The beneficial effect of reduced elongation growth on submergence tolerance of rice

Adverse effects of elongation growth on tolerance to completesubmergence for up to 14 d were evaluated in rice seedlingsof cultivars which differed in submergence tolerance. Thereis a good negative correlation between per cent survival andelongation growth of genotypes during complete submergence (r= – 0.81). When elongation growth underwater is minimizedby application of a gibberellin biosynthesis inhibitor, percent survival increases by as much as 50 times for one cultivar.These effects are likely related to elongation growth since(i) addition of gibberellin had the opposite effect by reducingsurvival, and (ii) when the elongation inhibitor and gibberellinwere added together, there was no effect on elongation growthand the per cent survival did not change. A GA-deficient mutantof rice which had little elongation ability during submergenceshowed a high level of submergence tolerance when plants weresubmerged at equal initial dry weights and carbohydrate levelsrelative to a submergence-tolerant cultivar. These results areconsistent with the hypothesis that elongation growth competeswith maintenance processes for energy and hence reduces survivalduring submergence. The impact of these findings is that inenvironments where elongation ability is not required, thereis a potential to increase submergence tolerance of agriculturallyimportant cultivars by selecting for least elongation, at leastduring periods of complete submergence. Furthermore, this trade-offbetween stimulated elongation growth and submergence tolerancewill have important ecological consequences for the distributionof plant species in different flood-prone environments. Key words: Gibberellin, growth, Oryza sativa, rice, submergence     

Molecular and Physiological Mechanisms of Flooding Avoidance and Tolerance in Rice

Submergence is one of the major constraints in rice production. The main factor limiting rice survival during submergence is oxygen deprivation. To cope with flooding conditions, rice has developed two survival strategies: either rapid elongation of the submerged tissues to keep up with the rising water level or no elongation to save carbohydrate resources for maintenance of energy production under submerged and concomitant hypoxic conditions. The survival strategies used by rice have been studied quite extensively and the role of several phytohormones in the elongation response has been established. The mechanisms of submergence tolerance include metabolic changes, for instance, the shift to an ethanolic fermentation pathway, reduced elongation growth to save carbohydrates and energy for maintenance processes, and protective antioxidant systems. Current molecular technology can provide tools for the understanding of mechanisms developed by rice to survive submergence. In addition, cloning of genes related to submergence tolerance might open new ways to genetic improvement of this crop.     

水稻的耐淹性状及其Sub1基因

近4年来, 水稻(Oryza sativa L.)耐淹性状及其分子机理的研究取得了长足的进展。水稻植株的耐淹性状主要由Submergence-1 (Sub1) 基因控制。Sub1通过调节乙烯和赤霉素介导的反应, 抑制淹水期间的伸长生长和减缓碳水化合物消耗来控制耐淹性状。文章介绍了水稻应对淹涝胁迫的两种策略, 影响耐淹性状的主要生理因素, Sub1基因定位以及它控制耐淹性状的机理; 阐述了Sub1基因在我国杂交水稻生产方面的应用前景。     

Oxygen deprivation stress in a changing environment

Past research into flooding tolerance and oxygen shortages inplants has been motivated largely by cultivation problems ofarable crops. Unfortunately, such species are unsuitable forinvestigating the physiological and biochemical basis of anoxia-toleranceas selection has reduced any tolerance of anaerobiosis and anaerobicsoil conditions that their wild ancestors might have possessed.Restoration of anoxia-tolerance to species that have lost thisproperty is served better by physiological and molecular studiesof the mechanisms that are employed in wild species that stillpossess long-term anoxia-tolerance. Case studies developingthese arguments are presented in relation to a selection ofcrop and wild species. The flooding sensitivity and metabolismof maize is compared in relation to rice in its capacity foranaerobic germination. The sensitivity of potato to floodingis related to its disturbed energy metabolism and inabilityto maintain functioning membranes under anoxia and postinoxia.By contrast, long-term anoxia-tolerance in the American cranberry(Vaccinium macrocarpon) and the arctic grass species Deschampsiaberingensis can be related to the provision and utilizationof carbohydrate reserves. Among temperate species, the sweetflag (Acorus calamus) shows a remarkable tolerance of anoxiain both shoots and roots and is also able to mobilize carbohydrateand maintain ATP levels during anoxia as well as preservingmembrane lipids against anoxic and post-anoxic injury. Phragmitesaustralis and Spartina alterniflora, although anoxia-tolerant,are both sulphide-sensitive species which can pre-dispose themto the phenomenon of die-back in stagnant, nutrient-rich water.Glyceria maxima adapts to flooding through phenological adaptationswith a seasonal metabolic tolerance of anoxia confined to winterand spring which, combined with a facility for root aerationand early spring growth, allows rapid colonization of siteswith only shallow flooding. The diversity of responses to floodingin wild plants suggests that, depending on the life strategyand habitat of the species, many different mechanisms may beinvolved in adapting plants to survive periods of inundationand no one mechanism on its own is adequate for ensuring survival. Key words: Anoxia, hypoxia, flooding, Zea mays, Solanum tuberosum, Oryza sativa, Acorus calamus, Phragmites australis, Glyceria maxima, cranberry     

Genetic engineering of rice capable of synthesizing fructans and enhancing chilling tolerance

Fructans are water-soluble fructose oligomers and polymers thatare based on sucrose, and have been implicated in protectingplants against water stress. Rice (Oryza sativa L.) is highlysensitive to chilling temperatures, and is not able to synthesizefructans. Two wheat fructan-synthesizing enzymes, sucrose:sucrose1-fructosyltransferase, encoded by wft2, or sucrose:fructan6-fructosyltransferase, encoded by wft1, were introduced intorice plants, and rice transformants that accumulate fructanswere successfully obtained. The mature leaf blades of transgenicrice lines with wft2 or wft1 accumulated 16.2 mg g^–1 FWof oligo- and polysaccharides mainly composed of inulin oligomersof more than DP7, and 3.7 mg g^–1 FW of oligo- and polysaccharides,mainly composed of phlein oligomers of more than DP15, respectively.The transgenic rice seedlings with wft2 accumulated significantlyhigher concentrations of oligo- and polysaccharides than non-transgenicrice seedlings, and exhibited enhanced chilling tolerance. Theoligo- and polysaccharide concentrations of seedlings expressingwft1 were obviously lower than those of lines expressing wft2,and no correlation between oligo- and polysaccharide concentrationsand chilling tolerance was detected in wft1-expressing ricelines. The results suggest that transgenic rice lines expressingwheat-derived fructosyltransferase genes accumulated large amountsof fructans in mature leaf blades and exhibited enhanced chillingtolerance at the seedling stage. This is the first report owingthat fructan accumulation enhanced tolerance to non-freezinglow temperatures. Key words: Chilling tolerance, fructan, fructosyltransferase, Oryza sativa, transgenic plant     

Plasma Membrane Permeability of Root-Tip Cells Following Temporary Exposure to Al Ions Is a Rapid Measure of Al Tolerance among Plant Species

No correlations were recognized between Al tolerance among fourplant species, rice (Oryza sativa L.), maize (Zea mays L.),pea (Pisum sativum L.), and barley (Hordeum vulgare L.), inrank order of Al tolerance, and cation exchange capacities ofroot-tip (0-1 cm) cells or of their cell walls. The plasma membraneof root-tip of Al sensitive plant species (pea and barley) wasconsiderably permeabilized with elongation of root in Al-freesolution following 0.5 h pretreatment with Al. K⁺ release fromand Al permeation into the protoplasts isolated from the root-tipof Al-sensitive plant species were more significant than thosefor Al-tolerant plant species (rice and maize) on 10 or 30 mintreatment with Al. The permeability of the plasma membrane forprotoplasts isolated from Al sensitive plant species was considerablyincreased by treatment with hy-potonic Al-free control solutionfollowing 10 min pretreatment with Al. To our knowlege, theseare the most rapid responses to Al ions reported to date, i.e.,within 0.5 h in whole plant and within 10 min in protoplast.These results suggest that a temporary contact with Al ionsirreversibly alters the plasma membrane of root-tip cells ofAl-sensitive plant species: the cells become more leaky andrigid due to binding of Al ions to the plasma membrane. (Received January 5, 1998; Accepted February 26, 1998)     

Relationship between Coleoptile Elongation and Alcoholic Fermentation in Rice Exposed to Anoxia. II. Cultivar Differences

The relationship between coleoptile elongation and survivalvs. alcoholic fermentation of rice under anoxia is examinedusing eight cultivars differing in submergence tolerance. Anoxiawas imposed on either 1 or 4 d aerated seeds either by N₂ flushingsubmerged tissues or by incubating tissues in stagnant deoxygenatedagar at 0·1% w/v; the latter simulated the stagnant conditionsof waterlogged soil. Two cultivars that were most tolerant tosubmergence also had the greatest tolerance to anoxia, whilea submergence intolerant cultivar was also intolerant to anoxia. Coleoptile growth under anoxia was related to rates of ethanolsynthesis (R_E), however differences between growth during anoxiaand survival after anoxia indicated that post-anoxic injurymay also be important in rice seeds exposed to anoxia. The correlationbetween coleoptile growth and R_E measured on a tissue basisusing intact seeds was r² = 0·67 among six varietiesover 0-3 d anoxia. This correlation improved to about r² = 0·85using R_E of (embryos plus coleoptiles) over 0-3 d, or coleoptilesat 3 d after anoxia. Coleoptile growth of individual seeds wasusually poorly correlated to R_E in these cultivars at 2-3 dafter anoxia. When coleoptiles of similar lengths were obtainedfrom different cultivars using 4 d aerated seeds, there weredifferences in R_E and coleoptile growth which were related tocoleoptile growth during 3 or 5 d anoxia, either on a tissue(r² = 0·85) or a fresh weight basis (r² = 0·70-0·97respectively). Results are discussed in relation to factorswhich may limit ethanol synthesis in rice exposed to anoxiaand the importance of growth to the survival of seeds and matureplants during submergence in the natural environment.Copyright1994, 1999 Academic Press Anoxia, ethanol, alcoholic fermentation, Oryza sativa L., rice, submergence     

Desiccation Tolerance and Potential Longevity of Developing Seeds of Rice (Oryza sativa L.)

The longevity and desiccation tolerance of samples of seedsof a japonica rice (Oryza sativa L.) harvested serially duringdevelopment from plants grown in two temperature regimes, viz28/20 °C and 32/24 °C (12/12 h) were determined. Massmaturity (defined as the end of the seed-filling phase) occurred19·7 and 18·3 d after 50% anthesis, respectively.Longevity (determined at 40 °C with 15% moisture contentand quantified by the value of the constant K_i of the seed viabilityequation) improved during seed development and maturation until17 and 14 d after mass maturity in the cooler and warmer regimes,respectively, but declined thereafter. Changes in K_i with timewere similar in the two environments until mass maturity, butthe increase in K_i values after mass maturity was much greaterin the cooler regime. Tolerance of desiccation to low (4%) moisturecontents improved until 22 and 14 d after mass maturity in thecooler and warmer regimes, respectively, when maturation dryinghad reduced seed moisture contents naturally to 24 and 32% moisturecontent, respectively. Further delays to seed harvest reduceddesiccation tolerance, particularly in the warmer environment.Comparison among 15 samples of seeds harvested at differenttimes in the two environments showed a strong correlation (r= 0·947, P < 0·01) between longevity (K_i) anddesiccation tolerance (to 4% moisture content). Hence, it issuggested that the regulation of desiccation tolerance to lowmoisture contents and potential air-dry longevity during seeddevelopment and maturation determined here may have a commoncause.Copyright 1994, 1999 Academic Press Oryza sativa L., rice, desiccation tolerance, genebanks, seed development, seed longevity, temperature     

Genes/QTLs affecting flood tolerance in rice

The adaptation of deepwater rice to flooding is attributed to two mechanisms, submergence tolerance and plant elongation. Using a QTL mapping study with replicated phenotyping under two contrasting (water qualities) submergence treatments and AFLP markers, we were able to identify several genes/QTLs that control plant elongation and submergence tolerance in a recombinant inbred rice population. Our results indicate that segregation of rice plants in their responses to different flooding stress conditions is largely due to the differential expression of a few key elongation and submergence tolerance genes. The most important gene was QIne1 mapped near sd-1 on chromosome 1. The Jalmagna (the deepwater parent) allele at this locus had a very large effect on internal elongation and contributed significantly to submergence tolerance under flooding. The second locus was a major gene, sub1(t), mapped to chromosome 9, which contributed to submergence tolerance only. The third one was a QTL, QIne4, mapped to chromosome 4. The IR74 (non-elongating parent) allele at this locus had a large effect for internal elongation. An additional locus that interacted strongly with both QIne1 and QIne4 appeared near RG403 on chromosome 5, suggesting a complex epistatic relationship among the three loci. Several QTLs with relatively small effects on plant elongation and submergence tolerance were also identified. The genetic aspects of these flooding tolerance QTLs with respect to patterns of differential expression of elongation and submergence tolerance genes under flooding are discussed. Received: 13 December 1999 / Accepted: 14 March 2000<@head-com-p1a.lf>Communicated by G. Wenzel     

Does ethylene induce elongation of the rice coleoptile through auxin?

Ethylene stimulated the elongation of intact rice (Oryza sativaL.) coleoptiles in which endogenous growth had been stoppedcompletely by decapitation and red light. p-Chlorophenoxyisobutyricacid slightly inhibited endogenous growth, but not the ethyleneinduced growth. Thus, ethylene could stimulate the elongationof coleoptiles in which the auxin level was considered to bevery low. ¹ Present address: Institute for Agricultural Research, TohokuUniversity, Katahira, Sendai 980, Japan. (Received February 16, 1979; )     

褐飞虱翊型分化遗传规律的研究

以褐飞虱Nilaparvata lugens (Stal)长翅型(macroptery,简称M)、短翅型(brachyptery,简称B)的遗传纯系为实验材料,进行亲本、稻株生育期、虫口密度三因子交互实验。结果表明：(1)在环境条件(指稻株生育期、虫口密度等)一致时,B♀×B♂、B♀×M♂、M♀×B♂、M♀×M♂四种亲本组合的F1代短翅型成虫百分率分别为：98%、92%、64%、29%,各亲本组合间差异极显著;(2)亲本相同时,将F1代褐飞虱初孵若虫多头词养(多于l0头/株)在黄熟期稻株上,其长翅型雌、雄成虫百分数均高于灌浆期稻株上;(3)单头饲养实验中发现,不论亲本组合、稻株生育期如何,雌虫绝大多数分化为短翅型,而雄虫则几乎全为长翅型。这表明褐飞虱的翅型分化遗传由一个受多种因子影响的调控体系决定,且调控作用与性别有关。     

OsRAN2, essential for mitosis,enhances cold tolerance in rice by promoting export of intranuclear tubulin and maintaining cell division under cold stress

With global climate change, abnormally low temperatures have affected the world's rice production. Many genes have been shown to be essential for molecular improvement of rice cold‐tolerance traits. However, less is known about the molecular cellular mechanism of their response to cold stress. Here, we investigated OsRAN2 involved in regulation of cell division during cold stress in rice. Expression of OsRAN2 was increased under cold treatment, but not during salt and drought stress. The mean root mitotic index was closely related to the expression level of OsRAN2. Knockdown transgenic rice lines showed an aberrant organization of spindles during mitosis and stunted growth during development. Overexpression of OsRAN2 enhanced cold tolerance in rice. The transgenic rice overexpressing OsRAN2 showed maintained cell division, decreased proportion of cells with intranuclear tubulin and formation of a normal nuclear envelope under the cold condition. Our study suggests a mechanism for OsRAN2 in regulating cold resistance in rice by maintaining cell division through promoting the normal export of intranuclear tubulin at the end of mitosis. This insight could help improve the cold‐tolerance trait in rice.     

Role of the Aquaporin PIP1 Subfamily in the Chilling Tolerance of Rice

Although an association between chilling tolerance and aquaporinshas been reported, the exact mechanisms involved in this relationshipremain unclear. We compared the expression profiles of aquaporingenes between a chilling-tolerant and a low temperature-sensitiverice variety using real-time PCR and identified seven genesthat closely correlated with chilling tolerance. Chemical treatmentexperiments, by which rice plants were induced to lose theirchilling tolerance, implicated the PIP1 (plasma membrane intrinsicprotein 1) subfamily member genes in chilling tolerance. Ofthese members, changes in expression of the OsPIP1;3 gene suggestedthis to be the most closely related to chilling tolerance. AlthoughOsPIP1;3 showed a much lower water permeability than membersof the OsPIP2 family, OsPIP1;3 enhanced the water permeabilityof OsPIP2;2 and OsPIP2;4 when co-expressed with either of theseproteins in oocytes. Transgenic rice plants (OE1) overexpressingOsPIP1;3 showed an enhanced level of chilling tolerance andthe ability to maintain high OsPIP1;3 expression levels underlow temperature treatment, similar to that of chilling-tolerantrice plants. We assume that OsPIP1;3, constitutively overexpressedin the leaf and root of transgenic OE1 plants, interacts withmembers of the OsPIP2 subfamily, thereby improving the plants’water balance under low temperatures and resulting in the observedchilling tolerance of the plants.     

Promotion of the Growth of Self-Incompatible Pollen Tubes in Lily by cAMP

Cyclic AMP, forskolin and 3-isobutyl-1-methylxanthine promotedthe elongation of self-incompatible pollen tubes in Lilium longiflorum.It appears that a functional cAMP-regulated system that involvesadenylate cyclase and phosphodiesterase is present in the lilystyle and is involved in the regulation of elongation of self-incompatiblepollen tubes. (Received January 21, 1993; Accepted June 19, 1993)     

Patterns of Root Respiration Associated with the Induction of Aluminium Tolerance in Phaseolus vulgaris L.

Aluminium (Al) tolerance in an Al-tolerant cultivar of Phaseolusvulgaris L. (‘Dade’) was found to be an inducibletrait. Upon exposure to 10 µM Al, the rate of root elongationwas inhibited in comparison to controls. During the following72 h, the rate of elongation returned to levels comparable tocontrols. In contrast, root elongation of an Al-sensitive cultivar(‘Romano’) did not recover after exposure to Al.In Dade, the resumption of root elongation following exposureto Al was accompanied by increased rates of root respiration,whereas respiration rates slowly declined over the 72 h treatmentperiod in Romano. When partitioned into growth and maintenanceexpenditures, a larger proportion of root respiration of Dadeexposed to Al was allocated to maintenance processes, potentiallyreflecting diversion of energy to metabolic pathways that offsetthe adverse effects of Al toxicity. Romano did not show sucha pattern and respiration associated with both growth and maintenancewas reduced after exposure to Al. Root and shoot growth of bothcultivars were also measured to determine the effects of long-term(21 d) exposure to 10 µM Al. Dade plants exposed to Alexhibited enhanced growth in comparison to controls, whereasRomano plants were characterized by reduced shoot and root growth.Modelling the time-course of root respiration and measuringthe long-term growth responses to Al is a valuable method ofelucidating respiratory costs of stress tolerance. Key words: Aluminium, differential tolerance, maintenance respiration, Phaseolus vulgaris, root respiration     

Levels of {beta}-Glucan and Lignin in Elongating Internodes of Deepwater Rice

Partial submergence or treatment with either ethylene or gibberellicacid (GA₃ induces rapid growth in deepwater rice (Oryza sativaL.). We correlated the synthesis of two cell wall componentswith two phases of internodal elongation, namely (13,14)-ß-glucanformation with cell elongation and lignification with differentiationof the secondary cell wall and cessation of growth. The contentof ß-glucan was highest in the zone of cell elongationin internodes of air-grown plants and plants that were inducedto grow rapidly by submergence. In the intercalary meristemand in the differentiation zone of the internode, ß-glucanlevels were ca. 70% lower than in the zone of cell elongation.The outer cell layers, enriched in epidermis, contained moreß-glucan in submerged, rapidly growing internodesthan in air-grown, control internodes. The ß-glucancontent of the inner, parenchymal tissue was unaffected or slightlylowered by submergence. The epidermis appears to be the growth-limitingstructure of rapidly growing rice internodes. We hypothesizethat elevated levels of ß-glucan contribute to elongationgrowth by increasing the extensibility of the cell wall. Lignificationwas monitored by measuring the content of lignin and the activitiesof two enzymes of the lignin biosynthetic pathway, coniferylalcohol dehydrogenase (CAD) and phenylalanine ammonia-lyase(PAL), in growing and non-growing regions of the internode.Using submerged whole plants and GA₃-treated excised stem segments,we showed that lignin content and CAD activity were up to sixfoldlower in newly formed internodal tissue of rapidly growing ricethan in slowly growing tissue. No differences were observedin parts of the internode that had been formed prior to inductionof growth. PAL activity was reduced throughout the internodeof submerged plants. We conclude that lignification is one ofthe processes that is suppressed to permit rapid growth. ¹ This work was supported by the National Science Foundationthrough grants No. DCB-8718873 and DCB-9103747 and by the Departmentof Energy through grant No. DE-FGO2-90ER20021. M.S. was therecipient of a fellowship from the Max Kade Foundation.     

Effect of high temperature pre-treatment on elongation of the mesocotyl of rice seedlings I. Cell multiplication in the mesocotyl

Mesocotyl elongation in paddy rice seedlings of the japonicatype was markedly stimulated by high temperature pre-treatmentof seeds. Examination of the number of parenchyma cells perone cell layer in longitudinal sections of mesocotyl revealedthat stimulation may be mainly caused by cell multiplication. (Received February 24, 1970; )     

PHYSIOLOGICAL ACTIVITIES OF HELMINTHOSPOROL AND HELMINTHOSPORIC ACID: I. EFFECTS ON GROWTH OF INTACT PLANTS

Helminthosporol (H-ol) and helminthosporic acid (H-acid) wereeffective in promoting elongation of leaf sheaths of rice, Japanesebarnyard grass and dwarf maize (d-2 and d-5) and of hypocotylsof taisai (Brassica chinensis), but inactive in leaf sheathsof oat and wheat, hypocotyls of sesame and morning glory (Pharbitisnil) and epicotyls of Pharbitis and dwarf and tall peas. Onthe elongation of the leaf sheath of maize d-1, H-ol was promotivebut the activity of H-acid was doubtful. On hypocotyls of lettuceand daikon (Raphanus sativus), only H-acid was active. Multiplicationrate and size of fronds of Lemna perpusila were not affectedby either of the substances. Compared with gibberellic acid for the effect on the shoot growth,H-ol and H-acid were weak in activity and narrower in the scopeof plants that responded. H-ol and H-acid characteristicallypromoted the elongation of the primary root. Comparative effectivenessof H-ol and H-acid varied with plant species or parts examined. ¹ This study was supported in part by grant-in aid of the Ministryof Education (No. 0417). The results reported here were presentedat the Annual Meeting of the Botanical Society of Japan at Kanazawain 1964 (S).