin (Citrus resnhi Hort. ex Tanaka; Cleo) growing in containers of native sand in the greenhouse. At the end of the drought period, both species had similar minimum stem water potentials but Cleo had higher leaf relative water content (RWC) and higher leaf osmotic potential at full turgor () than Carr. Flooding had no effect on RWC but osmotic adjustment (OA) and were higher in Cleo than in Carr. Net CO₂ assimilation rate (A_CO2) in leaves was decreased more by drought than by flooding in both species but especially in Carr. Leaf water‐use efficiency (A_CO2/transpiration) was lower in Carr and was decreased more by DS and flooding stress than in Cleo. Higher values of intercellular CO₂ concentration (C_i) in stressed plants than in control plants indicated that non‐stomatal factors including chlorophyll degradation and chlorophyll fluorescence [maximum quantum efficiency of PSII (Fv/Fm, where Fm is the maximum fluorescence and F₀, minimum fluorescence in dark‐adapted leaves)] were more important limitations on A_CO2 than stomatal conductance. In both genotypes, leaf proline was increased by drought but not by flooding, whereas both stresses increased proline in roots. Soluble sugars in leaves were increased by DS, and flooding decreased leaf sugars in Cleo. In general, DS tended to increase the concentrations of Ca, K, Mg, Na and Cl in both leaves and roots, whereas flooding tended to decrease these ions with the exception of leaf Ca in Cleo. Based on water relations and net gas exchange, Cleo was more tolerant to short‐term DS and flooding stress than Carr.     

Sequence of physiological responses in groundnut (Arachis hypogaea L.) subjected to soil moisture deficit

Responses of drought-tolerant (DT) and drought-susceptible (DS) pot-grown groundnut (Arachis hypogaea L.) varieties to changes in leaf relative water content (RWC) were studied. Water stress (WS) was imposed on 30-day-old plants for 2 weeks. Leaf RWC decreased significantly under WS conditions with simultaneous decrease in net photosynthetic rate (P _N) and stomatal conductance (g _s). Even though no significant difference was observed between DT and DS varieties with regard to RWC, DT varieties were able to maintain significantly higher P _N than DS varieties. Higher values of water use efficiency (WUE) were also observed in DT varieties during WS conditions. The decline in P _N due to WS could be attributed to both reduction in g _s (i.e. stomatal limitation) and to reduction in chlorophyll content (Chl). No significant difference in leaf area index (LAI) was found between DT and DS types and LAI was not reduced by WS. Significant differences were found among the studied groundnut varieties, but not between DT and DS types, in terms of root, aboveground, and total dry mass. These growth parameters significantly decreased under WS conditions. Based on the results, a sequence of physiological responses in groundnut crop subjected to WS was postulated.     

Photosynthetic acclimation to elevated CO2 in wheat cultivars

Wheat (T. aestivum) cvs. Kalyansona and Kundan grown under atmospheric (CA) and elevated CO₂ concentrations (650±50 cm³ m^-3 - CE) in open top chambers were examined for net photosynthetic rate (PN), stomatal limitation (l _s) of P _N, ribulose-1,5-bisphosphate carboxylase (RuBPC) activity, and saccharide content of the leaves. The P _N values of both CA- and CE-grown plants compared at the same CO₂ concentration showed a down regulation under CE at the post-anthesis stage. The negative acclimation of P _N appeared to be due to both stomatal and mesophyll components, and the RuBPC activity got also adjusted. There was a decrease in activation state of RuBPC under CE. In connection with this, an increased accumulation of saccharides in wheat leaf under CE was observed. Kalyansona, owing to its larger sink potential in terms of the number of grains, showed a greater enhancement under CE in both post-ear emergence dry matter production and grain yield. Under CE, this cultivar also showed a lower down regulation of P _N than Kundan.     

Chlorophyll fluorescence imaging of photosynthetic activity in sun and shade leaves of trees

The differences in pigment levels, photosynthetic activity and the chlorophyll fluorescence decrease ratio R _Fd (as indicator of photosynthetic rates) of green sun and shade leaves of three broadleaf trees (Platanus acerifolia Willd., Populus alba L., Tilia cordata Mill.) were compared. Sun leaves were characterized by higher levels of total chlorophylls a + b and total carotenoids x + c as well as higher values for the weight ratio chlorophyll (Chl) a/b (sun leaves 3.23–3.45; shade leaves: 2.74–2.81), and lower values for the ratio chlorophylls to carotenoids (a + b)/(x + c) (with 4.44–4.70 in sun leaves and 5.04–5.72 in shade leaves). Sun leaves exhibited higher photosynthetic rates P _N on a leaf area basis (mean of 9.1–10.1 μmol CO₂ m⁻² s⁻¹) and Chl basis, which correlated well with the higher values of stomatal conductance G _s (range 105–180 mmol m⁻² s⁻¹), as compared to shade leaves (G _s range 25–77 mmol m⁻² s⁻¹; P _N: 3.2–3.7 μmol CO₂ m⁻² s⁻¹). The higher photosynthetic rates could also be detected via imaging the Chl fluorescence decrease ratio R _Fd, which possessed higher values in sun leaves (2.8–3.0) as compared to shade leaves (1.4–1.8). In addition, via R _Fd images it was shown that the photosynthetic activity of the leaves of all trees exhibits a large heterogeneity across the leaf area, and in general to a higher extent in sun leaves than in shade leaves.     

不同施氮水平对巨桉幼树耐旱生理特征的影响

采用盆栽方法,研究了巨桉幼树在N0(不施氮)、N1(1.4g尿素·盆-1)、N2(2.8g尿素·盆-1)3个氮处理水平下,连续干旱不同时间[分别停水0(D0)、3、6、9、12、15、18d]时巨桉的生理响应。结果显示:(1)除D0外,试验期内N1和N2处理的巨桉叶片含水量(LWC)、叶片相对含水量(LRWC)和叶片保水力(LWHC)基本低于N0水平,尤其在干旱中期最为明显,表明在干旱胁迫前施氮可能对巨桉叶片水分生理产生负面作用。(2)干旱初期,氮处理间的可溶性蛋白(SP)和可溶性糖(SS)含量的差异不大,而干旱处理后期(9~18d),N0处理的SP和SS较初期明显增加,但N1和N2处理相对于N0变化较为平缓,表明施氮不利于SP和SS积累;N1和N2处理下脯氨酸(Pro)含量的增幅随着干旱胁迫时间的延长明显大于N0处理。(3)随干旱时间延长干旱程度加重,N1、N2处理巨桉叶片过氧化氢(H2O2)和丙二醛(MDA)含量明显高于N0处理,表明施氮使得巨桉在干旱条件下水分缺乏更为严重,产生更多的活性氧(ROS)。(4)整个干旱处理期内,施氮并未显著改变巨桉的超氧化物歧化酶(SOD)活性和抗坏血酸(AsA)含量,但N1和N2的过氧化物酶(POD)活性明显高于N0。(5)施氮增加了巨桉叶片的色素含量并在干旱初期和中期保持较高水平,在干旱初期(0~3d)增加了巨桉叶片的净光合速率(Pn),但随着干旱时间的延长而迅速下降;施氮(N1、N2)的蒸腾速率(Tr)和气孔导度(Gs)在干旱初期均显著小于N0,但在干旱中后期(6d以后)各处理间差异不显著且均处于极低水平。研究表明,水分充足时施氮有助于增强巨桉的光合同化能力,促进其生长,但遇到持续干旱时施氮更易面临水分亏缺,降低其抵抗干旱的能力,因此在巨桉人工林的经营管理过程中,不应在干旱或季节性干旱即将到来之前施氮,若干旱过程中需施氮则应采取灌溉等途径保证其充足的水分供应。