Water deficit, leaf rolling and susceptibility to photoinhibition in field grown sorghum

Chlorophyll fluorescence and gas-exchange techniques were used to investigate changes in photosynthelic performance in response to high light and mild water deficit, in two cultivars of the C., plant sorghum ( Sorghurn bicofor [L.] Moench). grown under field conditions. For all leaves fully exposed to the sun, the efficiency of phcttosystem 11 (PSII) showed a mid-day decline, hut with substantial over-night recovery: the magnitude of the mid-day decline was enhanced by water deficit. There was no corresponding decline in leaves not exposed to full sunlight, either because they were shaded by other leaves or else because of leaf-roiling. Net assimilation rates appeared more sensitive to water-deficit than was PSI1 efficiency. Shade-adapted leaves had lower rates of photosynthesis in full sun (and lower stomatal conductances) than well-exposed leaves. When these shade-adapted leaves were suddenly exposed to full sunlight, fluorescence quenching was slow. especially when plants were well-watered. For the latter, photochemical quenching (q_p)was small even after several minutes. indicating a continuing imbalance between energy funnelled to PSI1 and subsequent electron transport. Shade-adapted leaves that were water stressed were better able to withstand a sudden increase in irradiance than those that were well watered. It is suggested that the shade-adapted eaves from unirrigated plants. having a lower s'tomatal conductance than the irrigated leaves, had been acclimated by receiving energy in excess of that required to fix CO₂, thus leading to the operation of dissipative mechanisms. A shortened protocol for quenching analysis is proposed that enables non-photochemical quenching to be partitioned into rapidly and slowly relaxing components (the latter including photoinhibition) by relating results to a theoretical maximum yield of variable fluorescence. This is particularly suitable for screening field material.     

Infection time and density influence the response of sorghum to the parasitic angiosperm Striga hermonthica

Two cultivars of sorghum (CSH-1 and Ochuti) were grown in the presence and absence of the root hemiparasite Striga hermonthica in uniform conditions in the field in Kenya, Africa. S. hermonthica had a marked influence on growth and photosynthesis of 'CSH-1'; however, 'Ochuti' showed a less severe response to infection and tolerance of the parasite. The variation in genotype response might be partly explained by later attachment of the parasite and a lower level of infection. Laboratory studies were used to determine the importance of both variables in determining host response to infection. Early infection by S. hermonthica had a more negative effect on the host than late infection. The level of parasite biomass supported by the host also influenced host productivity but the relationship was nonlinear. Low degrees of parasite infection had a proportionately much greater effect on host grain weight than at greater parasite loading. Early infection of 'Ochuti' in laboratory conditions resulted in lower stem dry weight than in uninfected plants but not in smaller total plant biomass or lower rates of photosynthesis. In conclusion, the time of parasite attachment affected host performance and might explain much of the variation in host sensitivity both within and between studies. The level of parasite infection affected host performance to a lesser extent. In addition, late attachment and low levels of infection might have implications for control management strategies.     

The influence of the parasitic Striga species on the host crops sorghum and maize

The biology of the Striga species will be briefly described. Pot experiments indicate that damage to the host plant is very much greater than is explained by removal of photosynthates. A phytotoxin is almost certainly involved, which causes decreased shoot growth of the host plant but which also directly or indirectly encourages increased root growth. The significance of this change in root/shoot ratio will be discussed in relation to host resistance and various control measures.     

Development and assessment of fusarium oxysporum-based mycoherbicide for control of Striga Hermonthica in sorghum

Abstract

Experiments were carried out from 2002 to 2003 to determine the most suitable form of fungal delivery for possible use by farmers in biological control of Striga hermonthica. Six mycoherbicides were developed, based on Fusarium oxysporum isolated from wilted S. hermonthica. In mycoherbicide formulation, rock phosphate powder, sorghum bran and gum arabic powder were used as carriers. Besides its role as a carrier, gum arabic powder was used as a sticker. There were three carriers with two formulations each, making six treatments altogether. Living propagule studies were based on colony, mycelium and conidium number of F. oxysporum. In greenhouse evaluation of mycoherbicides, each kg sorghum seed was coated with 10 g mycoherbicide before sowing. Carrier rock phosphate powder with gum arabic powder as a sticking agent was the most suitable form of its delivery for use by peasant farmers.     

Inhibition of Striga seed germination associated with sorghum growth promotion by soil bacteria

Striga spp. are obligate parasitic weeds of tropical cereals and generally have the same host range as rhizospheric bacteria of the genus Azospirillum. Four strains of Azospirillum brasilense, isolated from soil where sorghum is grown, have been tested for their effect on germination of Striga hermonthica seeds and on cereal (Sorghum vulgare) growth. Two out of four strains assayed significantly inhibited germination of the parasite. Moreover, one of the two strains showed a plant growth promoting (PGPR) effect.     

The parasitic angiosperm Striga hermonthica can reduce photosynthesis of its sorghum and maize hosts in the field

Two cultivars of sorghum (CK60 and Ochuti) and one cultivarof maize (H511) were grown in field plots in western Kenya inthe presence or absence of the parasitic angiosperm Striga hermonthica,with or without a single addition of nitrogen fertilizer (150kg N ha^–1) using a factorial design. A progressive declinein rates of photosynthesis of Striga-infected plants were observedfor the sorghum cultivar CK60 from 30 d after planting (DAP)and for maize from 40 DAP, until measurements ended 63 DAP.At this time photosynthetic rates were 46% and 31% lower inthe Striga-infected sorghum and maize cultivars, respectively,compared to uninfected control plants. No decline in photosynthesiswas observed in the second sorghum cultivar studied, Ochuti,a local land race reported to show some tolerance to the parasite.The trends in photosynthesis reflected stunting of the cereals,as determined by the height of the youngest emerged ligule,however, only the grain yield of the sorghum cultivar CK60 wassignificantly reduced by the presence of the parasite. The nitrogenapplication influenced neither the growth nor the photosyntheticparameters measured, and possible explanations for the absenceof responses are discussed. It is concluded that S. hermonthicacan reduce photosynthetic rates of field-grown sorghum and maize,and suggest that an ability to maintain high rates of photosynthesiswhilst infected may be an important correlate of tolerance tothe parasite. Key words: Parasitic angiosperm, photosynthesis, nitrogen, tropical weeds, tropical agriculture     

Root nitrogen concentration of sorghum above 2% produces least Striga hermonthica seed stimulation

A series of pot and laboratory experiments was carried out to assess the effects of N status of sorghum roots and timing of N application (as NH₄NO₃) on the germination of Striga hermonthica seeds. Root N concentrations varied from 10 to 26 mg N g^?1. The cut root and the root exudates technique used in assaying S. hermonthica seed germination gave similar results. However, the cut root technique was easier to handle and more discriminating at low germination levels. S. hermonthica seed germination per unit sorghum root mass followed a broken‐stick model. It decreased with increasing root N concentrations, reaching lowest levels at a root N concentration of 19.5 mg N g^?1, after which no further reduction occurred. It was not possible to reduce S. hermonthica seed germination to a zero level. Timing of N application influenced the time a higher N concentration is reached, not the S. hermonthica seed germination. Both timing and rate of N application are important in maintaining root N concentrations above 19.5 mg N g^?1, thereby potentially reducing S. hermonthica germination in the field. Translation of results to reductions in infection levels and yield losses is hampered by density‐dependent relations after the S. hermonthica germination stage.     

Reduced sensitivity to photoinhibition following frost-hardening of winter rye is due to increased phosphate availability

The possibility of a role for phosphate metabolism in the photosynthetic regulation that occurs during frost hardening was investigated in winter rye (Secale cereale L. cv. Musketeer). Leaves of frost-hardened and non-hardened winter rye were studied during photosynthetic induction, and at steady state after being allowed to take up 20 mM orthophosphate through the transpiration stream for 3 h. At the growth irradiance (350 mol·m^-2·s^-1) frost-hardening increased the stationary rate of CO₂-dependent O₂ evolution by 57% and 25% when measured at 5 and 20° C, respectively. Frosthardening also reduced the lag phase to stationary photosynthesis by 40% at 5° C and decreased the susceptibility of leaves to oscillations during induction and after interruption of the actinic beam during steady-state photosynthesis. These responses are all indicative of increased phosphate availability in frost-hardened leaves. As reported previously by Öquist and Huner (1993, Planta 189, 150–156), frost-hardening also decreased the reduction state of Q_A, the primary, stable quinone acceptor of PSII, and decreased the sensitivity of winter rye to photoinhibition of photosynthesis. Non-hardened rye leaves fed orthophosphate also showed an increased photosynthetic capacity (25% at 20° C and light saturation), lower reduction state of Q_A, a reduced sensitivity to photoinhibition and lower susceptibility to oscillations resulting from a brief interruption of the actinic light. Thus, the data indicate that phosphate metabolism plays a key role in photosynthetic acclimation of winter rye to low temperatures.Abbreviations F_o and F_o minimal fluorescence when all PSII reaction centres are open in dark-and light-acclimated leaves, respectively - F_m and F_m maximal fluorescence when all PSII reaction centres are closed in dark-and light-acclimated leaves, respectively - F_v variable fluoresence (F_m -F_o) in dark-acclimated leaves - F_v variable fluorescence (F_m-F_o) in light-acclimated leaves - PCR photosynthetic carbon reduction - PPFD photosynthetic photon flux density - Q_A the primary, stable quinone acceptor of PSII - q_P photochemical quenching of fluorescence - q_N non-photochemical quenching of fluorescence This work was supported by the Swedish Natural Sciences Research Council. The authors are indebted to Dr. N. Huner, Department of Plant Sciences, UWO, London, Canada, for helpful discussions during the initiation of this work and for the gift of rye seeds.     

Effect of short-term heat treatment of rice seedlings on sensitivity of thylakoid membranes to photoinhibition

The after-effects of 24 h high temperature (35 or 45 °C) treatment on the photochemical activities and photooxidative lipid peroxidation, subsequent to their irradiation were studied in 7-d-old etiolated rice (Oryza sativa) seedlings. Photosystem (PS) 1 and PS 2 mediated photoreactions of thylakoids isolated from the seedlings exposed to high temperature did not differ significantly from the thylakoids isolated from control seedlings (25 °C). Hence, all kinds of tested thylakoids were equally efficient in capturing and utilizing radiant energy. The high irradiance induced loss in PS 2 activity and lipid peroxidation measured in terms of malondialdehyde production was more rapid in thylakoids isolated from stressed seedlings as compared to that of control seedlings. Thus the thylakoids isolated from the stressed seedlings were more prone to photodamage than those from the control seedlings.     

A time-course study of early establishment stages of parasitic angiosperm Striga asiatica on susceptible sorghum roots

A time-course study of the early establishment stages of Striga asiatica was carried out on a susceptible sorghum hybrid, CSH 1, using polyethylene bags and whole-root clearing and staining techniques. Preconditioned Striga seeds were applied to different aged segments of primary root but the results did not differ for these different aged segments. Most of the Striga seeds (63%) germinated within 24 h of inoculation on the host roots. The attachment of Striga radicles to host root was rapid and it occurred between 36 and 48 h after inoculation. Only 9% of the germinated Striga seeds attached to the host root but 65% of these attachments successfully penetrated through the epidermis and entered the host cortex within 72 h. Penetration through the cortical cells was difficult; only 17% of attachments were able to reach the endodermis. Penetration took from 12 to 43 h after the first appearance of haustorial cells in the cortex; a total of 84 to 120 h after inoculation on the host root. Penetration through the endodermis and establishment on the host stele was relatively easier, as most of the haustoria reaching the endodermis were able to establish on the host stele. But this is a slow process taking a minimum of 24 h, and a maximum of 60 h after first contact of haustorial cells with the endodermis. The minimum time taken from inoculation of ungerminated Striga seed on the host root to establishment is about 108 h. The results are discussed in relation to published reports on other parasitic species such as Agalinis purpurea.     

Estimation of the effect of photoinhibition on the carbon gain in leaves of a willow canopy

The occurrence of photoinhibition of photosynthesis in leaves of a willow canopy was examined by measuring the chlorophyll-a fluorescence ratio of F _V/F _M (F_M is the maximum fluorescence level of the induction curve, and F_V is the variable fluorescence, F _V=F _M–F ₀, where F₀ is the minimal fluorescence). The majority of the leaves situated on the upper parts of peripheral shoots showed an afternoon inhibition of this ratio on clear days. This was the consequence of both a decrease in F _M and a rise in F _O. In the same leaves the diurnal variation in intercepted photosynthetic photon flux density (PPFD) was monitored using leaf-mounted sensors. Using the multivariate method, partial least squares in latent variables, it is shown that the dose of PPFD, integrated and linearly weighted over the last 6-h period, best predicts photoinhibition. Photoinhibition occurred even among leaves that did not intercept PPFDs above 1000 mol·m^–2·s^–1. Exposure of leaves to a standard photoinhibitory treatment demonstrated that the depression in the F _V/F _M ratio was paralleled by an equal depression in the maximal quantum yield of CO₂ uptake and a nearly equal depression in the rate of bending (convexity) of the light-response curve of CO₂ uptake. As a result, the rate of net photosynthesis is depressed over the whole natural range of PPFD. By simulating the daily course in the rate of net photosynthesis, it is estimated that in the order of one-tenth of the potential carbon gain of peripheral willow shoots is lost on clear days as a result of photoinhibition. This applies to conditions of optimal temperatures. Photoinhibition is even more pronounced at air temperatures below 23° C, as judged from measurements of the F_V/F_M ratio on clear days: the afternoon inhibition of this ratio increased in a curvilinear manner from 15% to 25% with a temperature decrease from 23° to 14° C.Abbreviations and Symbols F_O minimum fluorescence - F_V variable fluorescence - F_M maximum fluorescence - PLS partial least squares in latent variables - PPFD photosynthetic photon flux density - VPD water vapour-pressure deficit This study was supported by the Swedish Natural Science Research Council. We are indebted to Dr. Jerry Leverenz (Department of Plant Physiology, University of Umeå, Sweden) for guidance with the modelling of the photosynthesis data.     

Overexpression of zeaxanthin epoxidase gene enhances the sensitivity of tomato PSII photoinhibition to high light and chilling stress

A tomato ( Lycopersicon esculentum Mill.) zeaxanthin epoxidase gene ( LeZE ) was isolated. The deduced amino acid sequence of LeZE showed high identities with zeaxanthin epoxidase in other plant species. Northern blot analysis showed that the mRNA accumulation of LeZE in the wild-type (WT) was not induced by light and temperature but regulated by the diurnal rhythm. The sense transgenic plants were obtained under the control of the cauliflower mosaic virus 35S promoter (35S-CaMV). Northern and western blot analysis confirmed that sense LeZE was transferred into the tomato genome and overexpressed. The ratio of (A + Z)/(V + A + Z) and the values of non-photochemical quenching were lower in transgenic plants than in WT plants under high light and chilling stress with low irradiance. The O₂ evolution rate and the maximal photochemical efficiency of PSII (Fv/Fm) in transgenic plants decreased more quickly during both stresses and recovered slower than that in WT under optimal conditions. These results suggested that overexpression of LeZE impaired the function of the xanthophyll cycle and aggravated PSII photoinhibition in tomato under high light and chilling stress.     

A change in the sensitivity of elongation factor G to oxidation protects photosystem II from photoinhibition in Synechocystis sp. PCC 6803

The repair of photosystem II (PSII) after photodamage is particularly sensitive to oxidative stress and inhibition of such repair is associated with the oxidation of specific cysteine residues in elongation factor G (EF-G), a key translation factor, in the cyanobacterium Synechocystis sp. PCC 6803. Expression of mutated EF-G with a target cysteine residue replaced by serine in Synechocystis resulted in the protection of PSII from photoinhibition. This protection was attributable to the enhanced repair of PSII via acceleration of the synthesis of the D1 protein, which might have been due to reduced sensitivity of protein synthesis to oxidative stress.     

Towards effective resistance to Striga in African maize

The fascinating biology of Striga parasitism is manifest through a series of signal exchanges between the parasite and its host. As an obligate root hemi-parasite, Striga development is cued to exudates and solutes of host roots but with negative ramifications on host plant health. Striga control in crops, via a variety of biotechnological approaches, needs to be based on increased understanding of this intricate biology. Maize has become the major cereal crop of Africa. However, this New World transplant has shown a paucity of Striga resistance characters relative to native sorghum. In this paper, we review growing evidence for maize genetic defenses against early pre-emergent phases of the Striga life cycle, when the tolls of parasitism are first manifest. Resistance characters first described in maize wild relatives have now been captured in Zea mays. The possible stacking of new and complementary sources of resistance in improved maize varieties targeted for Striga prone areas is discussed. An integrated approach combining genetic with other control measures is advocated with a more realistic view of the resource challenges prevalent in African agriculture.Key words: Striga, parasitic weed, maize, sorghum, resistance, integrated control     

The inhibited xanthophyll cycle is responsible for the increase in sensitivity to low temperature photoinhibition in rice leaves fed with glutathione

Exposure of intact rice leaves to an irradiance of 1000 μmol m⁻² s⁻¹ at 6 °C for 2 h caused severe photoinhibition of Photosystem II. The rate and extent of photoinhbition were greatly exacerbated in leaves fed with 10 mM reduced glutathione (GSH) or 10 mM cysteine. Analyses of antioxidant enzyme activities as well as the application of protein synthesis inhibitors revealed that the increased sensitivity to photoinhibition following GSH feeding was not related to its effect on cellular antioxidant systems. On the other hand, feeding with GSH markedly suppressed the formation of zeaxanthin and antheraxanthin via the xanthophyll cycle and its associated nonradiative energy dissipation in leaves chilled in high light, suggesting that the stimulating effect of exogenous GSH on photoinhibition may be attributable to its action on the xanthophyll cycle. In vitro experiments using isolated thylakoids indicated that GSH is a weak inhibitor of violaxanthin deepoxidation. The possible implications of these results are discussed. This revised version was published online in June 2006 with corrections to the Cover Date.     

A role of the C-terminal extension of the photosystem II D1 protein in sensitivity of the cyanobacterium Synechocystis PCC 6803 to photoinhibition.

The D1 protein, a key protein subunit of Photosystem II complex (PSII), is synthesised as a precursor (pD1) with a carboxyl-terminal extension. In the cyanobacterium Synechocystis sp. PCC 6803, this extension consists of 16 amino acid residues and it is cleaved by a specific protease in two putative steps with the final cleavage after the residue Ala344. In order to define the importance of the extension for the functioning of PSII, we constructed and characterized several site-directed mutants of Synechocystis that differ in the length and amino acid sequence of this extension. The mutant lacking the entire C-terminal extension exhibited slightly increased sensitivity to photoinhibition. Analysis of the PSII assembly in the mutant by the blue-native electrophoresis in combination with radioactive labelling revealed an increased level of the unassembled D1 protein in this strain. Replacement of the amino acid residue Asn359 by His or Asp also led to the higher vulnerability to photoinhibition of both mutants. In the Asn359His mutant, this vulnerability was accompanied by an increased level of the PSII core lacking CP43 indicating limitation of the repair cycle in the CP43 reassembly step.     

Photosystem II in a mutant of Chlamydomonas reinhardtii lacking the 23 kDa psbP protein shows increased sensitivity to photoinhibition in the absence of chloride

The psbP gene product, the so called 23 kDa extrinsic protein, is involved in water oxidation carried out by Photosystem II. However, the protein is not absolutely required for water oxidation. Here we have studied Photosystem II mediated electron transfer in a mutant of Chlamydomonas reinhardtii, the FUD 39 mutant, that lacks the psbP protein. When grown in dim light the Photosystem II content in thylakoid membranes of FUD 39 is approximately similar to that in the wild-type. The oxygen evolution is dependent on the presence of chloride as a cofactor, which activates the water oxidation with a dissociation constant of about 4 mM. In the mutant, the oxygen evolution is very sensitive to photoinhibition when assayed at low chloride concentrations while chloride protects against photoinhibition with a dissociation constant of about 5 mM. The photoinhibition is irreversible as oxygen evolution cannot be restored by the addition of chloride to inhibited samples. In addition the inhibition seems to be targeted primarily to the Mn-cluster in Photosystem II as the electron transfer through the remaining part of Photosystem II is photoinhibited with slower kinetics. Thus, this mutant provides an experimental system in which effects of photoinhibition induced by lesions at the donor side of Photosystem II can be studied in vivo.Abbreviations Chl chlorophyll - DCIP 2,6-dichlorophenolindophenol - DPC 2,2-diphenylcarbonic dihydrazide - HEPES 4-(2-hydroxyethyl)-1-piperazinethanesulfonic acid - P₆₈₀ the primary electron donor to PS II - PpBQ phenyl-p-benzoquinone - PS II Photosystem II - Q_A the first quinone acceptor of PS II - Q_B the second quinone acceptor of PS II - SDS sodium dodecyl sulfate - Tris tris(hydroxymethyl)aminomethane - Tyr_D accessory electron donor on the D₂-protein - Tyr_Z tyrosine residue, acting as electron carrier between P₆₈₀ and the water oxidizing system