Carbon isotope ratios in ear parts of triticale : influence of grain filling

Four triticale (×Triticosecale Wittmack) genotypes were grown under rainfed conditions with limited irrigation support in Lleida in northeast Spain. For each variety, samples consisting of 10 tillers with half-sterilized spikes were taken three times from anthesis to maturity. Carbon isotope ratios (δ¹³C) were then determined in water extracts from ear bracts (glumes, paleas, and lemmas), awns and flag leaves, and in powdered kernels. For the half-sterilized spikes, carbon isotope analysis was carried out separately in bracts and awns from fertile and nonfertile spikelets. The δ¹³C in the water-soluble fraction of awns, glumes, and glumells from fruitless spikelets was significantly higher than that from fertile spikelets sampled at mid-grain filling. Differences in δ¹³C among sterile and fertile spikelets were not significant in samples taken a few days after anthesis or at maturity. These results are in accordance with some degree of refixation by awns and ear bracts of the CO₂ respired by grains during grain filling. There was progressively higher δ¹³C from flag leaf blades to awns, glumes, and glumells. This variation in δ¹³C along plant parts may be caused by differences in the ratio of assimilation rate to CO₂-diffusive conductance. Values of δ¹³C of mature kernels were between the values at anthesis and mid-grain filling for the water-soluble fraction of flag leaves and inner bracts and were fairly similar to those of glumes and awns.     

Photosynthesis and Transpiration in Leaves and Ears of Wheat and Barley Varieties

Carbon exchange rate (CER) and transpiration were measured inflag leaves, whole ears, glumes (referring to the total areaof glumes and lemmas) and awns, in six hexaploid spring wheats(Triticum aestivum L.), three cultivated tetraploid spring wheats(T. turgidum L.), four wild tetraploid wheats (T. dicoccoides),eight six-rowed barleys (Hordeum vulgare L.) and five two-rowedbarleys (H. vulgare L.). Differences between varieties and between species in total earCER and transpiration were associated largely with differencesin ear surface area rather than with rates per unit area. Ratesof CER and transpiration per unit area of ears were 40–80%of those of flag leaves, depending on the species. However, since ear surface area was greater than flag leaf areaby a factor of 1.1, 3.9, 5.5 and 4.4, in hexaploid wheat, tetraploidwheat, six-rowed barley, and two-rowed barley, respectively,total ear CER reached up to 90% of that of the flag leaf. The contribution of awns to total ear CER depended largely ontotal awn surface area per ear, rather than on CER per unitawn area. Awns contributed about 40–80% of total spikeCER, depending on the species, but only 10–20% of spiketranspiration. The disproportionately small contribution ofawns to ear transpiration was caused by the very low rate oftranspiration per unit area of awns. Thus, while transpirationratio (CER/transpiration) was about the same in flag leavesand glumes, it was higher by several orders of magnitude inthe awns. A large amount of awns in the ear is therefore a drought adaptiveattribute in these cereals, for which tetraploid wheat exceededhexaploid wheat and six-rowed barley exceeded two-rowed barley. Key words: Carbon exchange rate, Transpiration, Barley, Wheat     

Photosynthetic pigment composition and photosystem II photochemistry of wheat ears.

The characteristics of pigment composition and photosystem II (PSII) photochemistry in the flag leaf and ear parts of wheat (Triticum aestivum L.) grown in the field was compared. At the early stage of flowering, awns and the flag leaf showed the highest values in the maximal efficiency of PSII photochemistry (Fv/Fm), actual PSII efficiency (phi(PSII)), photochemical quenching (qP), and the efficiency of excitation capture by open PSII centres (Fv/F'm), followed by glumes, lemmas, and paleae, respectively except that no differences in F'v/F'm were observed among glumes, leamms, and paleae. With progressing grain filling, there was a change in the photosynthetic pigment stoichiometry. In the ear parts, neoxanthin and antheraxanthin decreased equally with chlorophyll levels. Lutein and zeaxanthin decreased less than chlorophyll levels while beta-carotene and violaxanthin decreased faster than chlorophyll levels. No big differences in pigment composition were observed among different ear parts. For the flag leaf, neoxanthin and beta-carotene decreased concomitantly with chlorophyll, whereas lutein and xanthophyll cycle pigment were less affected, leading to increases in lutein/chlorophyll and xanthophyll cycle pigment/chlorophyll ratios. Fv/Fm, phi(PSII), qP, and F'v/F'm decreased gradually in the flag leaf and ear parts but to different extents. The largest changes were observed in awns, followed by the lemmas of floret 2, the lemmas of floret 1, glumes, and the flag leaf, respectively. The results suggest that during grain filling, a down-regulation of PSII associated with an increase of the de-epoxidation state of the xanthophyll cycle carotenoids occurred in the flag leaf but not in the ear parts.     

Ear photosynthesis, carbon isotope discrimination and the contribution of respiratory CO2 to differences in grain mass in durum wheat

The role of ear photosynthesis in grain filling was studied in a number of durum wheat (Triticum turgidum var durum L.) landraces and varieties from the Middle East, North Africa, and from the collections of ‘Institut National de la Recherche Agronomique’ (INRA, France) and ‘Centro International de Mejora de Maiz y Trigo’ (CIMMYT, Mexico). Plants were grown in the field in a Mediterranean climate. Flag leaves (blade plus sheath) and ears were kept in the dark from 1 week after anthesis to maturity which reduced grain weight by 22.4% and 59.0%, respectively. In a further experiment, the carbon isotope discrimination ratio (Δ) of ear bracts, awns and flag leaves was measured on samples taken at anthesis and on mature kernels. The mean value of Δ for the water soluble fraction of bracts (17.0‰) and awns (17.7‰) were lower than those of leaves (19.5‰) and fairly similar to those of kernels (17.4‰) averaged across all genotypes. Data indicate that most of the photosynthates in the grain come from ear parts and not from flag leaves. In addition, a higher water use efficiency (WUE) of ear parts than of the flag leaf is suggested by their lower Δ values. Gas exchange in ears and flag leaves was measured during grain filling. Averaged over all genotypes, CO₂ diffusive conductance was about five times higher in the flag leaf than in the spike (with distal portions of awns outside the photosynthetic chamber) 2 weeks after anthesis. In absolute terms, the dark respiration rate (R_d) was greater than the net photosynthesis rate (P_n) by a factor of 1.74 in the spike, whereas R_d was much smaller, only 22.1, 65.7 and 24.8% of P_n in blade, sheath and awns, respectively. Data indicate that photosynthesis, and hence the water use efficiency (photosynthesis/transpiration), is greatly underestimated in ears because of the high rates of respiration which diminish the measured rates of net CO₂ exchange. Results of ¹³C discrimination and gas exchange show that genotypes from North Africa have higher WUE than those from the Middle East. The high R_d values of ears as well as their low diffusive conductance suggest that CO₂ from respiration may be used as source of carbon for ear photosynthesis. In the same way, the anatomy of glumes, for example, supports the role of bracts using internal CO₂ as source of photosynthesis. In the first experiment, the Δ in mature grains from culms with darkened ears compared with control culms provided further evidence in support of this hypothesis. Thus, the Δ from kernels of control plants was 0.40 higher than that from ear-darkened plants, probably because of some degree of refixation (recycling) of respired CO₂ in the grains.     

Monocarpic senescence in wheat: Influence of sterile glumes and ear

The senescence of sterile glumes, flag leaf and the other two leaves below the ear of wheat ( Triticum aestivum L. cv. Sonalika) was studied in relation to grain development and surgical manipulation. The senescence of sterile glumes was faster than that of the leaves in terms of chlorophyll and protein degradation. The flag leaf senesced later than the other two leaves below it. Removal of sterile glumes markedly reduced the harvest index (crop: straw ratio) and average dry weight per grain as compared to removal of the flag leaf. Maximum grain weight was achieved after the glumes had senesced completely. Removal of the ear delayed senescence of all the three leaves. It is concluded that sterile glumes are important suppliers of assimilate for grain filling and that nutrient drainage is the primary cause of the monocarpic senescence in wheat.     

APETALA 2‐like genes AP2L2 and Q specify lemma identity and axillary floral meristem development in wheat

The spikelet is the basic unit of the grass inflorescence. In tetraploid (Triticum turgidum) and hexaploid wheat (Triticum aestivum), the spikelet is a short indeterminate branch with two proximal sterile bracts (glumes) followed by a variable number of florets, each including a bract (lemma) with an axillary flower. Varying levels of miR172 and/or its target gene Q (AP2L5) result in gradual transitions of glumes to lemmas, and vice versa. Here, we show that AP2L5 and its related paralog AP2L2 play critical and redundant roles in the specification of axillary floral meristems and lemma identity. AP2L2, also targeted by miR172, displayed similar expression profiles to AP2L5 during spikelet development. Loss‐of‐function mutants in both homeologs of AP2L2 (henceforth ap2l2) developed normal spikelets, but ap2l2 ap2l5 double mutants generated spikelets with multiple empty bracts before transitioning to florets. The coordinated nature of these changes suggest an early role of these genes in floret development. Moreover, the flowers of ap2l2 ap2l5 mutants showed organ defects in paleas and lodicules, including the homeotic conversion of lodicules into carpels. Mutations in the miR172 target site of AP2L2 were associated with reduced plant height, more compact spikes, promotion of lemma‐like characters in glumes and smaller lodicules. Taken together, our results show that the balance in the expression of miR172 and AP2‐like genes is crucial for the correct development of spikelets and florets, and that this balance has been altered during the process of wheat and barley (Hordeum vulgare) domestication. The manipulation of this regulatory module provides an opportunity to modify spikelet architecture and improve grain yield.     

Lack of C4 photosynthetic metabolism in ears of C3 cereals

There is continuing controversy over whether a degree of C₄ photosynthetic metabolism exists in ears of C₃ cereals. In this context, CO₂ exchange and the initial products of photosynthesis were examined in flag leaf blades and various ear parts of two durum wheat (Triticum durum Desf.) and two six-rowed barley (Hordeum vulgare L.) cultivars. Three weeks after anthesis, the CO₂ compensation concentration at 210 mmol mol^?1 O₂ in durum wheat and barley ear parts was similar to or greater than that in flag leaves. The O₂ dependence of the CO₂ compensation concentration in durum wheat ear parts, as well as in the flag leaf blade, was linear, as expected for C₃ photosynthesis. In a complementary experiment, intact and attached ears and flag leaf blades of barley and durum wheat were radio-labelled with ¹⁴CO₂ during a 10s pulse, and the initial products of fixation were studied in various parts of the ears (awns, glumes, inner bracts and grains) and in the flag leaf blade. All tissues assimilated CO₂ mainly by the Calvin (C₃) cycle, with little fixation of ¹⁴CO₂ into the C₄ acids malate and aspartate (about 10% or less). These collective data support the conclusion that in the ear parts of these C₃ cereals C₄ photosynthetic metabolism is nil.     

Endophytic Fungi in Four Winter Wheat Cultivars (Triticum aestivum L.) Differing in Resistance Against Stagonospora nodorum (Berk.) Cast. & Germ. =Septoria nodorum (Berk.) Berk.

Four winter wheat cultivars with different levels of resistance to Septoria nodorum were investigated at four locations during two vegetation periods. Forty plants per cultivar and site were collected at random at seven defined growth stages from crop emergence to harvest. Samples from roots, culms, leaves, glumes and kernels were examined for the occurrence of endophytic fungi after surface sterilization. 83% of the 26944 isolates sporulated and were assigned to 213 species. The most frequent were: Septoria nodorum (20.1%), Alternaria tenuissima (9.8%), Epicoccum purpurascens (9.1%), Idriella bolleyi (6.9%), Fusarium graminearum (5.3%), Fusarium culmorum (4.0%), Cladosporium oxysporum (3.7%), Didymella exitialis (3.1%), Fusarium nivale (2.8%) and Rhizoctonia solani (2.1%). Each species occurred preferentially in one or more plant organs. A factorial analysis of variance showed that plant organ, sampling site, vegetation, period and cultivar in decreasing order of importance influenced the quantitative and qualitative composition of the fungal populations. No relationship between endophytic fungi was found to be constantly antagonistic or mutualistic. Septoria nodorum was isolated mainly from culms. The number of S. nodorum isolates differed significantly between cultivars in culms and glumes but not in flag leaves. The results are discussed in relation to resistance breeding and the effect endophytic fungi, might have on yield.     

Gas exchange of ears of cereals in response to carbon dioxide and light

One cultivar each of spring wheat (Triticum aestivum L. cv. Arkas), oat (Avena sativa L. cv. Lorenz), and barley (Hordeum vulgare L. cv. Aramir) was chosen in order to study the relative contributions of individual bracts to the gas exchange of whole ears. The distribution and frequency of the stomata on the bracts were examined. Gas exchange was measured at normal atmospheric CO₂ (330 bar) and at high CO₂ (2000 bar) on intact ears and on ears from which glumes or lemmas and pleae (wheat and oat) or awns (barley) had been removed.The relative contribution to the gas exchange of the whole organ is highest for the awns of barley ears. In wheat, the contribution of the glumes is slightly higher than that of the inner bracts before anthesis. Two weeks after anthesis the inner bracts contribute more than the glumes. This tendency of increasing importance of the inner bracts is also found in oat ears, but the relative amount of CO₂ uptake by the glumes is higher than in wheat. These changes during ontogeny result from the better supply of light to the inner bracts caused by opening of the ears' structures during grain filling, which in part compensates for the decreasing photosynthetic capacity.The ratio of the photosynthesis rate at high CO₂ to that at normal CO₂ is lower for the glumes of oat and for the awns of barley than for the other bracts.Abbreviations A³³⁰, A²⁰⁰⁰ net photosynthesis rate, A³³⁰ at normal atmospheric CO₂ (330 bar), A²⁰⁰⁰ at high CO₂ (2000 bar) - PPFD photosynthetic photon flux density - p^c intercellular partial pressure of CO₂     

RuBP carboxylase activity in wheat following ear emergence and evaluation of the role of ear

Ribulose bisphosphate carboxylase activity and chlorophyll content were measured in the leaves and reproductive parts of two wheat cultivars after ear emergence. The chlorophyll content of the flag leaf was mostly higher than that of the awns and glumes. Awns had the highest chlorophyll content among reproductive parts. Light transmission to the lower leaves was higher in the dwarf cultivar Moti than in the medium tall cultivar Kalyansona. RuBP carboxylase activity in Kalyansona leaves was higher than in Moti leaves. In postanthesis stages there was no difference in RuBP carboxylase activity in the flag leaf and lower leaves between cultivars. Awns had the maximum activity of RuBP carboxylase followed by glumes and grains among ear parts. The relative capacity for photosynthesis in the ear parts was several times higher than in the flag leaf on a unit chlorophyll basis. It is suggested that in a crop canopy in the field, the spike(ear) may have a greater importance in grain development than has been previously estimated.     

Wheat nitrogen metabolism during grain filling: comparative role of glumes and the flag leaf

The mobilization of nitrogen (N) compounds and the roles played by glumes and the flag leaf during grain filling were studied in bread wheat (Triticum aestivum L. cv. Florida) grown under field conditions. Glumes lost twice as much of their total N content as that lost by the flag leaf between the milk and early dough stages. In the flag leaf, glumes and grains, Glu, Asp, Ser and Ala accounted for 85% of all the reductions in the free amino acid pool. Principal component analysis of free amino acid pools separated grains from the glumes and the flag leaf, suggesting grain specific regulations in the use of free amino acids in protein synthesis. In all three organs, no decrease in Gln was detected, probably due to steady glutamine synthetase (GS; EC 6.3.1.2) activities per soluble protein in both the flag leaf and glumes. Compared with the flag leaf, glumes presented relatively smaller amounts of the chloroplast GS associated isoform. This we show is due to a lower relative number of mesophyll cells in glumes as supported by the different anatomy and the cellular pattern of the GS immunolocalization. We argue that cellular distribution plays a key role in supporting metabolism to enable the various functions undertaken by glume tissue.     

Planting density effects on assimilation and partitioning of photosynthates during grain filling in the late-sown wheat

Leaf blades of the late-sown winter wheat produced the major portion, i.e., more than 60 %, of the total ¹⁴C-photosynthates at grain filling, but ear (rachis and glumes) only about 15 %, sheaths about 11 %, and stem internodes about 11 %. The change of plant density in this experiment had little influence on the ¹⁴CO2-photoassimilation of the ear (rachis and glumes), flag leaf lamina, sheaths and stem internodes, but markedly affected photosynthesis of the second, the third and lower leaves. The photosynthetic rate [expressed as specific radioactivity, s-¹ kg-¹(d.m.)] and the amount of ¹⁴CO₂ photosynthates decreased significantly in the second, the third and other lower leaves at a high plant density. Upon grain-filling of the late-sown wheat, the grain was the major importer of photosynthates. Yet partitioning to the stem internodes depended on the plant density. Stem was the importer of photosynthates at a low plant density, but the exporter at a high plant density. In plants at a low plant density a fairly large proportion of photosynthates was distributed into the roots. The middle and lower above-ground parts of the late-sown wheat at a high plant density decreased or lost their function early. As a result, the plant senesced earlier. However, the grain setting, filling and yielding were restricted. An appropriately low plant density was suitable for prolonging the function of the middle and lower organs, delaying the senescence of plant, increasing the source supply for grain filling, and improving the grain yield. This revised version was published online in August 2006 with corrections to the Cover Date.     

Effect of changed source/sink relations on proteolytic activities and on nitrogen mobilization in field-grown wheat (Triticum aestivum L.)

Nitrogen mobilization and the pattern of proteolytic enzymeswere investigated in leaves and glumes of field-grown winterwheat (Triticum aestivum L.) during maturation. Source/sinkrelations were changed by removal of the ear, the flag leafor the lower leaves shortly after anthesis. Removal of the earwas most effective, resulting in delayed senescence of the flagleaf with the chlorophyll, aminopeptidase and carboxypeptidaseactivities remaining high in contrast to the control, whereasneutral endopeptidase activity increased more slowly. No majorchanges were observed in the second leaf from the top in plantswith either ears or flag leaves removed. Nitrogen mobilizationand proteolytic activities in glumes and the remaining leaveswere influenced only slightly by leaf removal. In earless plants,nitrogen was transported from the second leaf into the leafsheath and stem, but in the flag leaf the total reduced nitrogenremained high and free amino groups increased. The increase in endopeptidase activity was influenced by thesource/sink relations. However, the accumulation of amino groupsand the increasing endopeptidase activity in the flag leaf ofearless plants suggest that the nitrogen sink capacity did notgreatly control protein degradation; it remains to be seen whetherphytohormones, accumulated amino acids or other factors delayedthe increase in endopeptidase activity. (Received September 3, 1979; )     

Persistence of photosynthetic components and photochemical efficiency in ears of water-stressed wheat (Triticum aestivum)

Ear photosynthesis may be an important source of C for grain growth in water-stressed plants of cereals. The main objectives of this work were to determine the stability of the photosynthetic apparatus and the photochemical efficiency of ears in plants subjected to post-anthesis drought. Plants of wheat ( Triticum aestivum L. cv. Granero INTA) were grown in pots under a rain shelter and subjected to water stress (soil water potential around −0.6 to −0.8 MPa) starting 4  days after anthesis. Post-anthesis drought substantially accelerated the loss of chlorophyll, Rubisco and the light-harvesting complex of photosystem II (LHCII) in the flag leaf, but the degradation of these photosynthetic components was much less affected by water deficit in awns and ear bracts. Quantum yield of PSII (Φ_PSII) decreased in leaves of water-stressed plants. In contrast, ear bracts had a higher Φ_PSII than leaves, and Φ_PSII of ear bracts did not decrease at all in response to drought. Removing the grains immediately before fluorescence measurements (less than 30 min) slightly reduced Φ_PSII, indicating that CO₂ supplied by grain respiration may contribute to the high photochemical efficiency of ears in droughted plants. However, other factors may be involved in maintaining high Φ_PSII, since even in the absence of grains Φ_PSII remained much higher in ear bracts than in the flag leaf. The relative stability of ear photosynthetic components and their relatively high photochemical efficiency may help to maintain ear photosynthesis during the grain filling period in droughted plants.     

Epidermal conductance in different parts of durum wheat grown under Mediterranean conditions: the role of epicuticular waxes and stomata

Abstract. Epidermal (non-stomatally-controlled) conductance from the fourth leaf, first node leaf, flag leaf and ear of durum wheat (Triticum turgidum var durum L.) grown under Mediterranean field conditions has been measured, along with leaf stomatal frequency and the amount and distribution of epicuticular waxes. Measurements were carried out on varieties and land-races from the Middle East, North Africa, ‘Institut National de la Recherche Agricole’ (INRA) and ‘Centra Internacional de Mejora de Maiz y Trigo’ (CIMMYT). Significant differences were observed among genotypes in the epidermal conductances (g_e) of the four organs. For each of the four organs tested, genotypes from the Middle East and CIMMYT showed higher g_e. values than those from North Africa and INRA. Ears showed epidermal conductances that were more than four times higher than those of leaves when g_e. values were expressed per unit dry weight. The amount of epicuticular waxes was higher in the fourth leaves, intermediate in the first node and flag leaves and lower in the ears. For each organ, g_e differences among genotypes were unrelated with the amount of epicuticular waxes. Removal of epicuticular waxes by dipping the organs into chloroform significantly increased the epidermal conductance for the fourth and first node leaves and the ear. However, this did not occur for the flag leaf. For the fourth leaf, g_e of intact leaves and g_e of leaves in which epicuticular waxes were removed were unrelated (r = -0.265). The regression coefficient of this relation for the first node and flag leaves showed values of 0.666 and 0.650 (P > 0.05), respectively, and values were even higher in the ear (r > m 0.892, P > 0.01). Scanning electron microscope analysis showed that wax bloom decreased from the fourth leaf to the flag leaf, whereas the extent of amorphous wax increased. Wax bloom in leaves consisted mainly of deposits of thin wax plates. In the ears and the adaxial surface of flag leaves, fibrillar waxes predominated. In the first node and flag leaves, the wax deposits on the adaxial side cover the surface of the leaf more densely and uniformly than those on the abaxial side. There was no significant correlation between g_e and total stomatal density, or between g_e and either adaxial or abaxial stomatal density for any sample of the three different leaves. The contribution of epicuticular waxes plus total stomatal frequency only explained 42.4, 11.8, 28.3 and 16% of g_e (per unit leaf area) variations for the fourth leaf, first node leaf, flag leaf and the combined variation of the three leaves together, respectively. From these results, it is concluded that complex interrelationship between different morphophysiological characteristics probably control g_e differences among genotypes and that these interrelationships differ for each different plant part.     

Free Polyamines in Senescing Wheat(Triticum aestivum L.)

The free polyamine content of flag leaves, peduncles, rachis,glumes, and grains of wheat (Triticum aestivum L., cv. Castell)plants, ripening under field conditions, has been investigatedduring three consecutive growing seasons. Putrescine was quantitativelythe most important of all polyamines detected in these organs.Concentrations were highest in the grains, glumes and flag leaves.No correlation was found between polyamine content and the onsetof senescence of flag leaves and other organs. Excised primaryleaves, however, showed a decrease in polyamine content in thedark and also in light/dark cycles, but in the latter case onlyafter an initial increase. Sink removal of otherwise intactwheat plants caused an accumulation of putrescine in flag leavesat the later stages of senescence, whereas removal of all otherleaves was without any significant effect. Putrescine was alsorecovered in phloem-exudate samples collected throughout theperiod of grain development. In both grains and glumes, peakconcentrations of polyamines were found early during seed development. Key words: Triticum aestivum, polyamines, ripening, senescence     

Optical and anatomical characteristics of bracts from the Chinese "glasshouse" plant, Rheum alexandrae Batalin (Polygonaceae), in Yunnan, China

Bracts that lacked chlorophyll were compared with rosette leaves on the Chinese glasshouse plant Rheum alexandrae Batalin. The structures were analyzed anatomically and with photospectrometry. Histological features were significantly different between the bracts and the rosette leaves. Epidermal pavement cells and palisade cells were larger in the bracts than in the rosette leaves, but the ratio of the intercellular spaces to the cells in the palisade layer was the same in the two structures. Absorption spectrum analyses of the half-translucent bracts showed that, despite their thinner structure, they absorbed ultraviolet (UV) radiation more efficiently than did the green rosette leaves. The characteristics of the R. alexandrae bracts were different from those of the Himalayan glasshouse plant Rheum nobile. These results suggest that R. alexandrae evolved bracts as an adaptive strategy to protect the inflorescence from UV radiation. Received: August 22, 2001 / Accepted: October 15, 2001     

ELE restrains empty glumes from developing into lemmas

Although there is evident homology among reproductive organs when comparing Poaceae(grass)and eudicots,the identity of grass specific organs,such as lodicules,palea,lemma,and glumes has been the subject of a vast and largely inconclusive discussion.Here we provide some direct evidence to support the idea that the empty glumes of rice(Oryza sativa)are counterparts of lemmas.We show that the development of empty glumes is regulated by ELE(elongated empty glume),which belongs to a plant specific novel gene family.Mutations at the ELE locus cause elongated empty glumes,which mimic the lemmas and have the epidermal morphology of lemmas with four or five vascular bundles.As a nuclear-localized gene,ELE is specifically expressed at the empty glumes of immature spikelets,and its ectopic expression causes many floral development defects,including lemma-like palea,extra palea-like structures,elongated lodicules,extra stamens and stigmas.Our result suggests that empty glumes are lemmas of the sterile florets located at the lateral side of the rice spikelet,and ELE acts as a regulator restraining its growth to maintain its small size in wild-type plants.     

Photosynthesis and transpiration of the flag leaf in four spring-wheat cultivars

Co₂ exchange and transpiration rates of the flag leaves of four spring wheat (Triticum aestivum L.) cultivars, namely Glenlea, Neepawa, Opal and Kolibri, were compared using infra-red gas-analysis technique. The plants were grown in a controlled environment under an 18-h photoperiod, with day and night temperatures of 20 and 15° C, respectively. The time course of the CO₂-exchange rate (CER) of the flag leaf differed among cultivars. CER began to decrease rapidly some 2 weeks after ear emergence in Glenlea, Neepawa and Kolibri, but only after 4 weeks in Opal. The decline in CER of Glenlea, Neepawa and Opal was continuous throughout the period of grain development whereas in Kolibri CER was maintained at a constant level between the 4th and 6th weeks after ear emergence. The transpiration rates of the flag leaves of the 4 cultivars did not change markedly until 6–7 weeks after ear emergence, indicating that the reduction in CER was not primarily a response to increased stomatal resistance to the diffusion of CO₂. Removing the ear of the main shoot of intact plants failed to depress CER of the subtending flag leaf until 5 weeks after ear removal. Removing the ears of all the tillers of plants in which all but 3 tillers had been removed at ear emergence did not depress CER until 4 weeks after ear emergence, but removal of the ear of the main shoot of plants where all the tillers had been removed at ear emergence reduced the CER of the flag leaf 2 weeks after ear removal. Removal of tillers at ear emergence had a marked effect on the time course of CER and transpiration rates of the flag leaf. Both CER and transpiration rates of a 4-tiller plant were maintained at a higher level throughout ear development as compared to those of a one-tiller plant. The transpiration rate of the flag leaf of Glenlea increased during the later part of the life of the leaf even for one-tiller plants with no ear, indicating that such a stomatal response may be part of the normal course of leaf aging and not a response to a feedback stimulus from the ear.