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1.
The morphological, biochemical, and physiological characteristics of silver birch (Betula pendula Roth), curly birch (B. pendula Roth var. carelica), and downy birch (B. pubescens Ehrh.) at early stages of ontogenesis in natural conditions were investigated. Some intra- and interspecific peculiarities of the morphophysiological properties were determined. Priority development of the underground mass for the seedlings of curly birch and that of the aboveground mass for the seedlings of silver birch and downy birch were found. The leaf of curly birch is developed more actively as compared with those in the other two species under equal potential possibility of the growth of their leaves by elongation. A high similarity of the mineral composition (C, N, P, K) of different forms and species of birches was demonstrated. For silver birch the maximum values of the stomatal conductance, rate of photosynthesis, and transpiration under similar atmospheric and soil conditions were obtained. In downy birch we found a higher efficiency of water use. The highest values of the maximum rate of ribulose 1,5-bisphosphate carboxylase/oxygenase carboxylation were found in curly birch. This may serve as an indicator of its shade tolerance, as compared with silver birch and downy birch.  相似文献   

2.
The finding of unique Chl d- and Chl f-containing cyanobacteria in the last decade was a discovery in the area of biology of oxygenic photosynthetic organisms. Chl b, Chl c, and Chl f are considered to be accessory pigments found in antennae systems of photosynthetic organisms. They absorb energy and transfer it to the photosynthetic reaction center (RC), but do not participate in electron transport by the photosynthetic electron transport chain. However, Chl d as well as Chl a can operate not only in the light-harvesting complex, but also in the photosynthetic RC. The long-wavelength (Qy) Chl d and Chl f absorption band is shifted to longer wavelength (to 750 nm) compared to Chl a, which suggests the possibility for oxygenic photosynthesis in this spectral range. Such expansion of the photosynthetically active light range is important for the survival of cyanobacteria when the intensity of light not exceeding 700 nm is attenuated due to absorption by Chl a and other pigments. At the same time, energy storage efficiency in photosystem 2 for cyanobacteria containing Chl d and Chl f is not lower than that of cyanobacteria containing Chl a. Despite great interest in these unique chlorophylls, many questions related to functioning of such pigments in primary photosynthetic processes are still not elucidated. This review describes the latest advances in the field of Chl d and Chl f research and their role in primary photosynthetic processes of cyanobacteria.  相似文献   

3.
In this study, we have compared the photosynthetic characteristics of two contrasting species of Tradescantia plants, T. fluminensis (shade-tolerant species), and T. sillamontana (light-resistant species), grown under the low light (LL, 50–125 µmol photons m?2 s?1) or high light (HL, 875–1000 µmol photons m?2 s?1) conditions during their entire growth period. For monitoring the functional state of photosynthetic apparatus (PSA), we measured chlorophyll (Chl) a emission fluorescence spectra and kinetics of light-induced changes in the heights of fluorescence peaks at 685 and 740 nm (F 685 and F 740). We also compared the light-induced oxidation of P700 and assayed the composition of carotenoids in Tradescantia leaves grown under the LL and HL conditions. The analyses of slow induction of Chl a fluorescence (SIF) uncovered different traits in the LL- and HL-grown plants of ecologically contrasting Tradescantia species, which may have potential ecophysiological significance with respect to their tolerance to HL stress. The fluorometry and EPR studies of induction events in chloroplasts in situ demonstrated that acclimation of both Tradescantia species to HL conditions promoted faster responses of their PSA as compared to LL-grown plants. Acclimation of both species to HL also caused marked changes in the leaf anatomy and carotenoid composition (an increase in Violaxanthin?+?Antheraxantin?+?Zeaxanthin and Lutein pools), suggesting enhanced photoprotective capacity of the carotenoids in the plants grown in nature under high irradiance. Collectively, the results of the present work suggest that the mechanisms of long-term PSA photoprotection in Tradescantia are based predominantly on the light-induced remodeling of pigment-protein complexes in chloroplasts.  相似文献   

4.
Mesophyll structure and content of photosynthetic pigments in the leaves of three species of steppe plants, Centaurea scabiosa L., Euphorbia virgata Waldst. et Kit., Helichrysum arenarium (L.) Moench, were investigated in four geographical sites of the Volga region and the Urals located in the forest-steppe and steppe zones. Variations of the studied parameters between geographical points depended both on the species and on the structural organization of the leaf. The highest level of variation was observed for leaf area and pigment content per unit leaf area, the size and the number of chloroplasts in the cell changed to a lesser extent. The leaf thickness, leaf area and mesophyll cell sizes mostly depended on the plant species. C. scabiosa had large leaves (40–50 cm2) with large thickness (280–290 μm) and large mesophyll cells (up to 15000 μm3). The leaves of H. arenarium and E. virgata were ten times smaller and characterized by 1.5 times smaller thickness and 2?3 times smaller cell size. Geographical location and climate of the region affected leaf density, proportion of partial tissue volume, and the ratio of the photosynthetic pigments. In the southern point of Volga region with the highest climate aridity, all studied species were characterized by maximum values of volumetric leaf density (LD), due to the high proportion of sclerenchyma and vascular bundles, and specificity of the mesophyll structure. With the decline in latitude, chlorophyll (Chl) and carotenoid (Car) contents in leaf area were reduced, the ratio Chl/Car was increased, and the ratio Chl a/b was declined. The reduction of the pigment content in the leaf in all species was associated with a reduction in the amount of Chl per chloroplast, and for C. scabiosa and H. arenarium it was associated also with the reduction of chloroplast amount in the leaf area. In turn, chloroplast number per leaf area and the total cell area (Ames/A) depended on the ratio of the number and size of mesophyll cells inherent to this plant species. At the same time, we found a similar mechanism of spatial organization of leaf restructuring for all studied species—decrease in Ames/A was accompanied by increasing in the proportion of intercellular air spaces in the leaf. It is concluded that variations in structural and functional parameters of the photosynthetic apparatus of steppe plants were associated with plant adaptation to climate features. General direction of the changes of leaf parameters of the studied species with aridity was the increase of LD and the decrease of pigment content per leaf area however the cellular mechanisms of changes in the pigment content and integral parameters of mesophyll were determined by the plant species properties.  相似文献   

5.
Four plant species, Elymus mollis Trin., Carex kobomugi Ohwi, Glehnia littoralis F. Schmidt ex Miq., and Vitex rotundifolia L.f., are dominant perennial species in coastal sand dunes of Korea. We examined a physiological adaptation of these species by measurements of diurnal variation in photosynthesis and chlorophyll (Chl) fluorescence and solute patterns in leaves during one season (June), which is favorable for plant growth of all four species. All four species adopted different strategies in order to utilize radiation and to maintain water status under a fluctuating microclimate. Although the lowest water contents among four plant species was found, E. mollis with a high Chl and K+ content showed better photosynthetic performance, with high stomatal conductance (g s), net photosynthetic rate (P N), instantaneous carboxylation efficiency (CE), and water-use efficiency. Midday depression of P N in E. mollis and G. littoralis, without a reduction of gs, was associated with a reduction in CE and maximum photochemical efficiency of PSII, indicating nonstomatal limitation. Photosynthesis depression in both C. kobomugi and V. rotundifolia, with relatively low g s values, could be attributed to both stomatal and nonstomatal limitations. The high storage capacity for inorganic ions in E. molli, C. kobomugi, and G. littoralis may play an efficient role in regulating photosynthesis and maintaining leaf water status through stomatal control, and can also play an important role in osmotic adjustment.  相似文献   

6.
Frequently asked questions about chlorophyll fluorescence,the sequel   总被引:2,自引:0,他引:2  
Using chlorophyll (Chl) a fluorescence many aspects of the photosynthetic apparatus can be studied, both in vitro and, noninvasively, in vivo. Complementary techniques can help to interpret changes in the Chl a fluorescence kinetics. Kalaji et al. (Photosynth Res 122:121–158, 2014a) addressed several questions about instruments, methods and applications based on Chl a fluorescence. Here, additional Chl a fluorescence-related topics are discussed again in a question and answer format. Examples are the effect of connectivity on photochemical quenching, the correction of F V /F M values for PSI fluorescence, the energy partitioning concept, the interpretation of the complementary area, probing the donor side of PSII, the assignment of bands of 77 K fluorescence emission spectra to fluorescence emitters, the relationship between prompt and delayed fluorescence, potential problems when sampling tree canopies, the use of fluorescence parameters in QTL studies, the use of Chl a fluorescence in biosensor applications and the application of neural network approaches for the analysis of fluorescence measurements. The answers draw on knowledge from different Chl a fluorescence analysis domains, yielding in several cases new insights.  相似文献   

7.
Different pigments often occur together and affect photosynthetic characteristics of the respective leaf portions. In this study, photosynthetic activity in variegated leaves of five cultivars of the ornamental and medicinal plant, Coleus × hybridus hort., was estimated by image analysis and point data measurements of major chlorophyll (Chl) fluorescence parameters and related to the amount of photosynthetic pigments measured with a Chl meter or spectrophotometrically in leaf extracts. Significant differences in Chl and carotenoid (Car) contents were noticed among differentially pigmented sectors of a leaf and among the cultivars. Although the higher Chl concentration was noticed in purple parts compared to green parts of the leaves, the values of minimal and maximal fluorescence yield at the dark- and light-adapted state (F0, Fm, F0', Fm', respectively) were a little lower than those in the green sectors, indicating photoprotective effects provided by anthocyanins and Car, more abundant in the red parts. The lowest Chl and Car content was detected in creamy-yellow and pink sectors and this contributed to low F0, Fm, and Fm', maximal quantum yield of PSII photochemistry, and nonphotochemical and photochemical quenching but high PSII maximum efficiency and effective quantum yield of PSII photochemistry. Both methods of Chl fluorescence analysis revealed heterogeneity in capture, transfer, and dissipation of excitation energy but Chl fluorescence imaging was more suitable in examining very narrow pigmented leaf areas.  相似文献   

8.
We investigated the influence of light quality on the vulnerability of pepper plants to water deficit. For this purpose plants were cultivated either under compact fluorescence lamps (CFL) or light-emitting diodes (LED) providing similar photon fluence rates (95 µmol m?2 s?1) but distinct light quality. CFL emit a wide-band spectrum with dominant peaks in the green and red spectral region, whereas LEDs offer narrow band spectra with dominant peaks at blue (445 nm) and red (665 nm) regions. After one-week acclimation to light conditions plants were exposed to water deficit by withholding irrigation; this period was followed by a one-week regeneration period and a second water deficit cycle. In general, plants grown under CFL suffered more from water deficit than plants grown under LED modules, as indicated by the impairment of the photosynthetic efficiency of PSII, resulting in less biomass accumulation compared to respective control plants. As affected by water shortage, plants grown under CFL had a stronger decrease in the electron transport rate (ETR) and more pronounced increase in heat dissipation (NPQ). The higher amount of blue light suppressed plant growth and biomass formation, and consequently reduced the water demand of plants grown under LEDs. Moreover, pepper plants exposed to high blue light underwent adjustments at chloroplast level (e.g., higher Chl a/Chl b ratio), increasing the photosynthetic performance under the LED spectrum. Differently than expected, stomatal conductance was comparable for water-deficit and control plants in both light conditions during the stress and recovery phases, indicating only minor adjustments at the stomatal level. Our results highlight the potential of the target-use of light quality to induce structural and functional acclimations improving plant performance under stress situations.  相似文献   

9.
The present study was conducted to determine the effect of exogenous application of brassinolide (BR) on Leymus chinensis grown under shade, i.e., control (100% natural light), mild shade (70% natural light), and moderate shade (50% natural light). Shade substantially enhanced the plant growth, synthesis of photosynthetic pigments, photosynthetic efficiency, and chlorophyll (Chl) fluorescence attributes of L. chinensis as compared with control. The order of increase was mild shade > moderate shade > natural light except Chl content, where the order of increase was moderate shade > mild shade > natural light. Likewise, application of BR resulted in further exacerbation of plant height, plant fresh and dry mass, but less in case of Chl and carotenoids contents, gas-exchange characteristics, and Chl fluorescence attributes. The results conclude that shade significantly enhanced plant growth through alterations in physiological attributes of L. chinensis, while, application of BR may not further improve the plant growth under shade.  相似文献   

10.
Changes of chlorophyll (Chl) a fluorescence and photosynthetic pigment contents were analysed in galled leaves (visibly damaged and undamaged parts) and intact leaves. The values of minimal fluorescence of the dark-adapted state, maximal quantum yield of PSII photochemistry, effective quantum yield of PSII photochemical conversion, and photochemical quenching coefficient decreased in Ulmus pumila L. leaves galled by Tetraneura ulmi (L.) and in U. glabra Huds. galled by Eriosoma ulmi (L.). Colopha compressa (Koch.) feeding affected these parameters only in damaged parts of U. laevis Pall. galled leaves. The increasing number of T. ulmi galls progressively decreased photosynthetic performance. In gall tissues of all analysed aphid species, the lowest photosynthetic pigment content was found, indicating that the photosynthetic capacity must have been low in galls. Significant reduction of Chl and carotenoid contents were observed in damaged and undamaged portions of galled leaves only in the case of T. ulmi feeding.  相似文献   

11.
The native alpine plant Saussurea superba is widely distributed in Qinghai–Tibetan Plateau regions. The leaves of S. superba grow in whorled rosettes, and are horizontally oriented to maximize sunlight exposure. Experiments were conducted in an alpine Kobresia humilis meadow near Haibei Alpine Meadow Ecosystem Research Station (37°29′–37°45′N, 101°12′–101°33′E; alt. 3200 m). Leaf growth, photosynthetic pigments and chlorophyll fluorescence parameters were measured in expanding leaves of S. superba. The results indicate that leaf area increased progressively from inner younger leaves to outside fully expanded ones, and then slightly decreased in nearly senescent leaves, due to early unfavorable environmental conditions, deviating from the ordinary growth pattern. The specific leaf area decreased before leaves were fully expanded, and the leaf thickness was largest in mature leaves. There were no significant changes in the content of chlorophylls (Chl) and carotenoids (Car), but the ratios of Chl a/b and Car/Chl declined after full expansion of the leaves. The variation of Chl a/b coincided well with changes in photochemical quenching (q P) and the fraction of open PSII reaction centers (q L). The maximum quantum efficiency of PSII photochemistry after 5 min dark relaxation (F (v)/F (m)) continuously increased from younger leaves to fully mature leaves, suggesting that mature leaves could recover more quickly from photoinhibition than younger leaves. The light-harvesting capacity was relatively steady during leaf expansion, as indicated by the maximum quantum efficiency of open PSII centers (\(F_{\text{v}}^{{\prime }}\)/\(F_{\text{m}}^{{\prime }}\)). UV-absorbing compounds could effectively screen harmful solar radiation, and are a main protection way on the photosynthetic apparatus. The decline of q P and q L during maturation, together with limitation of quantum efficiency of PSII reaction centers (L (PFD)), shows a decrease of oxidation state of QA in PSII reaction centers under natural sunlight. Furthermore, light-induced (Φ NPQ) and non-light-induced quenching (Φ NO) were consistent with variation of L (PFD). It is concluded that the leaves of S. superba could be classified into four functional groups: young, fully expanded, mature, and senescent. Quick recovery from photoinhibition was correlated with protection by screening pigments, and high level of light energy trapping was correlated with preservation of photosynthetic pigments. Increasing of Φ NPQ and Φ NO during leaves maturation indicates that both thermal dissipation of excessive excitation energy in safety and potential threat to photosynthetic apparatus were strengthened due to the declination of q P and q L, and enhancement of L (PFD).  相似文献   

12.
Chlorophyll (Chl) a fluorescence induction (transient), measured by exposing dark-adapted samples to high light, shows a polyphasic rise, which has been the subject of extensive research over several decades. Several Chl fluorescence parameters based on this transient have been defined, the most widely used being the FV [= (FM–F0)]/FM ratio as a proxy for the maximum quantum yield of PSII photochemistry. However, considerable additional information may be derived from analysis of the shape of the fluorescence transient. In fact, several performance indices (PIs) have been defined, which are suggested to provide information on the structure and function of PSII, as well as on the efficiencies of specific electron transport reactions in the thylakoid membrane. Further, these PIs have been proposed to quantify plant tolerance to stress, such as by high light, drought, high (or low) temperature, or N-deficiency. This is an interesting idea, since the speed of the Chl a fluorescence transient measurement (<1 s) is very suitable for high-throughput phenotyping. In this review, we describe how PIs have been used in the assessment of photosynthetic tolerance to various abiotic stress factors. We synthesize these findings and draw conclusions on the suitability of several PIs in assessing stress responses. Finally, we highlight an alternative method to extract information from fluorescence transients, the Integrated Biomarker Response. This method has been developed to define multi-parametric indices in other scientific fields (e.g., ecology), and may be used to combine Chl a fluorescence data with other proxies characterizing CO2 assimilation, or even growth or grain yield, allowing a more holistic assessment of plant performance.  相似文献   

13.
B. He  M. Gu  X. Wang  X. He 《Photosynthetica》2018,56(4):1147-1153
The photosynthesis was investigated 30 d after Pb treatment in Myrica rubra seedlings. The Pb treatment resulted in significantly increased Pb concentrations in shoots. Low Pb concentration exposure (≤2 mM) reduced the net photosynthetic rate (PN), transpiration rate (E), and stomatal conductance (gs) without affecting the intercellular CO2 concentration (Ci), chlorophyll (Chl) content, and Chl fluorescence parameters. At 10 d after severe Pb treatment (≥4 mM), PN was inhibited and accompanied by Chl damage, while at 30 d, the inhibition of PN was followed by an increase of Ci and a decrease of gs, E, Chl content, and Chl fluorescence parameters. M. rubra showed a promising prospect for use in the soil phytoremediation, when Pb concentration is low, but the remediation efficiency of M. rubra is limited if Pb exceeds 2 mM.  相似文献   

14.
Optical characteristics, contents of photosynthetic pigments, total soluble sugars, and starch, rates of gas exchange, chlorophyll (Chl) a fluorescence, and leaf water relations were analysed in three Vitis vinifera L. cultivars, Tinto Cão (TC), Touriga Nacional (TN), and Tinta Roriz (TR), grown in Mediterranean climate. Chl content was significantly lower in TC than in TN and TR leaves, while the Chl a/b ratio was higher. TR had the lowest net photosynthetic rate, stomatal conductance, and contents of soluble sugars and starch than TN and TC. In spite of low Chl content, TC showed the lowest photon absorbance and the highest photochemical efficiency of photosystem 2. TC had the lowest predawn and midday leaf water potential. The capability for osmotic adjustment was similar among cultivars and the calculated modulus of elasticity was higher in TC leaves. The typical lighter green leaves of TC seemed to be an adaptive strategy to high irradiance and air temperature associated to water stress.  相似文献   

15.
Drought stress has multiple effects on the photosynthetic apparatus. Herein, we aimed to study the effect of drought stress on fluorescence characteristics of PSII in leaves of Plectranthus scutellarioides and explore potentially underlying mechanisms. Plants of P. scutellarioides were grown in a greenhouse and subjected to drought (DS, drought-stressed) or daily irrigation (control group). Leaf chlorophyll (Chl) index and induction kinetics curves of Chl a fluorescence and the JIP-test were used to evaluate effects of drought lasting for 20 d. Our results showed that both the leaf and soil relative water content decreased with increasing treatment duration. The leaf Chl index was reduced to half in the DS plants compared with the control group after 20 d. The minimal fluorescence in the DS plants was higher than that in the control plants after 10 d of the treatment. Maximum photochemical efficiency and lateral reactivity decreased with increasing treatment duration in the DS plants. With the continuing treatment, values of absorption flux per reaction center (RC), trapped energy flux per RC, dissipated energy flux per RC, and electron transport flux per RC increased in the earlier stage in the DS plants, while obviously decreased at the later stage of the treatment. In conclusion, drought stress inhibited the electron transport and reduced PSII photochemical activity in leaves of P. scutellarioides.  相似文献   

16.
Cistus salvifolius L. is the most widely spread Cistus species around the Mediterranean basin. It colonizes a wide range of habitats growing from sea level to 1,800 m a.s.l., on silicolous and calcicolous soils, in sun areas as well as in the understory of wooded areas. Nevertheless, this species has been mainly investigated in term of its responsiveness to drought. Our aim was to understand which leaf traits allow C. salvifolius to cope with low-light environments. We questioned if biochemical and physiological leaf trait variations in response to a reduced photosynthetic photon flux density were related to leaf morphological plasticity, expressed by variations of specific leaf area (SLA) and its anatomical components (leaf tissue density and thickness). C. salvifolius shrubs growing along the Latium coast (41°43'N,12°18'E, 14 m a.s.l., Italy) in the open and in the understory of a Pinus pinea forest, were selected and the relationships between anatomical, gas exchange, chlorophyll (Chl) fluorescence, and biochemical parameters with SLA and PPFD variations were tested. The obtained results suggested long-term acclimation of the selected shrubs to contrasting light environments. In high-light conditions, leaf nitrogen and Chl contents per leaf area unit, leaf thickness, and Chl a/b ratio increased, thus maximizing net photosynthesis, while in shade photosynthesis, it was downregulated by a significant reduction in the electron transport rate. Nevertheless, the increased pigment-protein complexes and the decreased Chl a/b in shade drove to an increased light-harvesting capacity (i.e. higher actual quantum efficiency of PSII). Moreover, the measured vitality index highlighted the photosynthetic acclimation of C. salvifolius to contrasting light environments. Overall, our results demonstrated the morphological, anatomical, and physiological acclimation of C. salvifolius to a reduced light environment.  相似文献   

17.
Pigments absorbing 350–1,050 nm radiation have had an important role on the Earth for at least 3.5 billion years. The ion pumping rhodopsins absorb blue and green photons using retinal and pump ions across cell membranes. Bacteriochlorophylls (BChl), absorbing in the violet/blue and near infra red (NIR), power anoxygenic photosynthesis, with one photoreaction centre; and chlorophylls (Chl), absorbing in the violet/blue and red (occasionally NIR) power oxygenic photosynthesis, with two photoreaction centres. The accessory (bacterio)chlorophylls add to the spectral range (bandwidth) of photon absorption, e.g., in algae living at depth in clear oceanic water and in algae and photosynthetic (PS) bacteria in microbial mats. Organism size, via the package effect, determines the photon absorption benefit of the costs of synthesis of the pigment–protein complexes. There are unresolved issues as to the evolution of Chls vs. BChls and the role of violet/blue and NIR radiation in PS bacteria.  相似文献   

18.
Chlorophyll (Chl) deficiency in leaves of a plastome sunflower (Helianthus annuus L.) en:chlorina-5 mutant is due to the formation of smaller chloroplasts with a markedly reduced membrane system, as compared to the parent 3629 line. Abnormalities in the structure of the photosynthetic apparatus in the mutant can be mainly attributed to changes in the formation of photosystem I and its light-harvesting complexes. Chl deficiency in en:chlorina-5 correlated with its lower capability of synthesizing the first specific Chl precursor, 5-aminolevulinic acid (ALA) in the light. Light-independent stages of Chl biosynthesis in the mutant had the same efficiency as in leaves of the parent line. ALA formation in darkness and its conversion into protochlorophyllide did not depend on the extent of photosynthetic membrane development and photosynthetic activity.  相似文献   

19.
In the present study, photosynthetic traits and chlorophyll (Chl) fluorescence parameters of Populus sibirica grown under different irrigation regimes were investigated to estimate seedling growth and vitality for reforestation of a desertification area. According to our results, photosynthesis and Chl fluorescence were significantly affected by water deficit only under severe drought conditions.  相似文献   

20.
Wild progenitors of common wheat are a potential source of tolerance to biotic and abiotic stresses. We conducted a glasshouse pot experiment to study genotypic differences in response to drought stress in a collection of 180 accessions of Aegilops and Triticum along with one tolerant and one sensitive control variety. Several physiological traits and chlorophyll fluorescence parameters were evaluated. Our findings indicated that drought significantly reduced shoot fresh (59.45%) and dry (50.83%) weights, stomatal conductance (41.52%) and maximum photosynthetic capacity (41.06%), but increased initial fluorescence (28.10%). Drought stress also decreased the chlorophyll content, relative water content and maximum quantum efficiency by 14.90, 12.13 and 11.42%, respectively. Principal component analysis of the 182 individuals identified three components that explained 57.61 and 61.68% of the total variation in physiological and photosynthetic traits under control and stress conditions, respectively. When grouped into the 12 species tested, the three top components explained 78.22% of the total variation under drought. The means comparison, stress tolerance index and biplot analysis identified five accessions with superior tolerance to drought. Remarkably, four species of wild relatives—Ae. cylindrica (DC genome), Ae. crassa (DM genome), Ae. caudata (C genome) and T. urartu (Au genome)—responded well to drought stress with a lower percentage decline for most traits and high values for the first two components. The potential of these species offers further opportunities for analysis at the molecular and cellular levels to confront with drought stress through a physiological mechanism.  相似文献   

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