Correlation of Chlorophyll Meter Readings with Gas exchange and Chlorophyll Fluorescence in Flag Leaves of Rice (Oryza sativa L.) Plants

Abstract

The objective of this study was to establish the correlation of the chlorophyll meter (SPAD) readings with the contents of chlorophyll (Chl) and ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), the gross photosynthetic rate (P_g), and the maximum quantum yield of photosystem II (PSII) (F_v/F_m) in flag leaves of rice (Oryza sativa L.) in ripening stage. The SPAD readings significantly correlated with the Chl content, the Rubisco content, P_g and F_v/F_m (R² = 0.848, 0.648, 0.671 and 0.712, respectively), which suggests that the SPAD meter has the potential to estimate the photosynthetic capacity of the flag leaves. However, both P_g and F_v/F_m had a stronger relationship with the Rubisco content than the SPAD readings, indicating that the PSII photochemical and CO₂ assimilation capacities are strongly influenced by the Rubisco content. Therefore, accurate calibration would be indispensable to obtain the physiological information from the SPAD readings of flag leaves.     

Correlation of Leaf Nitrogen,Chlorophyll and Rubisco Contents with Photosynthesis in a Supernodulating Soybean Genotype Sakukei 4

Abstract

Soybean requires more nitrogen (N) than gramineous crops because it accumulates a large amount of N in seeds, and its photosynthetic rate per leaf N is low. The supernodulating genotype Sakukei 4 has a superior symbiotic N₂ fixation capability, and thereby is potentially high-yielding. In our previous study, Sakukei 4 was characterized by having a superior ability to maintain high leaf N content and high photosynthetic rate. The objectives of this study were to know photosynthetic characteristics of Sakukei 4 in detail, especially, the responses to CO₂ concentration and light intensity, and to elucidate how the photosynthetic characteristics of Sakukei 4 are associated with the amounts of photosynthesis-related N compounds (chlorophyll and Rubisco). The three genotypes (Sakukei 4 - supernodulating cultivar derived from Enrei, Enrei - normally nodulating cultivar, En1282-non-nodulating line derived from Enrei) were grown at various N levels in this study. The CO₂ exchange rate (CER) in Sakukei 4 was higher than, or equal to that in Enrei at wide ranges of CO₂ concentrations (150-700 μmol mol-¹) and light intensities (200-1,500 μmol m-² s-¹ PPFD). Sakukei 4 had higher leaf N (N_l), chlorophyll (Chl_L) and Rubisco (Rub_L) contents per leaf area, but lower chlorophyll and Rubisco contents per leaf N content (Chl_L/N_l, Rub_L/N_l) than Enrei. The specific leaf weight (SLW) and leaf area trended to be lower in Sakukei 4 than in Enrei. These results indicate that the superior photosynthetic rate in Sakukei 4 is attributed to higher total N, chlorophyll and Rubisco contents per leaf area, but not to high rate of allocation of total N to these N compounds.     

Screening of High kcat Rubisco among Poaceae for Improvement of Photosynthetic CO2 Assimilation in Rice

Abstract

The activity of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) is a major limitation of photosynthetic CO₂ assimilation in C₃ plants. In order to find useful Rubisco for improvement of photosynthesis in rice under elevated CO₂, we analyzed the catalytic turnover rate (k_cat) of Rubisco in Poaceae including C₃ alpine plants, C₃ cold-resistant plants and C₄ plants. Rubisco in these plants showed 1.1- to 2.8-fold higher k_cat than that in rice. However, the most of high k_cat Rubisco also showed a higher km for CO₂ (Kc) than that of rice, indicating that increase in k_cat led to decrease in the affinity for CO₂. Rubisco in Festuca ovina, Phleum pratense and Sorghum bicolor showed relatively high k_cat to Kc. Although the k_cat of Rubisco in F. ovina and P. pratense was not so high (1.5-1.6 fold relative to rice), the Kc was comparable to that in rice and the amino acid sequence of RbcL shared higher identity to that in rice than that in S. bicolor. By contrast, Rubisco of S. bicolor showed considerably high k_cat (2.5-fold relative to rice), which is considered to be the most important factor for improvement of photosynthesis. In our estimation, the expression of high k_cat Rubisco of F. ovina and S. bicolor in rice could significantly enhance CO₂ assimilation at Ci of 50 Pa, the level assumed to be reached by the middle of this century.     

Characteristics of Gas Exchange and Chlorophyll Fluorescence during Senescence of Flag Leaf in Different Rice (Oryza sativa L.) Cultivars Grown under Nitrogen-Deficient Condition

Abstract

Effects of nitrogen (N) deficiency on photosynthetic gas exchange and photosystem II (PSII) photochemistry of flag leaves during grain-filling stage were investigated in six rice cultivars, Kasalath (a conventional indica), IR36 (an improved indica), Shirobeniya (a conventional japonica), Nipponbare (an improved japonica), Akenohoshi (an improved japonica-indica intermediate type) and BSI429 (an improved tropical japonica, a new plant type line) grown hydroponically in N-sufficient (NS) and N-deficient (ND) solution. From 3 to 24 days after heading (DAH), net photosynthetic rate (P_n), maximum quantum yield of photosystem II (PSII) (F_v/F_m), quantum yield of PSII electron transport ( Φ_psii), and contents of chlorophyll (Chl) and ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) in the flag leaves decreased, particularly under the ND condition in all six cultivars. A substantial difference was observed among the ND plants for the sustainability index (SI, the ratio of the value at 24 DAH to that at 3 DAH) of P_n, F_v/F_m, Φ_psii, Chl content and Rubisco content; SIs of those parameters of Akenohoshi, BSI429, Nipponbare and Shirobeniya were higher than those of IR36 and Kasalath. The SI of P_n showed significant positive correlations with those of F_v/F_m, Φ_psii, and the contents of Chl and Rubisco under the ND condition. It was concluded that the sustainability of photosynthesis in the flag leaves was mainly due to those of PSII photochemistry and electron transport, which was associated with the maintenance of Chl and Rubisco under the ND condition.     

Varietal Differences in Photosynthetic Rates in Rice Plants,with Special Reference to the Nitrogen Content of Leaves

Abstract

The photosynthetic rate in the fl ag leaf of rice at the full heading stage was examined in three japonica varieties, Koshihikari, Aikoku and Asanohikari, and the indica high-yielding variety Takanari at the same level of leaf nitrogen. At an ambient CO₂ concentration of 350 µL L-₁, Takanari had a higher photosynthetic rate and stomatal conductance than the japonica varieties when plants were compared at a leaf nitrogen content of approximately 1.5 g m-₂. Stomatal conductance increased considerably with increases in leaf nitrogen content in the japonica varieties. As a result, at a leaf nitrogen content of approximately 2.0 g m-₂, differences in terms of the photosynthetic rate among varieties were small. By contrast, there were no clear varietal differences in Rubisco content at any identical nitrogen content of leaves. We conclude that stomatal conductance is responsible for the varietal differences in photosynthetic rate examined at the same leaf nitrogen content.     

Effects of elevated atmospheric CO2 concentration on morphology of leaf blades in Chinese yam

ABSTRACT

The effects of elevated carbon dioxide concentration on the morphology of leaf blades in two Chinese yam lines under different temperature conditions were determined. Plants were grown under two [CO₂] levels, ambient (about 400 µmol mol^?1) and elevated (ambient + 200 µmol mol^?1) in the daytime, and two mean air temperature regimes, approximately ambient temperature (22.2°C) and high temperature (25.6°C). The palisade layer was thicker under elevated [CO₂] than under ambient [CO₂] in both temperature regimes, and the whole yam leaf blade was thicker under elevated [CO₂] than under ambient [CO₂] in the approximately ambient temperature regime. The numbers of chloroplasts per palisade cell and spongy cell as well as per unit profile area of palisade cell, number of starch grains per chloroplast, profile area of the starch grain, and starch-to-chloroplast area ratio in both palisade and spongy cells were higher under elevated [CO₂] than under ambient [CO₂] in both temperature regimes. Furthermore, the stomatal density on the abaxial side of the leaf blade in Chinese yam was greater under elevated [CO₂] than under ambient [CO₂] under both temperature regimes, and stomatal-pore length was higher under elevated [CO₂] than under ambient [CO₂] in the approximately ambient temperature regime. These results indicate that elevated [CO₂] positively affects the photosynthetic apparatus. The results of this study provide information for understanding the response characteristics of the leaf blade under elevated [CO₂] and a possible explanation for the positive photosynthetic responses of Chinese yam to elevated [CO₂] in our previous study.

List of Abbreviations:[CO₂]: carbon dioxide concentration     

Cytoplasmic evaluations during substitution backcrossing inSolanum

Summary Sterility-resistant cytoplasmic factors ofSolanum tuberosum spp.andigena (Juz. & Buk.) Hawkes and ofSolanum phureja (Juz. & Buk.) were combined with chromosomal genes ofS. tuberosum ssp.tuberosum. Eleven morphological characters of reciprocal F₁ and BC₁ progenies were monitored to evaluate rate of progression towards characteristics occurring in the recurrent ssp.tuberosum parent. Female and male fertilities of F₁ and BC₁ populations were also evaluated. Cytoplasmic factors did not influence morphological parameters consistently, except those relating directly to fertility. Differences between reciprocal progeny depended more upon the genes of individual parents used in crosses than on cytoplasm source. No consistent reciprocal differences occurred in tuber characters and those differences which arose seem to reflect a maternal rather than a cytoplasmic influence. F₁ and BC₁ progenies containing the sterility-resistant cytoplasmic factors of ssp.andigena or ofS. phureja had higher fertility than their respective reciprocal progenies with cytoplasm of ssp.tuberosum. Pennsylvania Agricultural Experiment Station Journal Series No 6212.     

Effects of temperature on growth and photosynthesis in the seedling stage of the sheath blight-resistant rice genotype 32R

The 32R rice genotype is resistant to sheath blight disease (ShB), with a high-yield potential. We examined effects of temperature on the plant responses of 32R in comparison with those of the ShB-susceptible rice genotype (29S) and Nipponbare (Nb, a Japonica standard cultivar). The seedlings at the 4th leaf stage of rice genotypes were exposed to 14/14, 19/14, 25/20, 31/26, 37/32 and 37/37 °C (day/night) for 5, 10 and 15 days. The dry weight, leaf area, photosynthesis, contents of ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco) and chlorophyll contents were examined. The dry weight showed lower in 32R than in 29S and Nb at a low temperature, and total dry weight correlated strongly with root dry weight and leaf area. The relative growth rate (RGR) correlated strongly with the net assimilation rate (NAR). Rubisco, chlorophyll contents and the photosynthetic rates were limited at a low temperature and showed lower in 32R than in 29S and Nb. The strong correlations between Rubisco and the rates of maximum photosynthesis and initial slope were found in 32R, but not found in 29S and Nb. In addition, RGR and NAR of 32R correlated positively with Rubisco. These suggest that 32R contains traits of cold-sensitive genotypes that are related to limiting Rubisco at a low temperature, thus diminishing photosynthesis and limiting plant growth. Differences of growth among 32R, 29S and Nb were discussed in the relation of genotypes.     

Rubisco small subunits of C4 plants,Napier grass and guinea grass confer C4-like catalytic properties on Rubisco in rice

ABSTRACT

Overexpression of Rubisco small subunit (RbcS) of C₄ plant, sorghum (sorghum bicolor) was shown to enhance the catalytic turnover rate (k _cat) of Rubisco in rice (Oryza sativa). In this study, the effects of other Rubisco small subunits of C₄ plants, Napier grass (Pennisetum purpureum) and guinea grass (Megathyrsus maximus) on kinetic properties of Rubisco in rice were studied. The expression levels of Napier grass RbcS (NgRbcS) and guinea grass RbcS (GgRbcS) proteins accounted for 41% and 45% of total RbcS, respectively in homozygous overexpression lines. The k _cat and K _m for CO₂ (Kc) of Rubisco were increased in all transgenic lines. Interestingly, the k _cat was markedly higher in NgRbcS homozygous line, whereas K _c was notably higher in GgRbcS homozygous line. Although its effects depend on species, these results suggest that the introduction of C₄ RbcS are effective approaches to alter the catalytic properties of Rubisco in rice.     

Analyzing the impact of high temperature and CO2 on net photosynthesis: biochemical mechanisms,models and genomics

Determining plant response at the biochemical/physiological level to a changing global environment is a prerequisite for constructing accurate models to predict plant productivity. High temperature and CO₂ impact plant biomass accumulation by altering the rate of net photosynthesis such that the measured rates differ greatly from the potential rates predicted from commonly used models. Such models are based on assumptions pertaining to biochemical limitations to net photosynthesis by the capacity of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), the enzyme that ultimately limits the rate of photosynthetic CO₂ fixation in C3 and C4 plants, or the capacity of electron transport to supply energy for Calvin cycle activity. Here we provide evidence that the impact of high temperature and CO₂ on net photosynthesis can be accurately calculated from predictive models based solely on Rubisco kinetics if the modeled rate of photosynthesis is adjusted for heat and CO₂-induced changes in the activation state of Rubisco. The activation state of Rubisco, which is regulated by the activity of Rubisco activase, thus appears to be the primary limitation to net photosynthesis at high temperature and/or CO₂. This limitation to net photosynthesis has not been incorporated into common biochemical models, thus compromising their effectiveness. The expression of activase mRNA was not indicative of the central role of activase in the response of photosynthesis to high temperature thus revealing limitations of using a broad genomics approach to identify the impact of environmental stress on plant metabolism.     

玉米矮花叶病抗病基因Rscmv1的精细定位

以感病亲本Mo17为轮回亲本、抗病亲本四一为供体亲本,构建近等基因系,对抗病基因Rscmv1进行精细定位。结果表明,在自交系四一中定位了两个抗病基因Rscmv1和Rscmv2,其中Rscmv1是抗源中普遍存在的主要抗病基因。以选育的近等基因系BC4F3、BC4F4和BC4F5为定位群体,在抗病基因区域内开发了9个有多态性的BAC-SSR标记,将Rscmv1定位在BAC-SSR标记A5-1和B2-1之间,其遗传距离分别为0.3 cM和0.6 cM,2个标记之间的物理距离为1.38 Mb。     

Glu A1和 Glu D1位点高分子量谷蛋白亚基共同缺失对弱筋小麦品质的影响

为了研究高分子量谷蛋白亚基(HMWGS)缺失对小麦品质的影响,以 GluA1和 GluD1位点HMWGS共同缺失材料2GS041410作供体亲本,以弱筋小麦品种扬麦13和扬麦18作轮回亲本进行回交,构建不同遗传背景的BC¹F³和BC²F³群体,测定受体、供体亲本的品质,以及回交群体BC¹F³和BC²F³中 GluA1和 GluD1位点HMWGS共同缺失纯合单株及两位点均正常表达纯合单株的品质。结果表明,供体2GS041410具有较低的SDS沉降值、较短的面团形成时间和稳定时间;在BC¹F³和BC²F³中, GluA1和 GluD1位点HMWGS共同缺失对蛋白含量影响不显著,但可显著或极显著降低SDS沉降值和水溶剂保持力(SRC)。 GluA1和 GluD1位点HMWGS双缺失在弱筋小麦品质育种中具有一定的应用价值。     

Marker-Assisted Selection of Xa21 Conferring Resistance to Bacterial Leaf Blight in indica Rice Cultivar LT2

Bacterial leaf blight of rice (BLB), caused by Xanthomonas oryzae pv. oryzae, is one of the most destructive diseases in Asian rice fields. A high-quality rice variety, LT2, was used as the recipient parent. IRBB21, which carries the Xa21 gene, was used as the donor parent. The resistance gene Xa21 was introduced into LT2 by marker-assisted backcrossing. Three Xoo races were used to inoculate the improved lines following the clipping method. Eleven BC₃F₃ lines carrying Xa21 were obtained based on molecular markers and agronomic performance. The 11 lines were then inoculated with the three Xoo races. All the 11 improved lines showed better resistance to BLB than the recipient parent LT2. Based on the level of resistance to BLB and their agronomic performance, five lines (BC₃F₃ 5.1.5.1, BC₃F₃ 5.1.5.12, BC₃F₃ 8.5.6.44, BC₃F₃ 9.5.4.1 and BC₃F₃ 9.5.4.23) were selected as the most promising for commercial release. These improved lines could contribute to rice production in terms of food security.     

Cytoplasmic substitution inSolanum. II. Tuber characteristics of reciprocal backcross progeny

Summary Recurrent backcrosses (BC) were used to introduce nuclear factors ofS. tuberosum L. ssp.tuberosum into the cytoplasm of Andean potatoesS. tuberosum ssp.andigena (Juz. and Buk.) Hawkes andS. phureja Juz. and Buk. Each generation BC was produced in reciprocal directions to test the effect of cytoplasmic substitution on yield. This report analyses tuber number, tuber weight, and tuber length of BC₂ and BC₃ substitution into ssp.andigena cytoplasm and BC₂ and BC₂ substitutions intoS. phureja cytoplasm. Direction of cross had no consistent significant effect on yield components of ssp.andigena substitution lines. InS. phureja reciprocal progenies direction of cross was responsible for significant differences of some reciprocal sets. When there were significant differences the higher yield usually occurred when ssp.tuberosum was the pistillate parent, but in some progenies when ssp.tuberosum was the pistillate parent the yield was significantly lower than in the reciprocals. It is suggested that differences between reciprocals resulted from chromosomally encoded gene action, resulting from maternal and/or paternal effects, rather than cytoplasmic factors. Authorized for publication as paper No. 7494 in the Journal Series of The Pennsylvania Agriculture Experiment Station.     

分子标记结合表型鉴定选育兼抗赤霉病、白粉病和条锈病弱筋小麦新品种扬麦38

长江中下游麦区是中国弱筋小麦优势产业带,小麦赤霉病、白粉病和条锈病是该麦区主要病害,当前弱筋小麦主导品种综合抗性较弱,影响其生产安全。为培育多抗优质弱筋小麦品种,以高产中筋小麦品种扬麦16为轮回亲本,以兼抗白粉病、条锈病的软质小麦92R137为供体亲本,构建了BC₁群体,利用分子标记在BC₁F₂代基础农艺性状较优良的株行中筛选抗白粉病基因 Pm21、抗条锈病基因 Yr26和软质麦相关基因 Pinb-D1a均纯合的单株,并鉴定BC₁F₆代对赤霉病、白粉病和条锈病的抗性,同时检测籽粒硬度、湿面筋含量、面团形成时间、稳定时间等重要品质指标以及小区产量,最终育成高抗赤霉病、免疫白粉病和高抗条锈病的弱筋小麦新品种扬麦38,于2022年通过国家农作物品种审定委员会审定。     

转Cry1A.401基因抗虫玉米CM8302和CM8303对亚洲玉米螟的抗性研究

采用农杆菌介导法获得转Cry1A.401基因抗虫玉米CM8302和CM8303,通过多代回交与自交获得BC₄F₁、BC₄F₂和BC₄F₃世代的转基因玉米材料。利用qRT-PCR和ELISA方法明确转基因抗虫玉米叶片、花丝和子粒均有Cry1A.401基因表达,不同世代间目的基因表达量无显著差异。田间接虫鉴定结果表明,CM8302和CM8303三个世代心叶期和吐丝期的抗性等级均为1级,抗性水平为高抗。室内生测结果表明,亚洲玉米螟幼虫取食CM8302心叶、花丝和子粒第6天存活率分别为0、4.0%和4.0%;取食CM8303心叶、花丝和子粒第6天存活率分别为0、4.0%和2.0%。分析显示转基因抗虫玉米CM8302和CM8303的心叶、花丝和子粒均具有显著杀虫效果且遗传稳定。     

玉米回交后代群体中供体基因组成分分析

以郑58为受体亲本,昌7-2为供体亲本,通过杂交、回交和79对亲本间有多态的SSR标记,分析BC₂F₁和BC₃F₁世代供体的基因组成分。结果表明：BC₂F₁世代单株内的供体染色体片段数平均为11.4个,BC₃F₁世代为4.8个,从BC₂F₁到BC₃F₁单株内的供体染色体片段数减少了57.6%;BC₂F₁世代单株内的供体染色体片段长度平均为99.3 cM,BC₃F₁世代为87.1 cM,BC₃F₁较BC₂F₁减少了12.3%;BC₂F₁世代单株内的供体基因组大小平均为1 132.8 cM,BC3F1世代为421.4 cM,BC₃F₁较BC₂F₁减少了62.8%。     

Effects of atmospheric CO2 concentration and defoliation on the growth of Themeda triandra

The effects of elevated atmospheric carbon dioxide (CO₂) concentration (700 μmol mol^?1) on defoliated (three clippings at 3‐week intervals) and undefoliated plants were determined for the C₄ grass Themeda triandra, Forsk. The elevated CO₂ concentration significantly increased leaf regrowth following defoliation, and total leaf production was greatest in this treatment. Shoot biomass of undefoliated plants was also increased under the elevated CO₂ concentration treatment. The primary effect of the elevated CO₂ concentration in both defoliated and undefoliated plants was an increase in individual leaf length and mass of dry matter, linked to a higher leaf water content and increased photosynthetic rates at the canopy level. Photosynthetic down‐regulation at the leaf level occurred, but this was compensated for by increased assimilation rates and greater canopy leaf area at the elevated CO₂ concentration. Increases in leaf and sheath growth of defoliated plants in the elevated CO₂ concentration treatment were lost following a final 3‐week reversion to ambient CO₂ concentration, but occurred in plants exposed to the elevated CO₂ concentration for the final 3‐week period only. In conclusion, elevated atmospheric CO₂ concentration increases shoot growth via increased leaf extension, which is directly dependent on stimulation of concurrent photosynthesis. CO₂ responsiveness is sustained following moderate defoliation but is reduced when plants experience reduced vigour as a result of maturation or high frequency of defoliation.     

Effects of elevated CO2 concentration on growth and photosynthesis of Chinese yam under different temperature regimes

Chinese yam (‘yam’) was grown at different carbon dioxide concentrations ([CO₂]), namely, ambient and elevated (ambient + 200 μmol mol^?1), under low- and high-temperature regimes in summer and autumn, separately. For comparison, rice was also grown under these conditions. Mean air temperatures in the low- and high-temperatures were respectively 24.1 and 29.1 °C in summer experiment and 20.2 and 24.9 °C in autumn experiment. In summer experiment, yam vine length, leaf area, leaf dry weight (DW), and total DW were significantly higher under elevated [CO₂] than ambient [CO₂] in both temperature regimes. Additionally, number of leaves, vine DW, and root DW were significantly higher under elevated [CO₂] than under ambient [CO₂] in the low-temperature regime. In autumn experiment, tuber DW was significantly higher under elevated [CO₂] than under ambient [CO₂] in the high-temperature regime. These results demonstrate that yam shows positive growth responses to elevated [CO₂]. Analysis of variance revealed that significant effect of [CO₂] × air temperature interaction on yam total DW was not detected. Elevated-to-ambient [CO₂] ratios of all growth parameters in summer experiment were higher in yam than in rice. The results suggest that the contribution of elevated [CO₂] is higher in yam than in rice under summer. Yam net photosynthetic rate was significantly higher under elevated [CO₂] than under ambient [CO₂] in both temperature regimes in summer. Elevated [CO₂] significantly affected on the rate in yam but not in rice in both experiments. These findings indicate that photosynthesis responds more readily to elevated [CO₂] in yam than in rice.     

5份新引进的人工合成六倍体小麦农艺及生理性状的评价

为挖掘人工合成六倍体小麦（synthetic hexaploid wheat,SHW）在育种中的应用价值,利用从国际玉米小麦改良中心引进的5份SHW（SHW1、SHW2、SHW3、SHW4、SHW5）分别与普通小麦驻麦762进行杂交、回交,构建相应回交群体（BC₂）,综合分析其农艺、产量、光合、叶绿素、水分利用效率等性状特点。结果表明,SHW的株高较高,穗下节和穗下茎较长,收获系数和产量低,分蘖多,部分材料千粒重较高;BC₂可将SHW的高分蘖、高生物量和驻麦762的多花多实、矮秆、高收获系数等性状聚合;SHW叶片的叶绿素含量在开花期之后维持较高水平,BC₂-SHW1、BC₂-SHW4和BC₂-SHW5在开花期的叶绿素含量均高于双亲,灌浆期BC₂的叶绿素含量整体低于SHW和高于驻麦762;SHW在拔节期、开花期、灌浆期有明显的光合优势,BC₂的光合速率从抽穗期之后明显低于双亲;驻麦762在抽穗期、开花期的瞬时水分利用效率最高;SHW在拔节期和灌浆期有较高的瞬时水分利用效率,在部分BC₂中得以遗传。SHW可以在小麦育种中用于农艺性状的改良及叶绿素含量和瞬时水分利用效率的提升。