干旱胁迫下油橄榄品种光合特性研究

为揭示油橄榄(Olea europaea L.)耐旱性与光合特性之间的关系,以筛选出的适宜于半干旱川西南地区种植的7个引进油橄榄品种为供试材料,采用盆栽模拟干旱胁迫的方法,研究持续干旱胁迫对其光合特性的影响。结果表明:(1)随着干旱胁迫程度加剧,7个油橄榄品种叶片相对含水量均显著降低,至干旱胁迫后期(25d),各品种叶片均出现大幅失水,其中品种‘科拉蒂’失水率最高(45.79%),而品种‘小苹果’失水率最低(25.52%),说明‘小苹果’叶片在干旱胁迫下较其他油橄榄品种具有更高保水能力。(2)随着干旱胁迫程度加剧,7个油橄榄品种叶片光合色素含量均不同程度降低,表明光合色素分解量大于合成量;干旱胁迫持续25d时,品种‘豆果’的叶绿素a和叶绿素b含量下降幅度最大(P0.05),品种‘皮削利’类胡萝卜素含量下降幅度最大(P0.05),而品种‘小苹果’叶绿素a含量下降幅度最小。(3)随着干旱胁迫的持续进行,各油橄榄品种叶片净光合速率(Pn)、蒸腾速率(Tr)、气孔导度(Gs)和胞间CO2浓度(Ci)均不同程度降低,而水分利用效率(WUE)则呈上升趋势;干旱胁迫期间,品种‘佛奥’的Pn、Tr和Ci以及‘皮削利’的Gs降幅均高于其他品种,而‘小苹果’的Pn、Gs和Ci降幅均为最小且WUE上升幅度最大。研究发现,在持续干旱胁迫条件下,油橄榄幼苗叶片均大幅失水,光合色素结构被破坏、色素分解、含量降低,同时气孔关闭蒸发减少,光合作用减弱,而供试油橄榄品种中‘小苹果’对干旱胁迫的适应性最强,适宜于在半干旱的川西地区种植。     

Differences in abscisic acid concentration in roots and leaves of two young Olive (<Emphasis Type="Italic">Olea europaea</Emphasis> L.) cultivars in response to water deficit

Differences in abscisic acid (ABA) accumulation between two olive cultivars were studied by enzyme-linked immunosorbent assay in roots and leaves, leaf water potential (Ψ_l), stomatal conductance (g _s) as well as photosynthetic rate (A) were also determined in well-watered (WW) and water-stressed (WS) plants of two olive cultivars ‘Chemlali’ and ‘Chetoui’. ‘Chemlali’ was able to maintain higher leaf CO₂ assimilation rate and leaf stomatal conductance throughout the drought cycle when compared with ‘Chetoui’. Furthermore, leaf water potential of ‘Chemlali’ decreased in lower extent than in Chetoui in response to water deficit. Interestingly, significant differences in water-stress-induced ABA accumulation were observed between the two olive cultivars and reflect the degree of stress experienced. Chemlali, a drought tolerant cultivar, accumulated lower levels of ABA in their leaves to regulate stomatal control in response to water stress compared to the drought sensitive olive cultivar ‘Chetoui’ which accumulated ABA in large amount.     

Drought-induced oxidative damage and antioxidant responses in peanut (<Emphasis Type="Italic">Arachis</Emphasis><Emphasis Type="Italic">hypogaea</Emphasis> L.) seedlings

Two cultivars of peanut (Arachis hypogaea L.) which were designated as resistant (Florispan) and sensitive (Gazipasa) according to their growth retardation under drought stress conditions were compared for their oxidative damage and antioxidant responses. Sixteen days-old peanut seedlings were subjected to PEG-6000 solutions of two different osmotic potentials; −0.4 and −0.8 MPa, and various growth parameters, photosystem II activity, changes in malondialdehyde (MDA), hydrogen peroxide (H₂O₂) and proline levels, activities of ascorbate peroxidase (APX), catalase (CAT), peroxidase (POX) and gluthatione reductase (GR) enzymes were determined. Both cultivars exhibited water deficit at −0.8 MPa osmotic potential of PEG-6000 and H₂O₂ levels significantly increased during exposure to −0.4 MPa osmotic potential. However, H₂O₂ levels were under control in both cultivars at exposure to −0.8 MPa osmotic potential. Significant proline accumulation was observed in the tissues of cv. Florispan at −0.8 MPa osmotic potential, whereas proline accumulation did not appear to be an essential part of the protection mechanism against drought in cv. Gazipasa. No significant variation in chlorophyll fluorescence values were detected in neither of the cultivars. Enzyme activity measurements revealed that Gazipasa copes well with lesser magnitudes of drought stress by increasing the activity of mainly APX, and during harsh stress conditions, only APX maintains its activity in the tissues. In cultivar Florispan, GR activity appears to take role in lesser magnitudes of drought stress, whereas CAT and APX activities appear to be very crucial antioxidative defenses during intense stress conditions. The results indicate that, the level of proline and activities of the enzymes CAT and APX are important mechanisms for the maintenance of drought tolerance in peanut plants.     

Physiological and anatomical changes induced by drought in two olive cultivars (cv Zalmati and Chemlali)

Photosynthetic gas exchange, vegetative growth, water relations and fluorescence parameters as well as leaf anatomical characteristics were investigated on young plants of two Olea europaea L. cultivars (Chemlali and Zalmati), submitted to contrasting water availability regimes. Two-year-old olive trees, grown in pots in greenhouse, were not watered for 2 months. Relative growth rate (RGR), leaf water potential (Ψ_LW) and the leaf relative water content (LWC) of the two cultivars decreased with increasing water stress. Zalmati showed higher values of RGR and LWC and lower decreased values of Ψ_LW than Chemlali, in response to water deficit, particularly during severe drought stress. Water stress also caused a marked decline on photosynthetic capacity and chlorophyll fluorescence. The net photosynthetic rate, stomatal conductance, transpiration rate, the maximal photochemical efficiency of PSII (F _v/F _m) and the intrinsic efficiency of open PSII reaction centres (F′ _v/F′ _m) decreased as drought stress developed. In addition, drought conditions, reduced leaf chlorophyll and carotenoids contents especially at severe water stress. However, Zalmati plants were the less affected when compared with Chemlali. In both cultivars, stomatal control was the major factor affecting photosynthesis under moderate drought stress. At severe drought-stress levels, the non-stomatal component of photosynthesis is inhibited and inactivation of the photosystem II occurs. Leaf anatomical parameters show that drought stress resulted in an increase of the upper epidermis and palisade mesophyll thickness as well as an increase of the stomata and trichomes density. These changes were more characteristic in cv. ‘Zalmati’. Zalmati leaves also revealed lower specific leaf area and had higher density of foliar tissue. From the behaviour of Zalmati plants, with a smaller reduction in relative growth rate, net assimilation rate and chlorophyll fluorescence parameters, and with a thicker palisade parenchyma, and a higher stomatal and trichome density, we consider this cultivar more drought-tolerant than cv. Chemlali and therefore, very promising for cultivation in arid areas.     

The relations between antioxidant enzymes and chlorophyll fluorescence parameters in common bean cultivars differing in sensitivity to drought stress

Effects of the antioxidant system and chlorophyll fluorescence on drought tolerance of four common bean (Phaseolus vulgaris L.) cultivars were studied. The cultivars were positioned in the order of a decrease in their drought tolerance: Yakutiye, Pinto Villa, Ozayse, and Zulbiye on the basis of changes in the water potential, stomatal conductance, photosynthetic pigment content, and lipid peroxidation. Under drought conditions, the level of H₂O₂ was not changed in cv. Pinto Villa but decreased in other cultivars. Antioxidant enzymes (superothide dismutase (SOD), guaiacol peroxidase (GPX), catalase (CAT), ascorbate peroxidase (APX), and glutathione reductase (GR)) were generally activated in all cultivars. Interestingly, CAT, APX, and GR activities were not changed in cv. Pinto Villa, APX activity decreased in cv. Yakutiye, and CAT activity was not changed in cv. Zulbiye. The increases in SOD and GPX activities in cv. Ozayse were higher than in other cultivars. Drought stress reduced the effective quantum yield of PS2 (Φ_PS2) and the photochemical quenching (q_p), while it increased nonphotochemical quenching (NPQ) in all cultivars. The reduction or increase was more pronounced in cv. Zulbiye. There were generally significant correlations between q_p, NPQ, and ROS scavenging by SOD and APX. Also, there were significant correlations between SOD and q_p in tolerant cultivars and APX and q_p in sensitive ones. The results indicate that activation of SOD and APX was closely related to the efficiency of PS2 in common bean cultivars. This interaction was essential for protection of photosystems and plant survival under drought.     

Changes in lipid composition, water relations and gas exchange in leaves of two young ‘Chemlali’ and ‘Chetoui’ olive trees in response to water stress

The comparative responses of two young olive trees (Olea europaea L. ‘Chemlali’ and ‘Chetoui’) to drought stress were investigated during 1 month. Three-month-old own-rooted plants were subjected to two irrigation treatments: WW (well watered plants that were irrigated with fresh water to maintain a soil water content close to field capacity), and WS (water stressed plants by withholding water). Leaf water potential, gas exchange and leaf lipid composition were studied. ‘Chemlali’ was able to maintain higher leaf CO₂ assimilation rate and leaf stomatal conductance throughout the drought cycle compared to ‘Chetoui’. Water stress induced a larger decrease in the total lipid content in ‘Chetoui’ than in ‘Chemlali’. Interestingly, the constitution of different lipid classes was highly altered in ‘Chetoui’. Lipid changes in Chemlali, a drought tolerant cultivar, revealed more stability of its cellular membranes to drought stress as compared to the drought susceptible olive cultivar, Chétoui. Furthermore, in comparison to the controls, drought stressed plants showed an increase in the degree of unsaturation of leaf lipids in the two olive cultivars. Moreover, the results observed in Chemlali showed that besides changes in lipids composition this cultivar may have an efficient defence strategy which can be related on antioxidative production against oxidative stress.     

Physiological responses of different olive genotypes to drought conditions

Gas exchange rates, chlorophyll fluorescence, pressure–volume relationships, photosynthetic pigments, total soluble sugars, starch, soluble proteins and proline concentrations were investigated in five Olea europaea L. cultivars with different geographical origins (Arbequina, Blanqueta, Cobran?osa, Manzanilla and Negrinha) grown under Mediterranean field conditions. We found considerable genotypic differences among the cultivars. Comparing the diurnal gas exchange rates, we observed that Cobran?osa, Manzanilla and Negrinha had high photosynthetic rate than Arbequina and Blanqueta. The first group reveals to be better acclimated to drought conditions, and appears to employ a prodigal water-use strategy, whereas Blanqueta and Arbequina, with high water-use efficiency, appear to employ a conservative water-use strategy. The degree of midday depression in photosynthesis was genotype dependent, with a maximum in Arbequina and a minimum in Negrinha. The reductions in the photosynthetic rate were dependent from both stomatal and non-stomatal limitations. Elastic adjustment plays an important role as drought tolerance mechanism. The group of cultivars that employ a prodigal water-use strategy revealed high tissue elasticity, whereas Arbequina and Blanqueta revealed high tissue rigidity. We also identified the existence of drought tolerance mechanisms associated with soluble proteins accumulation in the foliage. The high levels of soluble proteins in Arbequina may represent an increased activity of oxidative stress defence enzymes and may also represent a reserve for post stress recovery. In all cultivars, especially in Manzanilla, free proline was accumulated in the foliage. The discussed aspects of drought stress metabolism may have an adaptative meaning, supporting the hypothesis that olive cultivars native to dry regions, such as Cobran?osa, Manzanilla and Negrinha, have more capability to acclimate to drought conditions than cultivars originated in regions with a more temperate climate, like Arbequina and Blanqueta.     

Ionic and Osmotic Effects of Salinity on Single-Leaf Photosynthesis in Two Wheat Cultivars with Different Drought Tolerance

The effects of iso-osmotic salinity and drought stresses on leaf net photosynthetic rate (P _N) in two wheat (Triticum aestivum L.) cultivars BR 8 and Norin 61, differing in drought tolerance, were compared. In drought-sensitive Norin 61, the decline of P _N was larger than that in drought-tolerant BR 8. Under NaCl treatment, P _N decreased in two phases similarly in both cultivars. In the first phase, photosynthetic depression was gradual without any photochemical changes. In the second phase, photosynthetic depression was rapid and accompanied with a decline of the energy conversion efficiency in photosystem 2 (_PS2). Our observations suggest that the osmotic factor may induce a gradual depression of photosynthesis due to stomatal closure under both stress treatments. However, under NaCl treatment, a ionic factor (uptake and accumulation of excess Na⁺) may have direct effects on electron transport and cause more severe photosynthetic depression. The drought tolerance mechanism of BR 8 was insufficient to maintain single-leaf photosynthesis under salinity.     

Differential fungal colonization and physiological defense responses of new olive cultivars infected by the necrotrophic fungus <Emphasis Type="Italic">Verticillium dahliae</Emphasis>

New olive cultivars adapted to Tunisia’s growing conditions were examined for their resistance to verticillium wilt (VWO) to determine whether differences in susceptibility among currently grown cultivars might contribute to the management of this disease. Based on the evaluation of 14 cultivars, 10 were classified as susceptible or extremely susceptible (Chetoui, Chemlali, Rkhami, Jarboui, Zalmati, Jarboui, Oueslati, Manzanille, Picholine and Frangivento), 2 as moderately susceptible (Koroneiki and Coratina), and 2 as resistant (Meski and Sayali) to VWO. Three cultivars with different susceptibility levels were selected to examine the levels of hydrogen peroxide (H₂O₂), soluble sugars (SS), soluble proteins (SP), total polyphenols (TP), lipid peroxidation, activities of antioxidant enzymes, and fungal biomass in planta. V. dahliae DNA occurred early in the roots at 15 dpi and reached a maximum of 3.507 and 2.52 ng/100 ng of plant DNA, respectively, in the extremely susceptible and resistant cultivars. Fungal DNA in the stems occurred at 30 dpi and increased slowly to reach a maximum of 0.23 ng/100 ng of total DNA in the extremely susceptible cultivars. We showed that the amount of fungal DNA in planta was roughly correlated with the susceptibility to VWO (P < 0.0001; r = 0.95). The comparison of cultivars at the physiological level indicated that olive resistance is roughly correlated with the antioxidant enzymes activity, H₂O₂ concentration, and TP and SP contents. The results of this study open new perspectives for olive genetic improvement programs aiming at developing new cultivars resistant to this wilt.     

Effects of cadmium stress upon activities of antioxidative enzymes, photosynthetic rate, and production of phytochelatins in leaves and chloroplasts of wheat cultivars differing in yield potential

We tested the mode of action of Cd on photosynthesis and activities of ATP-sulfurylase (ATP-S), catalase (CAT), superoxide dismutase (SOD), ascorbate peroxidase (APX), glutathione reductase (GR), and on contents of phytochelatins (PCs) and glutathione (GSH) in two cultivars of wheat (Triticum aestivum L.) PBW-343 and WH-542 differing in yield potential. Cd treatment increased Cd content and photosynthetic activity in PBW-343 more than in WH-542. The activities of APX, GR, ATP-S, and synthesis of PCs and GSH were also increased by Cd, but the CAT and SOD activities were inhibited in both the cultivars. The efficient functioning of antioxidative enzymes, production of PCs and GSH, helped in counteracting the effects of Cd namely in PBW-343, protected photosynthetic ability, and increased the tolerance to Cd.     

Overexpressing <Emphasis Type="Italic">SgNCED1</Emphasis> in Tobacco Increases ABA Level,Antioxidant Enzyme Activities,and Stress Tolerance

Abscisic acid (ABA) regulates plant adaptive responses to various environmental stresses. 9-cis-epoxycarotenoid dioxygenase (NCED) is the key enzyme of ABA biosynthesis in higher plants. A NCED gene, SgNCED1, was overexpressed in transgenic tobacco plants which resulted in 51–77% more accumulation of ABA in leaves. Transgenic tobacco plants decreased stomatal conductance, transpiration rate, and photosynthetic rate but induced activities of superoxide dismutase (SOD), catalase (CAT), and ascorbate-peroxidase (APX). Hydrogen peroxide (H₂O₂) and nitric oxide (NO) in leaves were also induced in the transgenic plants. Compared to the wild-type control, the transgenic plants improved growth under 0.1 M mannitol-induced drought stress and 0.1 M NaCl-induced salinity stress. It is suggested that the ABA-induced H₂O₂ and NO generation upregulates the stomatal closure and antioxidant enzymes, and therefore increases drought and salinity tolerance in the transgenic plants.     

Different mechanisms of photosynthetic response to drought stress in tomato and violet orychophragmus

Carbonic anhydrase (CA) catalyzes reversible hydration of CO₂ and it can compensate for the lack of H₂O and CO₂ in plants under stress conditions. Antioxidative enzymes play a key role in scavenging reactive oxygen species and in protecting plant cells against toxic effects. Tomato represents a stress-sensitive plant while violet orychophragmus belongs to adversity-resistant plants. In order to study the drought responses in tomato and violet orychophragmus plants, CA and antioxidative enzyme activities, photosynthetic capacity, and water potential were determined in plants under drought stress. We found that there were similar change trends in CA activity and drought tolerance in violet orychophragmus, and in antioxidative enzymes and drought tolerance in tomato plants. Basic mechanisms of drought resistance should be identified for understanding of breeding measures in plants under stress conditions.     

Responses of two field-grown coffee species to drought and re-hydration

The gas exchange, parameters of chlorophyll fluorescence, contents of pigments, and activities of superoxide dismutase (SOD), ascorbate peroxidase (APX), as well as lipid peroxidation were investigated in two field-grown coffee species, Coffea arabica and C. liberica, exposed to drought and re-hydration. Drought caused a more pronounced inhibition of net photosynthetic rate in C. liberica compared to C. arabica. The de-epoxidation of xanthophyll cycle pigments at midday estimated by leaf reflectance was much higher in C. arabica than in C. liberica, but no significant change was found in response to drought. Under moderate drought, the activities of SOD and APX increased significantly only in C. arabica. The maximum photochemical efficiency of photosystem 2, PS2 (F_v/F_m) at predawn did not change and there was no lipid peroxidation during this time. Under severe drought F_v/F_m decreased and initial fluorescence (F₀) increased for both species, and SOD activity increased, APX activity remained relatively high, and malondialdehyde (MDA) accumulated in C. arabica, while APX decreased in C. liberica. The photosynthetic apparatus of C. arabica was completely recovered after 5 d of re-irrigation as indicated by the restoration of F_v/F_m to the control values. A lack of recovery upon rewatering of C. liberica indicated irreversible damage to PS2. Hence compared to C. liberica, C. arabica possesses a higher desiccation-induced antioxidative protection and higher portion of the total pigment pool used in photoprotection, which might aid alleviating photoinhibitory damage during desiccation and photosynthesis recovery when favourable conditions are restored.The research was financially supported by the project of the Chinese Academy of Sciences (KSCX2-SW-104).     

Photosynthesis, respiration and antioxidant enzymes in pepper leaves under drought and heat stresses

Leaf gas exchange, chlorophyll fluorescence and activities of antioxidant enzymes were studied in two pepper (Capsicum annuum L.) cultivars grown under drought (D) and heat (H), as well as under drought and heat in combination (HD). The drought-tolerant cv. Zhengjiao 13 exhibited greater net photosynthetic rate (P_N) and cytochrome respiratory pathway activity (R_SHAM), and lower contents of superoxide radical and hydrogen peroxide, as compared to the drought-sensitive cv. Longkouzaojiao. The P_N and R_SHAM decreased and ROS production increased under D and HD in both cultivars. As compared to the Longkouzaojiao, Zhengjiao 13 retained higher non-photochemical quenching (NPQ), photorespiration rate (R_L), and alternative respiratory pathways (R_KCN) under D and HD. Drought increased the superoxide dismutase (SOD) and ascorbate peroxidase (APX) activities in the cytosol, chloroplasts and mitochondria in the two cultivars. Although SOD and APX activities decreased in Longkouzaojiao under HD, SOD activity increased in Zhengjiao 13. There was no H-induced reactive oxygen species production due to increase of R_L, NPQ, R_SHAM, R_KCN and activities of SOD and APX. However, H slightly decreased the P_N. The results indicated that HD was more detrimental than either stress alone.     

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.     

Drought tolerance of photosynthetic electron transport under CO2‐enriched and normal air in cereal species

The quantum yield of photosynthetic electron transport (ΦPSII), evaluated by means of chlorophyll (Chl) fluorescence analysis, has proven to be a useful screening test for drought tolerance in durum wheat (Triticum durum Desf.). To explore the potential of this parameter further in detecting drought-tolerant genotypes, three cereal species were studied; ΦPSII measurements were carried out under two different gas mixtures, at three points of the induction curve (to obtain the maximal ΦPSII and both the transient and steady-state actual ΦPSII), and at three different water stress levels (moderate, severe and drastic). The species investigated were durum and bread wheat (Triticum aestivum L.) and barley (Hordeum vulgare L.); two cultivars per species, characterized by different levels of drought tolerance, were tested. The two gas mixtures used were normal air (21% O₂, 0.035% CO₂ in N₂) to monitor the whole photosynthetic process under physiological conditions, and CO₂ enriched-low O₂ air (1% O₂, 5% CO₂ in N₂) to monitor ΦPSII reduction under stress mainly related to Calvin cycle activity. When ΦPSII related to both assimilatory and non-assimilatory metabolism was evaluated, the cultivar differences observed under normal Air were more representative of the agronomic performance upon drought stress than under high CO₂-low O₂ air. Maximal ΦPSII showed no difference among either cultivars, gas mixtures or stress levels, the efficiency of excitation capture being highly resistant to drought. The ΦPSII evaluated during the transient yielded predictable values in respect of drought tolerance for durum wheat and barley cultivars, highlighting the key role of regulatory processes such as the Mehler peroxidase reaction and possibly also cyclic electron transport, in preventing overreduction under stress. The results clearly show that when Chl fluorescence analysis is used as a parameter in plant breeding, different experimental conditions should be used depending on the physiological mechanism that is bred or selected for.     

Physiochemical and antioxidant responses of the perennial xerophyte Capparis ovata Desf. to drought

Caper (Capparis ovata Desf.) is a perennial shrub (xerophyte) and drought resistant plant which is well adapted to Mediterranean Ecosystem. In the present study we investigated the plant growth, relative water content (RWC), chlorophyll fluorescence (F_V/F_M), lipid peroxidation (TBA-reactive substances content) as parameters indicative of oxidative stress and antioxidant enzymes such as superoxide dismutase (SOD), ascorbate peroxidase (APX), peroxidase (POX), catalase (CAT) and glutathione reductase (GR) in relation to the tolerance to polyethylene glycol mediated drought stress in C. ovata seedlings. For induction of drought stress, the 35 days seedlings were subjected to PEG 6000 of osmotic potential −0.81 MPa for 14 days. Lipid peroxidation increased in PEG stressed seedlings as compared to non-stressed seedlings of C. ovata during the experimental period. With regard to vegetative growth, PEG treatment caused decrease in shoot fresh and dry weights, RWC and F_V/F_M but decline was more prominent on day 14 of PEG treatment. Total activity of antioxidative enzymes SOD, APX, POX, CAT and GR were investigated in C. ovata seedlings under PEG mediated drought. Induced activities of SOD, CAT and POX enzymes were high and the rate of increment was higher in stressed seedling. APX activity increased on both days of PEG treatment, however, increase in GR activity was highest on day 14 of drought stress. We concluded that increased drought tolerance of C. ovata is correlated with diminishing oxidative injury by functioning of antioxidant system at higher rates under drought stress.     

Cassava biology and physiology

Cassava or manioc (Manihot esculenta Crantz), a perennial shrub of the New World, currently is the sixth world food crop for more than 500 million people in tropical and sub-tropical Africa, Asia and Latin America. It is cultivated mainly by resource-limited small farmers for its starchy roots, which are used as human food either fresh when low in cyanogens or in many processed forms and products, mostly starch, flour, and for animal feed. Because of its inherent tolerance to stressful environments, where other food crops would fail, it is often considered a food-security source against famine, requiring minimal care. Under optimal environmental conditions, it compares favorably in production of energy with most other major staple food crops due to its high yield potential. Recent research at the Centro Internacional de Agricultura Tropical (CIAT) in Colombia has demonstrated the ability of cassava to assimilate carbon at very high rates under high levels of humidity, temperature and solar radiation, which correlates with productivity across all environments whether dry or humid. When grown on very poor soils under prolonged drought for more than 6 months, the crop reduce both its leaf canopy and transpiration water loss, but its attached leaves remain photosynthetically active, though at greatly reduced rates. The main physiological mechanism underlying such a remarkable tolerance to drought was rapid stomatal closure under both atmospheric and edaphic water stress, protecting the leaf against dehydration while the plant depletes available soil water slowly during long dry periods. This drought tolerance mechanism leads to high crop water use efficiency values. Although the cassava fine root system is sparse, compared to other crops, it can penetrate below 2 m soil, thus enabling the crop to exploit deep water if available. Leaves of cassava and wildManihotpossess elevated activities of the C₄ enzyme PEP carboxylase but lack the leaf Kranz anatomy typical of C₄ species, pointing to the need for further research on cultivated and wild Manihot to further improve its photosynthetic potential and yield, particularly under stressful environments. Moreover, a wide range in values of K _m (CO₂) for the C₃ photosynthetic enzyme Rubisco was found among cassava cultivars indicating the possibility of selection for higher affinity to CO₂, and consequently higher leaf photosynthesis. Several plant traits that may be of value in crop breeding and improvement have been identified, such as an extensive fine root system, long leaf life, strong root sink and high leaf photosynthesis. Selection of parental materials for tolerance to drought and infertile soils under representative field conditions have resulted in developing improved cultivars that have high yields in favorable environments while producing reasonable and stable yields under stress.     

Cassava biology and physiology

Cassava or manioc (Manihot esculenta Crantz), a perennial shrub of the New World, currently is the sixth world food crop for more than 500 million people in tropical and sub-tropical Africa, Asia and Latin America. It is cultivated mainly by resource-limited small farmers for its starchy roots, which are used as human food either fresh when low in cyanogens or in many processed forms and products, mostly starch, flour, and for animal feed. Because of its inherent tolerance to stressful environments, where other food crops would fail, it is often considered a food-security source against famine, requiring minimal care. Under optimal environmental conditions, it compares favorably in production of energy with most other major staple food crops due to its high yield potential. Recent research at the Centro Internacional de Agricultura Tropical (CIAT) in Colombia has demonstrated the ability of cassava to assimilate carbon at very high rates under high levels of humidity, temperature and solar radiation, which correlates with productivity across all environments whether dry or humid. When grown on very poor soils under prolonged drought for more than 6 months, the crop reduce both its leaf canopy and transpiration water loss, but its attached leaves remain photosynthetically active, though at greatly reduced rates. The main physiological mechanism underlying such a remarkable tolerance to drought was rapid stomatal closure under both atmospheric and edaphic water stress, protecting the leaf against dehydration while the plant depletes available soil water slowly during long dry periods. This drought tolerance mechanism leads to high crop water use efficiency values. Although the cassava fine root system is sparse, compared to other crops, it can penetrate below 2 m soil, thus enabling the crop to exploit deep water if available. Leaves of cassava and wild Manihot possess elevated activities of the C₄ enzyme PEP carboxylase but lack the leaf Kranz anatomy typical of C₄ species, pointing to the need for further research on cultivated and wild Manihot to further improve its photosynthetic potential and yield, particularly under stressful environments. Moreover, a wide range in values of K _m (CO₂) for the C₃ photosynthetic enzyme Rubisco was found among cassava cultivars indicating the possibility of selection for higher affinity to CO₂, and consequently higher leaf photosynthesis. Several plant traits that may be of value in crop breeding and improvement have been identified, such as an extensive fine root system, long leaf life, strong root sink and high leaf photosynthesis. Selection of parental materials for tolerance to drought and infertile soils under representative field conditions have resulted in developing improved cultivars that have high yields in favorable environments while producing reasonable and stable yields under stress.     

An integrated diagnostic approach to understand drought tolerance in mulberry (Morus indica L.)

Four popular mulberry cultivars (Morus indica L. cvs.V-1, MR-2, S-36 and K-2) were assessed for drought tolerance with an integration of selective approaches. The potted plants were subjected to two watering treatments for 75 days: control pots were watered up to 100% field capacity (FC) and stressed pots were maintained at 25–30% FC. Net photosynthetic rate (P_n), stomatal conductance (g_s), transpiration rate (E) and instantaneous water use efficiency (WUE_i) were the key parameters to assess photosynthetic gas exchange performance. Drought caused marked down-regulation in leaf gas exchange in all cultivars (cvs) except V-1 which maintained better P_n, g_s, E and higher WUE_i under severe water deficit. All the four cvs also showed differential antioxidative responses under water stress. Higher concentrations of carotenoids, ascorbic acid, glutathione, α-tocopherol and proline were observed in the leaf extracts of V-1, while minimum accumulation of those metabolites was recorded with K-2 and S-36. An endogenous loss of α-tocopherol and higher lipid peroxidation were encountered in K-2, S-36 and MR-2, whereas V-1 showed minimum lipid peroxidation under water deficit regimes. Comparative morpho-anatomical analysis revealed a well-developed root system and a better anatomical architecture in V-1 which could further contribute tolerance during drought stress.