Growth under elevated atmospheric CO(2) concentration accelerates leaf senescence in sunflower (Helianthus annuus L.) plants

Some morphogenetic and metabolic processes were sensitive to a high atmospheric CO(2) concentration during sunflower primary leaf ontogeny. Young leaves of sunflower plants growing under elevated CO(2) concentration exhibited increased growth, as reflected by the high specific leaf mass referred to as dry weight in young leaves (16days). The content of photosynthetic pigments decreased with leaf development, especially in plants grown under elevated CO(2) concentrations, suggesting that high CO(2) accelerates chlorophyll degradation, and also possibly leaf senescence. Elevated CO(2) concentration increased the oxidative stress in sunflower plants by increasing H(2)O(2) levels and decreasing activity of antioxidant enzymes such as catalase and ascorbate peroxidase. The loss of plant defenses probably increases the concentration of reactive oxygen species in the chloroplast, decreasing the photosynthetic pigment content as a result. Elevated CO(2) concentration was found to boost photosynthetic CO(2) fixation, especially in young leaves. High CO(2) also increased the starch and soluble sugar contents (glucose and fructose) and the C/N ratio during sunflower primary leaf development. At the beginning of senescence, we observed a strong increase in the hexoses to sucrose ratio that was especially marked at high CO(2) concentration. These results indicate that elevated CO(2) concentration could promote leaf senescence in sunflower plants by affecting the soluble sugar levels, the C/N ratio and the oxidative status during leaf ontogeny. It is likely that systemic signals produced in plants grown with elevated CO(2), lead to early senescence and a higher oxidation state of the cells of these plant leaves.     

Plant growth in relation to ontogenetic changes in productive characteristics of individual leaves I. growth and productive characteristics ofHelianthus annuus andZinnia elegans leaves

Growth and productive characteristics of successive leaves were ontogenetically examined using sunflower and zinnia plants grown in the field. For the purpose of more advanced comprehension, ontogenetic behavior of productive characteristics was formulated with the schematic leaf growth model. The largest leaf in zinnia appeared around the middle leaf order, but did not show the highest photosythetic rate. The decreasing pattern of relative leaf area expansion rate did not significantly differ among individual leaves, and seemed inherent to species. The ontogenetic changes in net photosynthetic rate per unit leaf area showed convex curves, while those in dark respiratory rate per unit leaf area rapidly decreased with leaf development and then became constant. In part, the rapid increase of photosynthetic rate in young leaves was supported by enhancement of light utilization efficiency, along with increase of chlorophyll content. The approach of leaf angle to the horizontal was more or less accompanied by photosynthetic development and leaf expansion. It was suggested that photosynthetic maturation in leaves of the sun-leaf type appears at leaf age equivalent to 0.35 to 0.46 of leaf life span. Ontogenetic pattern of all productive characteristics basically differed little among successive leaves.     

Translocation of Radioactive Carbon after the Application of C-Alanine and CO(2) to Sunflower Leaves

(14)C-(UL)-l-Alanine was applied to the surface of mature leaves at the second node of sunflower (Helianthus annuus L. cv Commander) plants, under illumination. The alanine was absorbed during a 4-hour period, and some of it was metabolized by the absorbing tissue. After a lag period of about 15 minutes from first application, distribution of (14)C through the plant proceeded in much the same pattern as when (14)CO(2) is assimilated by similar leaves. Most, if not all, of the (14)C exported from the absorbing regions was in sucrose. Only minute amounts appeared in alanine or other amino acids in surrounding parts of the leaf blade or in the petiole, although these were strongly labeled in the tissue absorbing (14)C-alanine.When (14)CO(2) was supplied for 15 minutes to leaves of different ages, amino acids were lightly labeled in the leaf blade. Mature green leaves exported only sucrose. Yellowing leaves on 60-day-old plants exported a variety of substances including amino acids.     

The polarity of endogenous regulatory substances inBryophyllum crenatum leaves and stems

In isolated leaves ofBryophylluni crenatum the intensity of marginal bud formation decreases from the apex towards the blade base, which is associated with the decreasing content of enioganous gibberallins. As proved by Dostál (1930), the formation of marginal buds on transversely divided blade of the isolated leaf increases in comparison with the non-divided control leaf. The results of our experiments have revealed that the increase in the formation of marginal buds in the blade transversely divided into the apical, middle and basal parts is connected with the increasing level of endogenous gibberellins, especially in the apical part. This rising level appears as early as 7 days following blade division,i.e. at the time preceding the formation of marginal bud bases. InBryophyllum crenatum plants the level of endogenous cytokinins was estimated in apical, middle and basal leaves, as well as in adjacent internodia. Maximum content was found out in the leaves from the middle stem part, which is probably associated with the capacity of this part to form marginal buds spontaneously also in intact plants. However, prior to flowering the maximum of cytokinin activity is shifted to the apical stem part.     

Leaf and stem growth in the Lepidodendrales

Measurement of leaf length and stem diameter in three Upper Carboniferous l.epidodendron species shows an exponential relationship between these parameters. Larger axes bear longer leaves even in material which had apparently completed its primary' growth. It is suggested that this is a direct product of the determinate growth pattern of the ontogeny of arborescent lycopods. Measurements of Cyclostigma from the Irish Devonian are not explicable in these terms, and it is suggested that some specimens were immature at fossilization. It is recommended that the term leaf cushion be applied to the expanded leaf base in the Lepidodendrales at all growth stages, and not only after leaf abscission. Problems in the terminology of 'young' and 'old' growth stages in these determinate plants are discussed.     

Content of endogenous cytokinins in intact and transversally cut leaf blades of Bryophyllum crenatum

An increased growth of marginal buds on excised leaves from vegetative plants of Bryophyllum crenatum following transversal separation of leaf blades into three segments (apical, medial and basal) as compared with non-separated leaves, was associated with increased endogeneous cytokinin-like substances. This cycokinin increase can be demonstrated seven days after the separation and before the growth of marginal bud primordia. The level of endogenous cytokinin-like substances increases in a similar direction as the formation and growth of marginal buds, from the base of leaf blade to its apex.

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Zusammenfassung Der Anstieg von Marginalknospenbildung an den von den vegetativen Pflanzen Bryophyllum crenatum isolierten Blättern, der durch eine Querteilung der Blattspreiten in drei Segmente (apikales, mittleres und basales) gegenüber den intakten Kontrollblattspreiten nachweisbar ist, ist mit dem Anstieg der endogenen Zytokininähnlichen Stoffe verknüpft. Diesen Anstieg des Zytokininspiegels in der geteilten Blattspreite, gegenüber der intakten, kann man binnen sieben Tage nach der Querteilung der Blattspreite beweisen, d.h. in der Zeit vor dem Wachstum der Marginalknospenanlagen. Der endogene Zytokininspiegel steigt dabei von der Basis zum Apex der Blattspreite an, ähnlich wie die Marginalknospenbildung.

     

Studies of silica in the oat plant

Summary Oat plants,Avena sterilis L., were grown on soils in which the concentration of monosilicic acid in the soil solution, that is the level of supply of silica, ranged from 7 to 67 ppm SiO₂. Analyses at intervals throughout the growing period showed that the level of supply affected the amount and concentration of silica in the plant but not the pattern of its distribution among the parts.At maturity the caryopsis contained only 0.5 to 0.8 per cent of the total silica in the tops while the other parts of the inflorescence contained 40.7 to 41.3 per cent. The leaves (blade and sheath) contained 42.5 to 45.0 per cent of the total silica and the stems contained 7.8 to 10.9 per cent; the remainder was present in small sterile tillers.The concentration of silica in the dry matter was highest in the palea, lemma glumes, awn, and leaves. Among the leaves, the flag leaf had the highest silica content, both in terms of concentration in the dry matter and amount per leaf. The distribution of silica along a leaf followed a hyperbolic curve, the concentration being highest at the apex and lowest at the base of the blade.The chemistry of silica and the pattern of its distribution in the tops suggest that monosilicic acid and water move concomitantly in the transpiration stream and that solid silica is deposited in greatest quantities in those parts and regions from which water is lost in greatest quantities.     

ANAEROBIC ELEVATION OF ETHYLENE CONCENTRATION IN WATERLOGGED PLANTS

Anaerobic elevation of ethylene concentration in waterlogged and non-waterlogged Helianthus annuus L. and Lycopersicon esculentum Mill. was studied. A balloon method was devised to provide an anaerobic atmosphere around the intact sunflower stem. Anaerobic conditions were also produced by bubbling nitrogen into the floodwater. Ethylene concentration in the stem of waterlogged plants was higher when nitrogen was bubbled through the floodwater than when aerated, the effect being greater for the soil-grown plants than for the sand-cultured plants. Ethylene concentration in the stem of waterlogged plants was highest in the region exposed to anaerobiosis, and less with increasing distance or height on the non-waterlogged part of the stems. Intact sunflower stems increased their ethylene concentration in that part of the stem which was maintained in an oxygen-free atmosphere. The results suggest that enhanced ethylene production in waterlogged plants primarily occurs in the waterlogged part of roots and stems.     

Hydraulic architecture of plants of Helianthus annuus L. cv. Margot: evidence for plant segmentation in herbs

The hydraulic architecture of sunflower (Helianthus annuus L. cv. Margot) was studied in terms of the partitioning of the hydraulic conductance (Kleaf) of leaves inserted at progressively more apical nodes both in growing plants (GP) and in plants at full anthesis (mature plants, MP). Leaf conductance to water vapour (gL), leaf water potential (PsiL), leaf water potential at zero turgor (Psi tlp), and leaf osmotic potential at full turgor (pi0) were also measured. Sunflower plants showed gL and Kleaf values significantly increasing in the acropetal direction, while PsiL of basal leaves was significantly more negative than that of distal leaves; Psi tlp markedly decreased in the acropetal direction in MP so that leaves of MP retained increasingly more turgor the more apical they were. This hydraulic pattern, already present in very young plants (GP), strongly favours apical leaves. These data suggest that the progressive leaf dieback starting from the stem base, as observed when the inflorescence of sunflower reached maturity, might be due to time-dependent loss of hydraulic conductance. In fact, Kleaf loss was correlated with PsiL drop and stomatal closure. Leaf dehydration was aggravated by solute exportation from the basal towards the apical leaves, as revealed by the acropetal decrease of pi0. Kleaf was shown to be linearly and positively related to the prevailing ambient irradiance during plant growth, thus suggesting that leaf hydraulics is very sensitive to environmental conditions. It was concluded that the pronounced apical dominance of some sunflower cultivars is determined, among other factors, by plant hydraulic architecture.     

胡杨茎叶化学计量和异速生长关系特征随发育阶段变化的性别差异

胡杨(Populus euphratica Oliv.)是荒漠河岸林防风固沙和水土保持的雌雄异株树种。该研究以不同径阶(8、12、16、20 cm)的胡杨雌雄株为研究对象,通过当年生茎、叶化学计量元素(C、N、P、K)含量及生长关系分析,探讨不同器官化学计量随发育阶段的变化及异速生长关系的性别差异。结果表明：(1)胡杨雌雄株叶片C含量表现为大径阶(20 cm)显著高于小径阶(8 cm),而其叶片N含量随着径阶的增加显著增加;雌雄株茎、叶化学计量随着径阶的增加总体上呈增加趋势,且C、N含量均与径阶呈显著正相关关系;随着径阶的增加,雌株叶片P含量呈下降趋势,与径阶呈显著负相关关系,而雌雄株茎的P、K含量呈上升趋势,且与径阶呈显著正相关关系。(2)雌株各径阶叶片的N含量及8、12、20 cm径阶的叶片P含量均显著大于相应雄株,8、16、20 cm径阶当年生茎C含量以及20 cm径阶茎N、P含量均显著高于相应雄株。(3)雌株叶片C与N在20 cm径阶的斜率指数最大,而雄株在12 cm径阶斜率指数最大,雌雄株在各发育阶段N与P的变化较稳定;在相同C含量时,雄株茎能获得更多的N含量,雌株茎在相同N的情况下能获得更多P元素。研究发现,胡杨雌雄株间茎、叶化学计量元素含量和异速生长关系特征在不同发育阶段存在着明显性别差异,成熟雌株叶片需要更多的化学计量特征含量来满足生殖需求,总体反映了自身生长及环境适应的养分分配策略。     

Light conditions alter accumulation of long chain polyprenols in leaves of trees and shrubs throughout the vegetation season

In many plants belonging to angiosperms and gymnosperms the accumulation in leaves of long chain polyprenols and polyprenyl esters during growth in natural habitats depends on the light intensity. The amount of polyprenols in leaves is also positively correlated with the thickness of the leaf blade (SLA, specific leaf area). The polyprenol content of leaves shows seasonal changes with a maximum in autumn and a minimum in early summer with the difference between poorly and well illuminated plants persisting throughout the vegetation season.     

HETEROPHYLLIC DEVELOPMENT IN MUEHLENBECKIA (POLYGONACEAE)

Flowering shoots of Muehlenbeckia platyclados Meisn. bear only reduced scale leaves which resemble the membranous sheath portion (ochrea) of leaves of other members of the Polygonaceae. Shoots propagated from cuttings bear enlarged foliage leaves with distinct lamina, petiole, and ochrea zones. The developmental basis for this heterophylly is explored in order to determine whether scale leaves resemble foliage leaves in their pattern of ontogeny or are developmentally unique. SEM and histological analyses have shown that scale leaves and foliage leaves are distinctive from inception. The scale leaf arises as a collarlike growth and extends over the shoot apex as a hooded sheath without evidence of blade initiation. By contrast, the first stage of foliage-leaf ontogeny is the differentiation of the distal lamina from the future leaf base. As the foliage-leaf ochrea encircles the stem axis, the lamina grows erect and projects from the abaxial surface of the sheath. Lamina reduction coupled with ochrea elaboration in intermediate leaf types indicate a homology between the entire scale leaf and foliage-leaf ochrea. Despite this homology, the production of the bladeless scale leaf does not involve a mere suppression of the foliage-leaf lamina. Erect growth of the saccate ochrea of the foliage leaf contrasts with the hooded expansion of the scale. Early histological differences, including contrasting rates of cell differentiation, also distinguish the two organs. This disparity in modes of growth and differentiation from inception results from separate, predetermined courses of ontogeny. Unlike other plants studied, leaf size and degree of leaf elaboration decrease with shoot meristem enlargement in Muehlenbeckia. Leaf packing does increase with shoot development and may contribute to variations in leaf morphology. It is concluded that the peculiarities of the heterophyllic leaf sequence in Muehlenbeckia are a property of the shoot system as a whole.     

Some Soybean Cultivars Have Ability to Induce Germination of Sunflower Broomrape

Sunflower broomrape is a noxious parasitic weed which has caused severe damage to crop ecosystems. Trap crops can release a mixture of allelochemicals to induce the germination of sunflower broomrape. We studied the allelopathic effects of soybean on sunflower broomrape. Fourteen common soybean cultivars were grown in pots. Samples were collected from soybean plants and rhizosphere soil at five growth stages (V1, V3, V5, R2, and R4). The allelopathic effects of soybean reached highest at the V3 stage. Methanolic extracts of soybean roots induced higher broomrape germination than methanolic extracts of stems or leaves. The germination rates induced by root extracts (10-fold dilution) were positively correlated with germination rates induced by stem (10-fold dilution) and leaf extracts (10-fold dilution). The broomrape germination rates induced by root extracts were also positively correlated with soybean nodule diameter and dry weight. The results indicated that soybeans could induce sunflower broomrape germination. We conclude that soybean has the potential to be used as a trap crop for sunflower broomrape.     

Effects of Ethylene on Photosynthesis and Partitioning in Tomato, Lycopersicon esculentum Mill

The extended period of ethylene release from ethephon (2-chloroethylphosphonicacid) after application to intact tomato plants has provideda model system in which the effects of ethylene on photosyntheticmetabolism and carbon partitioning has been studied. Ethylenerelease from leaf tissue after ethephon treatment was 10 timesgreater than that from untreated control leaves. The specificactivity of 14C2H4 released from [14C] ethephon remained constantover several days demonstrating that the ethylene was derivedfrom the applied ethephon. The ethephon-treated plants exhibitedextreme epinasty of the leaves and 24 h after application theflower buds in the first visible cluster had abscised, leafexpansion at the apex had ceased and developing adventitiousroots were visible on the lower stem. Rates of steady-state photosynthesis, respiration, photorespirationand transpiration were the same in treated and control leaves24 h after ethephon application. Both treated and control leavespartitioned similar proportions of newly-fixed 14C from 14CO²into neutral (46.4%), acidic (14.0%), basic (5.0%) and insoluble(34.0%) leaf fractions under steady-state conditions. The speedof 11C-assimilate movement in the stems of control plants (3.62±0.42cm min-1 towards the apex and 4.03±0.15 cm min-1 towardsthe roots) was more rapid than in the ethephon-treated plants(2.90±0.31 cm min-1 upwards and 2.59±0.22 cm min-1downwards). Furthermore, in the control plants 20.0±5.4%of the 14C exported to the plant from the source leaf was transportedtowards the developing flower cluster and young leaves. Twenty-fourhours after ethephon application only 6.5 ±1.7% of theexported 14C was translocated towards the shoot. Contrary tosome reports ethylene did not affect steady-state gas exchangeprocesses while carbon partitioning was significantly alteredindicating that ethylene effects on photosynthetic carbon metabolismare indirect and not due to direct effects on photosyntheticprocesses per se. Key words: Ethylene, photosynthesis, partitioning     

Role of Ethylene in Induction of Flooding Damage in Sunflower

The possibility that symptoms of flooding damage in plants are primarily caused by an accumulation of ethylene was investigated using pot-grown sunflower (Helianthus annuus) plants. When plants were flooded to the basal pairs of leaves, ethylene in roots and stems below the water line began to increase. This coincided with the start of hypocotyl hypertrophy and new root formation in hypocotyls, which continued for 14-16 days. There were highly significant correlations between ethylene concentration and number of roots and hypocotyl diameter. After approximately 4 days of flooding, ethylene concentrations in stems between nodes for the 1st and 3rd basal pairs of leaves started to increase, coinciding with initiation of chlorophyll breakdown and epinasty of the 2nd basal pairs of leaves. Thus, there were correlations between ethylene concentration and chlorophyll breakdown and epinasty. The lower the leaves, the more chlorophyll breakdown among 1st, 2nd, 3rd, and 4th basal pairs of leaves. The longer the flooding, the more severe the flooding damage; and even when returned to normal condition, plants flooded longer than 3 days were not able to recover from flooding damage. A gas chromatographic study revealed that Ethephon was absorbed by roots and decomposed to ethylene in the plant. Damage symptoms caused by soil application of Ethephon, such as reduced stem height, chlorophyll breakdown, epinasty of the 2nd basal pairs of leaves, and hypocotyl hypertrophy, were almost identical with those caused by soil flooding treatment. Microscopic studies revealed that radially enlarged cells and increased intercellular spaces in the cortex were the major contribution to the increased hypocotyl diameter in both flooded and Ethephon-treated plants. It is concluded that the increase in ethylene concentration in flooded plants is largely, although not exclusively, responsible for flooding damage symptoms.     

Ontogenic change in leaf shape and crown form of a tropical tree,Scaphium macropodum (Sterculiaceae) in Borneo

Crown architecture was analyzed forScaphium macropodum (Sterculiaceae), a common shade-tolerant emergent tree of a tropical rain forest in West Kalimantan, Indonesia. Saplings and poles shorter than 12 m in height had no branches, and gathered their leaves at the ends of the stem. The leaves changed from entire to palmately-parted with increasing tree size. The parted leaves increased the light penetration through the clustered foliage. The size of leaves including the blade and petiole ranged from 22 cm to 147 cm. Because the weight of petiole per blade increased with leaf size, the leaf could not be enlarged infinitely. Taller trees with lateral branches bore small (about 40 cm in length) entire leaves. The light intensity in the forest increased from the ground to about 12 m tall and was nearly constant from 12 m to 18 m. Crown architecture ofS. macropodum adapted to this light environment. The monoaxial trees lower than 12 m could thus increase the amount of light with vertical elongation, and the branched trees higher than 12 m could increase it by means of lateral extension of crown area.     

Tissue Distribution of Glutamate Synthase and Glutamine Synthetase in Rice Leaves : Occurrence of NADH-Dependent Glutamate Synthase Protein and Activity in the Unexpanded, Nongreen Leaf Blades

To further explore the function of NADH-dependent glutamate synthase (GOGAT), the tissue distribution of NADH-GOGAT protein and activity was investigated in rice (Oryza sativa L.) leaves. The distributions of ferredoxin (Fd)-dependent GOGAT, plastidic glutamine synthetase, and cytosolic glutamine synthetase proteins were also determined in the same tissues. High levels of NADH-GOGAT protein (33.1 μg protein/g fresh weight) and activity were detected in the 10th leaf blade before emergence. The unexpanded, nongreen portion of the 9th leaf blade contained more than 50% of the NADH-GOGAT protein and activity per gram fresh weight when compared with the 10th leaf. The expanding, green portion of the 9th leaf blade outside of the sheath contained a slightly lower abundance of NADH-GOGAT protein than the nongreen portion of the 9th blade on a fresh weight basis. The fully expanded leaf blades at positions lower than the 9th leaf had decreased NADH-GOGAT levels as a function of increasing age, and the oldest, 5th blade contained only 4% of the NADH-GOGAT protein compared with the youngest 10th leaf blade. Fd-GOGAT protein, on the other hand, was the major form of GOGAT in the green tissues, and the highest amount of Fd-GOGAT protein (111 μg protein/g fresh weight) was detected in the 7th leaf blade. In the nongreen 10th leaf blade, the content of Fd-GOGAT protein was approximately 7% of that found in the 7th leaf blade. In addition, the content of NADH-GOGAT protein in the 10th leaf blade was about 4 times higher than that of Fd-GOGAT protein. The content of plastidic glutamine synthetase polypeptide was also the highest in the 7th leaf blade (429 μg/g fresh weight) and lowest in nongreen blades and sheaths. On the other hand, the relative abundance of the cytosolic glutamine synthetase polypeptide was the highest in the oldest leaf blade, decreasing to 10 to 20% of that value in young, nongreen leaves. These results suggest that NADH-GOGAT is important for the synthesis of glutamate from the glutamine that is transported from senescing source tissues through the phloem in the nongreen sink tissues in rice leaves.     

广谱抗真菌转基因水稻植株化学成分与结构动态变化

【背景】种植广谱抗真菌水稻可能会带来一定的环境生物安全问题,对其植株的化学成分进行实质等同性分析是转基因水稻安全性评价的重要内容之一。【方法】以表达广谱抗真菌蛋白转基因水稻转品1和转品8及其相应非转基因水稻七丝软粘的秸秆为研究材料,采用化学法和扫描电镜技术分析外源基因的导入对水稻秸秆化学成分以及组织显微结构的影响。【结果】（1）在整个生长发育过程中,广谱抗真菌转基因水稻转品1和转品8与其非转基因水稻七丝软粘叶片、叶鞘和茎的纤维素、半纤维素、木质素以及粗蛋白含量的变化趋势基本一致,且品种间化学成分的含量不存在显著差异。（2）广谱抗真菌转基因水稻叶片表皮的硅质瘤状结构以及气孔的形状和致密程度与其非转基因水稻七丝软粘相似;茎壁、厚壁组织、薄壁组织以及大小维管束的形态和分布情况未发生明显变化。【结论与意义】表达广谱抗真菌蛋白转基因水稻秸秆的化学成分和组织显微结构与非转基因水稻基本一致。这为广谱抗真菌转基因水稻的环境安全性评估提供了依据。     

The effect of waterlogging on the growth and ethylene content of Eucalyptus robusta Sm. (Swamp Mahogany)

Summary When waterlogged over a period of 80 days plants of Eucalyptus robusta Sm. showed symptoms of leaf chlorosis, epinasty and premature abscission, reduction of stem elongation, stem hypertrophy and formation of adventitious shoots; chlorophyll content was reduced and soluble protein content of the upper leaves increased. Waterlogging doubled the rate of release of ethylene from roots and stems within 6 days, but had no effect on the ethylene concentration of leaves.