A comparison of the submergence response of deepwater and non-deepwater rice

Twelve cultivars of rice (Oryza sativa L.), representing deepwater, short-statured, and semidwarf types, were tested for their response to submergence. The magnitude of the response varied between cultivars; however, all cultivars responded to submergence by rapid growth once internodal elongation had started. Three of these cultivars were tested for elongation capacity at four ages. The deepwater rice was capable of rapid internodal elongation in response to submergence at 4 weeks of age. Growth of the short-statured and semidwarf cultivars was not stimulated by submergence until about 10 weeks of age. In air, the internodes of deepwater rice grew slower than did those of the short-statured and semidwarf cultivars. We also investigated the elongation response of stem sections of all 12 cultivars to an atmosphere containing 3% O₂, 6% CO₂, 91% N₂ (all by volume), and 1 microliter per liter ethylene. We found that the response of each of the non-deepwater cultivars was qualitatively and quantitatively similar to that of the deepwater rice.     

Ethylene-induced stem growth of deepwater rice is correlated with expression of gibberellin- and abscisic acid-biosynthetic genes

Ethylene decreases the content of endogenous abscisic acid (ABA) and increases the level of bioactive gibberellin A₁ (GA₁) in the submerged internodes of deepwater rice. During partial submergence, internodes of deepwater rice undergo rapid elongation as a result of ethylene accumulation in the internodal lacunae. In anin vitro experiment using stem sections from deepwater rice, treatment with 5 μL L^-1 ethylene promoted stem growth by up to 3.2-foId times over air treatment. Expression patterns were analyzed for genes that encode GA- and ABA-biosynthesis enzymes to determine any possible molecular basis for the changes observed in GA₁ and ABA contents as a result of ethylene action. Expression of theOsGA20ox2 andOsGA20ox4 genes, which encode GA 20-oxidase, and of theOsGA3ox2 gene, which encodes the enzyme that converts GA₂₀ to CA₁, was up-regulated, whereas that of three ABA-biosynthetic genes —OsNCED1, OsNCED2, andOsNCEDS-was down-regulated in the presence of ethylene. These results indicate that GA and ABA contribute equally to the submergence-or ethylene-induced stem elongation of deepwater rice via the coordinated and opposite regulation of biosynthesis.     

Effect of submergence on translocation,starch content and amylolytic activity in deep-water rice

Submergence induces rapid internodal elongation in deep-water rice (Oryza sativa L. cv. Habiganj Aman II). We investigated the metabolic activities which help to support such fast growth. Three days of submergence in water under continuous light led to the mobilization of 65% of the starch from those regions of rice internodes which had been formed prior to submergence. Disappearance of starch was accompanied by a 70-fold enhancement of amylolytic activity. Similar increases in amylolytic activity were detected in response to ethylene and gibberellic acid. Submergence also caused a 26-fold increase in the translocation of newly synthesized photosynthetic assimilates from the leaves to the internodes and younger regions of the culms. These physiological processes are likely to provide the metabolic energy required for internodal elongation in response to submergence.Abbreviation GA₃ gibberellic acid     

Interactions between abscisic acid, ethylene and gibberellin in internodal elongation in floating rice: the promotive effect of abscisic acid at low humidity

The enhancement of internodal elongation in floating or deepwater rice (Oryza sativa L. cv. Habiganj Aman II) by treatment with ethylene or gibberellic acid (GA₃) at high relative humidity (RH) is inhibited by abscisic acid (ABA). Here, we examined the interactive effects of ethylene, gibberellin (GA) and ABA at low RH on internodal elongation of deepwater rice stem segments. Although ethylene alone hardly promoted internodal elongation of stem sections at 30% RH, it enhanced the internodal elongation induced by GA₃. Application of ABA alone to stem segments had no effect on internodal elongation. However, in the presence of ethylene and GA₃ at 30% RH, ABA further promoted internodal elongation. This promotive effect of ABA was not found in the internodes of stem segments treated either with ethylene or with GA₃ at 30% RH or in the internodes of stem segments treated with ethylene and/or GA₃ at 100% RH.     

The role of ethylene in the growth response of submerged deep water rice

We investigated the effect of partial submergence on internode elongation in a Bangladesh variety of floating or deep water rice (Oryza sativa L., cv. Habiganj Aman II). In plants which were at least 21 days old, 7 days of submergence led to a 3- to 5-fold increase in internodal length. During submergence, the ethylene concentration in the internodes increased from about 0.02 to 1 microliters per liter. Treatment of nonsubmerged plants with ethylene also stimulated internode elongation. When ethylene synthesis in partially submerged plants was blocked with aminooxyacetic acid and aminoethoxyvinylglycine, internode elongation was inhibited. This growth inhibition was reversed when ethylene biosynthesis was restored with 1-aminocyclopropane-1-carboxylic acid (ACC). Radio-labeling studies showed that ethylene in floating rice was synthesized from methionine via ACC. Internodal tissue from submerged plants had a much higher capacity to form ethylene than did internodal tissue from nonsubmerged plants. This increase in ethylene synthesis appeared to be due to enhanced ACC formation rather than to increased conversion of ACC to ethylene. Our results indicate that ethylene produced during submergence is required for the stimulation of growth in submerged floating rice plants.     

Low levels of oxygen promote internodal elongation in floating rice independently of enhanced ethylene production

Submergence induces rapid elongation of internodes in floating rice(Oryza sativa L. cv. Habiganj Aman II). The initial signalfor such internodal elongation has been considered to be the reduced partialpressure of oxygen in submerged internodal cavities, which promotes theelongation of internodes through the enhancement of ethylene synthesis. Weexamined the relationship between low oxygen pressure and ethylene production inthe rapid elongation of floating rice internodes using ethylene biosynthesisinhibitors, aminooxyacetic acid (AOA) and CoCl₂. When floating ricestem segments were incubated in an atmosphere of low O₂, internodalelongation accelerated and ethylene production increased. However, in stemsegments treated with AOA or CoCl₂, low O₂ stillstimulated the elongation of internodes although the ethylene production by theinternodes was less than by those in control stem segments where internodalelongation was not promoted. These results indicate that low O₂ iscapable of causing rapid elongation of internodes of floating rice independentlyof enhanced production of ethylene. In addition to low O₂,submergence, ethylene and gibberellic acid each enhanced the production ofethylene by internodal tissues, suggesting that enhanced ethylene production isa common phenomenon accompanied by the acceleration of internodal elongation infloating rice.     

Regulation of growth in stem sections of deep-water rice

Submergence in water greatly stimulates internodal elongation in excised stem sections of deep-water rice (Oryza sativa L. cv. Habiganj Aman II) and inhibits growth of leaf blades and leaf sheaths. The highest rates of internodal growth have been observed in continuous light. Very little growth occurs in submerged sections kept in darkness or incubated under N₂ in the light. The effect of submergence on the growth of deep-water rice is, at least in part, mediated by C₂H₄, which accumulates in the air spaces of submerged sections. This accumulation results from increased C₂H₄ synthesis in the internodes of submerged sections and reduced diffusion of C₂H₄ from the tissue into the water. Increased C₂H₄ levels accelerate internodal elongation and inhibit the growth of leaves. Compounds capable of changing the rate of C₂H₄ synthesis, namely aminoethoxyvinylglycine, an inhibitor of C₂H₄ synthesis, and 1-aminocyclopropane-1-carboxylic acid, the immediate, precursor of C₂H₄, have opposite effects on growth of internodes and leaves. The enhancement of internodal elongation by C₂H₄ is particularly pronounced in an atmosphere of high CO₂ and low O₂. The increase in C₂H₄ synthesis in internodes of submerged sections is primarily triggered by reduced atmospheric concentrations of O₂. The rate of C₂H₄ evolution by internodes isolated from stem sections and incubated in an atmosphere of low O₂ is up to four times greater than that of isolated internodes incubated in air. In contrast, C₂H₄ evolution from the leaves is reduced under hypoxic conditions. The effect of submergence on growth of stem sections of deep-water rice can be mimicked by exposing non-submerged sections to a gas mixture which is similar to the gaseous atmosphere in the internodal lacunae of submerged sections, namely 3% O₂, 6% CO₂, 91% N₂ (by vol.) and 1 l l^-1 C₂H₄. Our results indicate that growth responses obtained with isolated rice stem sections are similar to those of intact deep-water rice plants.Abbreviations ACC 1-aminocyclopropane-1-carboxylic acid - AVG aminoethoxyvinylglycine     

Changes in expansin activity and cell wall susceptibility to expansin action during cessation of internodal elongation in floating rice

We investigated the involvement of expansin action in determining the growth rate of internodes of floating rice (Oryza sativa L.). Floating rice stem segments in which rapid internodal elongation had been induced by submergence for 2 days were exposed to air or kept in submergence for 2 more days. Both treatments reduced the elongation rate of the internodes, and the degree of reduction was much greater in air-exposed stem segments than in continually submerged segments. These rates of internodal elongation were correlated with acid-induced cell wall extensibility and cell wall susceptibility to expansins in the cell elongation zone of the internodes, but not with extractable expansin activity. These results suggest that the reduced growth rate of internodes must be due, at least in part, to the decrease in acid-induced cell wall extensibility, which can be modulated through changes in the cell wall susceptibility to expansins rather than through expansin activity. Analysis of the cell wall composition of the internodes showed that the cellulosic and noncellulosic polysaccharide contents increased in response to exposure to air, but they remained almost constant under continued submergence although the cell wall susceptibility to expansins gradually declined even under continued submergence. The content of xylose in noncellulosic neutral sugars in the cell walls of internodes was closely and negatively correlated with changes in the susceptibility of the walls to expansins. These results suggest that the deposition of xylose-rich polysaccharides into the cell walls may be related to a decrease in acid-induced cell wall extensibility in floating rice internodes through the modulation of cell wall susceptibility to expansins.     

The biophysical basis of elongation growth in internodes of deepwater rice

Partial submergence induces rapid internodal elongation in deepwater rice (Oryza sativa L., cv Habiganj Aman II). We measured in vivo extensibility, tissue tension, hydraulic conductance and osmotic potential in the region of cell elongation in the uppermost internode. The in vivo extensibility of the internode, measured by stretching of living tissue with a custom-made constant stress extensiometer, rose rapidly following submergence of the plant. Both the elastic (E_el) and plastic (E_pl) extensibility increased when growth of the internode was induced. The submerged internode displayed tissue tension (elastic outward bending of longitudinally split internode sections); in air-grown control internodes, no such bending occurred. The hydraulic conductance, estimated from the kinetics of tissue shrinkage in 0.5 molar mannitol and subsequent swelling in distilled water, was not changed by submergence. The osmotic potential, measured with a dew-point hygrometer using frozen-thawed tissue, was only 18% less negative in the submerged internode than in the air-grown control. This indicates that osmoregulation takes place in rapidly elongating rice internodes. We suggest that the rapid expansion of the newly formed internodal cells of submerged plants is controlled by the yielding properties (E_pl) of the cell walls. Experiments with excised stem sections indicate that gibberellin is involved in increasing the E_pl of the elongating cell walls.     

Submergence Enhances Expression of a Gene Encoding 1-Aminocyclopropane-1-Carboxylate Oxidase in Deepwater Rice

Partial submergence greatly stimulates internodal growth indeepwater rice (Oryza sativa L.). Previous work has shown thatthe effect of submergence is, at least in part, mediated byethylene, which accumulates in the air spaces of submerged internodes.To investigate the expression of the genes encoding ethylenebiosynthetic enzymes during accelerated growth of deepwaterrice, we cloned a 1-aminocyclopropane- 1-carboxylate (ACC) oxidasecDNA (OSACO1) from internodes of submerged plants and measuredthe activity of the enzyme in tissue extracts with an improvedassay. We found an increase in ACC oxidase mRNA levels and enzymeactivity after 4 to 24 h of submergence. Thus, it is likelythat ethylene biosynthesis in internodes of deepwater rice iscontrolled, at least in part, at the level of ACC oxidase. (Received January 6, 1996; Accepted April 6, 1996)     

On the role of abscisic Acid and gibberellin in the regulation of growth in rice

Submergence induces rapid elongation of rice coleoptiles (Oryza sativa L.) and of deepwater rice internodes. This adaptive feature helps rice to grow out of the water and to survive flooding. Earlier, we found that the growth response of submerged deepwater rice plants is mediated by ethylene and gibberellin (GA). Ethylene promotes growth, at least in part, by increasing the responsiveness of the internodal tissue to GA. In the present work, we examined the possibility that increased responsiveness to GA was based on a reduction in endogenous abscisic acid (ABA) levels. Submergence and treatment with ethylene led, within 3 hours, to a 75% reduction in the level of ABA in the intercalary meristem and the growing zone of deepwater rice internodes. The level of GA₁ increased fourfold during the same time period. An interaction between GA and ABA could also be shown by application of the hormones. ABA inhibited growth of submerged internodes, and GA counteracted this inhibition. Our results indicate that the growth rate of deepwater rice internodes is determined by the ratio of an endogenous growth promoter (GA) and a growth inhibitor (ABA). We also investigated whether ABA is involved in regulating the growth of rice coleoptiles. Rice seedlings were grown on solutions containing fluridone, an inhibitor of carotenoid and, indirectly, of ABA biosynthesis. Treatment with fluridone reduced the level of ABA in coleoptiles and first leaves by more than 75% and promoted coleoptile growth by more than 60%. Little or no enhancement of growth by fluridone was observed in barley, oat, or wheat. The involvement of ABA in determining the growth rate of rice coleoptiles and deepwater rice internodes may be related to the semiaquatic growth habit of this plant.     

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.     

Ethylene and Gibberellin: Regulation of Internodal Elongation and Nodal Root Development in Floating Rice

Ethylene and GA₃ stimulated internodal elongation in the excisedstem sections of floating rice. The combined application ofethylene and GA₃ exerted a cooperative effect on internodalelongation, although the effect was variety dependent. Stimulativeeffect of ethylene on internodal growth in intact floating riceplants was virtually absent when the plants were pre-treatedwith Ancymidol, -cyclopropyl--(4-methoxyphenyl)-5-pyrimidinemethanol, an inhibitor of gibberellin biosynthesis. Submergenceof intact plants, which also induced internodal elongation,had no stimulative effect when the plants were pre-treated withAncymidol. Submergence of intact plants increased the endogenousgibberellin level. The internode of young, intact 9 day oldseedlings responded neither to submergence nor ethylene, butwhen seedlings were pre-treated with GA₃ they responded to eitherone. Nodal root development was also enhanced by either ethyleneor GA₃. Combined application of ethylene and GA₃ exerted a co-operativeeffect on nodal root development. Ancymidol-treated plants didnot produce nodal roots even though they were subjected to submergence,whereas nontreated control plants produced nodal roots normally. (Received September 12, 1984; Accepted February 15, 1985)     

Amylase activity and growth in internodes of deepwater rice

Isoelectrofocusing, product analysis, thermal denaturation studies and affinity chromatography on cycloheptaamylose-Sephadex were used to identify the amylolytic enzymes in internodes of deepwater rice (Oryza sativa L.). Amylolytic activity in internodes of deepwater rice consists of -amylase (sometimes separated into two isoforms) and of -amylase. During submergence of whole plants, -amylase activity increases in young, growing internodes, but -amylase activity declines. Although non-growing, mature internodes contain higher levels of -amylase than do the elongating younger internodes, the effect of submergence on amylase activities in both tissues follows the same trend. Submergence, gibberellic acid (GA₃) and ethylene all promote -amylase activity in growing and non-growing internodes of excised deepwater-rice stem sections. Inhibitor studies showed that submergence and ethylene promote -amylase activity in the absence of endogenous gibberellin (GA), and GA₃ enhances -amylase activity when ethylene action is inhibited. Therefore, ethylene and GA appear to increase -amylase activity independently of each other. Enhanced -amylase activities are probably responsible for the mobilization of carbohydrates which are needed to support internode elongation during submergence of deepwater rice.Abbreviations CHA cycloheptaamylose - GA₃ gibberellic acid - NBD 2,5-norbornadiene - TCY tetcyclacis     

深水稻节间伸长生长的机制

淹水可促进深水稻节间快速伸长生长,其主要受内源赤霉素、乙烯、脱落酸等激素信号分子的调控。淹水能促进深水稻植物株体内乙烯、赤霉素的生物合成、抑制脱落酸的生物合成,外源乙烯、赤霉素会加速深水稻节间伸长,而外源脱落酸抑制淹水节间的伸长,其中赤霉素是直接作用因子,乙烯能降低内源脱落酸水平、增加节地赤霉素的敏感性;还与渗透调节、细胞壁组份如膨胀素等有关,淹水及赤霉素都大大增加了膨胀素基因的表达。并就深水稻的     

深水稻节间伸长生长的机制

淹水可促进深水稻节间快速伸长生长,其主要受内源赤霉素、乙烯、脱落酸等激素信号分子的调控。淹水能促进深水稻植株体内乙烯、赤霉素的生物合成、抑制脱落酸的生物合成,外源乙烯、赤霉素会加速深水稻节间伸长,而外源脱落酸抑制淹水节间的伸长,其中赤霉素是直接作用因子,乙烯能降低内源脱落酸水平、增加节间对赤霉素的敏感性;还与渗透调节、细胞壁组份如膨胀素等有关,淹水及赤霉素都大大增加了膨胀素基因的表达。并就深水稻的进一步研究进行了展望。     

The effect of submergence,ethylene and gibberellin on polyamines and their biosynthetic enzymes in deepwater-rice internodes

Submergence and treatment with ethylene or gibberellic acid (GA₃) stimulates rapid growth in internodes of deepwater rice (Oryza sativa L. cv. Habiganj Aman II). This growth is based on greatly enhanced rate of cell-division activity in the intercalary meristem (IM) and on increased cell elongation. We chose polyamine biosynthesis as a biochemical marker for cell-division activity in the IM of rice stems. Upon submergence of the plant, the activity of S-adenosylmethionine decarboxylase (SAMDC; EC 4.1.1.50) in the IM increased six- to tenfold within 8 h; thereafter, SAMDC activity declined. Arginine decarboxylase (ADC; EC 4.1.1.19) showed a similar but less pronounced increase in activity. The activity of ornithine decarboxylase (ODC; EC 4.1.1.17) in the IM was not affected by submergence. The levels of putrescine and spermidine also rose in the IM of submerged, whole plants while the concentration of spermine remained low. The increase in SAMDC activity was localized in the IM while the activity of ADC rose both in the node and the IM above it. The node also contained low levels of ODC activity which increased slightly following submergence. Increased activities of polyamine-synthesizing enzymes in the nodal region of submerged plants probably resulted from the promotion of adventitious root formation in the node. Treatment of excised rice-stem sections with ethylene or GA₃ enhanced the activities of SAMDC and ADC in the IM and inhibited the decline in the levels of putrescine and spermidine. We conclude that SAMDC and perhaps also ADC may serve as biochemical markers for the enhancement of cell-division activity in the IM of deepwater rice.Abbreviations ADC arginine decarboxylase - GA gibberellin - IM intercalary meristem - ODC ornithine decarboxylase - SAM S-adenosylmethionine - SAMDC SAM decarboxylase     

Ethylene enhances gibberellin levels and petiole sensitivity in flooding-tolerant Rumex palustris but not in flooding-intolerant R. acetosa

The role of gibberellin (GA) and ethylene in submergence-induced petiole elongation was studied in two species of the genus Rumex. Analysis of endogenous GAs in the flooding-tolerant Rumex palustris Sm. and the intolerant Rumex acetosa L. by gas chromatography-mass spectrometry showed for both species the presence of GA₁, GA₄, GA₉, GA₁₉, GA₂₀ and GA₅₃. Gas chromatography-mass spectrometry analysis of R. palustris petiole tissue of submerged plants showed an increase in levels of 13-OH GAs, especially GA₁, compared with drained plants. This effect could be mimicked by application of 5 μL L⁻¹ ethylene. In R. acetosa, no differences between levels of GAs in drained or submerged plants were found. In R. palustris, both submergence and ethylene treatment sensitized petioles to exogenous gibberellic acid (GA₃). In R. acetosa the effect was opposite, i.e. submergence and ethylene de-sensitized petioles to GA₃. Our results demonstrate the dual effect of ethylene in the submergence response related to flooding tolerance, i.e. in the flooding-tolerant R. palustris ethylene causes an increased concentration of and sensitivity to GA with respect to petiole elongation while in the intolerant R. acetosa ethylene reduces growth independent of GAs. Received: 5 November 1996 / Accepted: 8 February 1997     

Comparison of the role of gibberellins and ethylene in response to submergence of two lowland rice cultivars, Senia and Bomba

We examined the gibberellin (GA) and ethylene regulation of submergence-induced elongation in seedlings of the submergence-tolerant lowland rice (Oryza sativa L.) cvs Senia and Bomba. Elongation was enhanced after germination to facilitate water escape and reach air. We found that submergence-induced elongation depends on GA because it was counteracted by paclobutrazol (an inhibitor of GA biosynthesis), an effect that was negated by GA₃. Moreover, in the cv Senia, submergence increased the content of active GA₁ and its immediate precursors (GA₅₃, GA₁₉ and GA₂₀) by enhancing expression of several GA biosynthesis genes (OsGA20ox1 and -2, and OsGA3ox2), but not by decreasing expression of several OsGA2ox (GA inactivating genes). Senia seedlings, in contrast to Bomba seedlings, did not elongate in response to ethylene or 1-aminocyclopropane-1-carboxylic-acid (ACC; an ethylene precursor) application, and submergence-induced elongation was not reduced in the presence of 1-methylcyclopropene (1-MCP; an ethylene perception inhibitor). Ethylene emanation was similar in Senia seedlings grown in air and in submerged-grown seedlings following de-submergence, while it increased in Bomba. The expression of ethylene biosynthesis genes (OsACS1, -2 and -3, and OsACO1) was not affected in Senia, but expression of OsACS5 was rapidly enhanced in Bomba upon submergence. Our results support the conclusion that submergence elongation enhancement of lowland rice is due to alteration of GA metabolism leading to an increase in active GA (GA₁) content. Interestingly, in the cv Senia, in contrast to cv Bomba, this was triggered through an ethylene-independent mechanism.     

Expansins and Internodal Growth of Deepwater Rice

The distribution and activity of the cell wall-loosening protein expansin is correlated with internodal growth in deepwater rice (Oryza sativa L.). Acid-induced extension of native cell walls and reconstituted extension of boiled cell walls were confined to the growing region of the internode, i.e. to the intercalary meristem (IM) and the elongation zone. Immunolocalization by tissue printing and immunoblot analysis, using antibody against cucumber expansin 29 as a probe, confirmed that rice expansin occurred primarily in the IM and elongation zone. Rice expansin was localized mainly around the vascular bundles at the base of the IM and along the inner epidermal cell layer surrounding the internodal cavity. Submergence greatly promoted the growth of rice internodes, and cell walls of submerged internodes extended much more in response to acidification than did the cell walls of air-grown internodes. Susceptibility of cell walls to added expansin was also increased in submerged internodes, and analysis by immunoblotting showed that cell walls of submerged internodes contained more expansin than did cell walls of air-grown internodes. Based on these data, we propose that expansin is involved in mediating rapid internodal elongation in submerged deepwater rice internodes.