Slow development of leaf photosynthesis in an evergreen broad-leaved tree, Castanopsis sieboldii: relationships between leaf anatomical characteristics and photosynthetic rate

Changes in net photosynthetic rate on a leaf area basis and anatomical properties during leaf development were studied in an evergreen broad‐leaved tree, Castanopsis sieboldii and an annual herb, Phaseolus vulgaris. In C. sieboldii, surface area of mesophyll cells facing the intercellular air spaces on a leaf area basis (S_mes) was already considerable at the time of full leaf area expansion (FLE). However, surface area of chloroplasts facing the intercellular air spaces on a leaf area basis (S_c), and chlorophyll and Rubisco contents on a leaf area basis increased to attain their maximal values 15–40 d after FLE. In contrast, in P. vulgaris, chloroplast number on a leaf area basis, S_c and S_mes at 10 d before FLE were two to three times greater than the steady‐state levels attained at around FLE. In C. sieboldii, the internal CO₂ transfer conductance (g_i) slightly increased for 10 d after FLE but then decreased toward the later stages. Limitation of photosynthesis by g_i was only about 10% at FLE, but then increased to about 30% at around 40 d after FLE. The large limitation after FLE by g_i was probably due to the decrease in CO₂ concentration in the chloroplast caused by the increases in thickness of mesophyll cell walls and in Rubisco content per chloroplast surface area. These results clearly showed that: (1) in C. sieboldii, chloroplast development proceeded more slowly than mesophyll cell expansion and continued well after FLE, whereas in P. vulgaris these processes proceeded synchronously and were completed by FLE; (2) after FLE, photosynthesis in leaves of C. sieboldii was markedly limited by g_i. From these results, it is suggested that, in the evergreen broad‐leaved trees, mechanical protection of mesophyll cells has priority over the efficient CO₂ transfer and quick construction of the chloroplasts.     

The consequences of delayed greening during leaf development for light absorption and light use efficiency

Many species of rainforest plants have an unusual form of leaf development such that leaves delay greening until after full leaf expansion. Chlorophyll accumulation was measured during leaf development in five woody rainforest species, three with white young leaves, and two with ‘normal’ greening. In the three species with white leaves, the chlorophyll content of the expanding leaves was about 0.4mg dm^?2, whereas in the two species with green young leaves, chlorophyll content was about 2.1 mg dm^?2. Chlorophyll accumulation in greenhouse and field experiments was independent of light level. During leaf expansion, species with delayed chloroplast development only absorb 18–25% of the maximum possible light, compared with 80% for species with normal greening. Furthermore, species with delayed greening have low chlorophyll contents and reduced absorption for at least 30 d after full expansion. At a PPFD typical of the forest under story, the photosynthetic light use efficiency based upon incident radiation was 0.030–0.036 for species with delayed chloroplast development and 0.068–0.085 for the two species with normal greening. The lower light use efficiency of white species was primarily due to decreased light absorption. However, they also had a slightly lower light use efficiency based upon absorbed radiation, suggesting that development of other components of the photo-synthetic apparatus also may be delayed. Despite the fact that delayed greening decreases light absorption and light use efficiency during leaf development, it is extremely common in shade-tolerant species. We suggest that an advantage of delayed greening is that resources are not invested in the leaf until it is fully expanded and better defended from herbivores.     

The genetic control of plastid division in higher plants

The division of plastids is an important part of plastid differentiation and development and in distinct cell types, such as leaf mesophyll cells, results in large populations of chloroplasts. The morphology and population dynamics of plastid division have been well documented, but the molecular controls underlying plastid division are largely unknown. With the isolation of Arabidopsis mutants in which specific aspects of plastid and proplastid division have been disrupted, the potential exists for a detailed knowledge of how plastids divide and what factors control the rate of division in different cell types. It is likely that knowledge of plant homologues of bacterial cell division genes will be essential for understanding this process in full. The processes of plastid division and expansion appear to be mutually independent processes, which are compensatory when either division or expansion are disrupted genetically. The rate of cell expansion appears to be an important factor in initiating plastid division and several systems involving rapid cell expansion show high levels of plastid division activity. In addition, observation of plastids in different cell types in higher plants shows that cell-specific signals are also important in the overall process in determining not only the differentiation pathway of plastids but also the extent of plastid division. It appears likely that with the exploitation of molecular techniques and mutants, a detailed understanding of the molecular basis of plastid division may soon be a reality.     

What is the role of cell division in leaf development?

An idea underlying a great deal of research and discussion in plant cell and developmental biology is that the spatial regulation of cell division plays a key role in plant development. In this article, the role of cell division in two aspects of leaf development is analysed: morphogenesis (leaf initiation, growth, and the generation of leaf shape) and histogenesis (the differentiation of leaf cells to form the various cell types that make up a functional leaf). The point of view that emerges from this analysis is that the rate and pattern of cell division is important for leaf development, but does not dictate leaf size, shape, or cell fate.     

Enzymes of chlorophyll biosynthesis

The enzymes responsible for chlorophyll biosynthesis in plants, algae and cyanobacteria are identified and described, with emphasis on their protein composition and structure, required cofactors, physical and catalytic properties, protein-protein interactions and allosteric modulation of activity. Properties and features of the pathway that enable it to operate in a coordinated way while using unstable and light-sensitive intermediates in potentially hostile biochemical environments are discussed. The evolutionary relationships and possible origins of the chlorophyll biosynthetic enzymes are also discussed.     

Effects of nitrogen deprivation on cell division and expansion in leaves of Ricinus communis L.

The effects of nitrogen deprivation on leaf extension, cell numbers and epidermal cell size were followed in leaves of Ricinus communis L. The extent to which reductions in final cell number or final epidermal cell size contributed to the reduction in final leaf size depended on the developmental stage of the leaf at the time of N deprivation. In leaves which already had their full complement of cells (leaf 2), the reduction in final leaf size following nitrogen deprivation was associated with a reduction in final cell size. In leaves that were at earlier stages of development at the onset of N deprivation (leaves 3 and 4), the reduction in final leaf size was greater than in leaf 2. In these younger leaves, the final cell size was even smaller than in leaf 2, but the greatest contribution to reduced final leaf size was a reduction in the number of cells produced. This accounted for approximately 80% of the reduction in final leaf size in leaf 4. During leaf development, the contribution from different tissue layers to the total cell number changed. In the smallest leaf sizes, the contribution from upper and lower epidermis and spongy parenchyma was greater than that from palisade parenchyma. As the leaf size increased, cells in the palisade parenchyma continued to divide for longer than in the other layers. At final leaf size, the contribution from the different tissue layers to total cell number was the same for leaves 2, 3 and 4, irrespective of N treatment. In these final leaf structures, palisade parenchyma contributed 60% of the total cell number. Thus, although nitrogen deprivation affected leaf size variously through cell division and cell expansion, depending on leaf developmental stage at the time of nitrogen deprivation, the ratio of cell numbers and sizes in different tissue layers, at final leaf size, was unaffected.     

4种阔叶树种叶中氮和磷的季节动态及其转移

从叶完全展开到生长季结束 ,对常绿阔叶树种日本米槠 (Castanopsis cuspidata(Thunb.) Schottky)和具柄冬青 (Ilexpedunculosa Miq)及落叶阔叶树种栎 (Quercus serrata Murr.)和栓皮栎 (Quercus variabilis Blume)叶片的 N和 P浓度、含量和养分转移进行了测定 .在生长期中日本米槠新叶的 N浓度在 5月为 36 .6 g/ kg,然后降到 15 .5和 17.5 g/ kg之间 ,其老叶的N浓度波动于 10 .4和 13.1g/ kg的范围内 ,而具柄冬青新叶的 N浓度从 2 7.3下降到 16 .0 g/ kg,接着上升到 18.3g/ kg,其老叶的 N浓度在 12 .0到 15 .5 g/ kg的范围内。栎和栓皮栎的叶 N浓度分别从 2 8.8下降到 18.1g/ kg和从 2 8.5下降到17.5 g/ kg。日本米槠新叶的 N含量从 1.5 4下降到 1.35 g/ m2 ,接着上升到 1.5 0 g/ m2 ,其老叶 N含量从 1.36下降到1.0 0 g/ m2 ,接着上升到 1.2 1g/ m2 ,而具柄冬青新叶的 N含量从 2 .2 5下降到 1.6 0 g/ m2 ,接着上升到 2 .2 0 g/ m2 ,其老叶的 N含量从 2 .13下降到 1.6 5 g/ m2。栎和栓皮栎的叶 N含量分别从 2 .10下降到 1.2 8g/ m2和从 2 .95下降到 2 .13g/ m2。日本米槠新叶的 P浓度由 3.39g/ kg降到 1.12和 1.15 g/ kg之间 ,其老叶的 P浓度变化于 0 .6 6和 0 .88g/ kg的范围内 ,而具柄冬青新叶的     

藏东南色季拉山林线过渡带7种灌木植物的叶氮回收潜力

在湿润的青藏高原东南部, 为什么常绿灌木广泛占据高海拔的林线过渡带及以上的高山带, 而落叶灌木只能零星分布?未来气候变暖对该区不同功能群物种的影响是否相同?通过测定西藏东南部色季拉山林线过渡带7种灌木凋落叶的氮含量, 比较了极端高海拔地区灌木不同表达单位的叶氮回收潜力在不同功能群间的差异, 以及不同海拔、不同坡向间的差异, 试图从养分限制的角度为解答上述科学问题提供基础数据。研究结果表明: 1)从基于单位质量叶氮含量(N_mass)的叶氮回收潜力来看, 常绿灌木裂毛雪山杜鹃(薄毛海绵杜鹃) (Rhododendron aganniphum var. schizopeplum)显著高于其他6种落叶灌木, 但由于受比叶重的影响, 基于单位面积叶氮含量(N_area)的叶氮回收潜力则表现为落叶灌木总体较高; 2)落叶灌木山生柳(Salix oritrepha)和拉萨小檗(Berberis hemsleyana)的叶氮回收潜力在不同海拔或不同坡向间均无显著差异, 但裂毛雪山杜鹃基于N_mass的叶氮回收潜力在高海拔地段明显偏高。在极端高海拔的林线过渡带, 通过降低凋落叶中的氮含量(增加叶氮回收潜力)以达到高效的养分利用可能是常绿灌木裂毛雪山杜鹃适应高寒胁迫环境的重要策略。与落叶灌木相比, 常绿灌木裂毛雪山杜鹃叶氮回收潜力对未来气候变暖可能更敏感。     

The Effect of Glucose on the Biochemical and Ultrastructural Characteristics of Developing Euglena Chloroplasts*

SYNOPSIS. Chloroplast development is inhibited in Euglena gracilis strain Z, when greened in a medium containing glucose. This inhibition is reflected not only in the pattern of chlorophyll accumulation but also in the chloroplast ultrastructure and activities of the 2 light reactions of photosynthesis. Chloroplasts of cells greening in the presence of glucose are delayed in déveloping certain structures. Photosystem I activity develops at about the same rate as that of the controls during the first 48 h of greening, after which it develops at a slower rate. The rate of development of photosystem II activity in cells greening in a glucose medium lags considerably behind that of the controls until the later hours of greening. There are similarities between glucose inhibition and chloramphenicol inhibition of chloroplast development. Glucose may inhibit a step in chloroplast development ultimately controlled by the chloroplast genome.     

Isolation and initial characterization of virescent mutants of Arabidopsis thaliana

In higher plants, development of the chloroplasts must be coordinated with development of the leaf. In order to study the signals that synchronize these two developmental processes, we have isolated virescent (delayed in greening) mutants of Arabidopsis thaliana. Two such mutants that have pale-green young leaves which gradually green more fully during leaf maturation have been partially characterized. The two, vir1 and vir2, are due to separate nuclear recessive mutations. The pale leaves of vir1 and vir2 both had reduced 77°K fluorescence emission at 730–734 nm relative to that at 686–687 nm, indicating a reduction in the relative amount of LHC I compared to WT. As leaves greened, the amount of LHC I increased to near wildtype levels. The shift in the fluorescence emission peak from 730 nm to 734 nm, characteristic of maturing LHC I, was seen for vir1, but not vir2, suggesting that vir1 is a regulatory mutant while vir2 may be defective in a specific aspect(s) of LHC I function.Abbreviations D dark - EMS ethyl methanesulfonate - er erecta - gl1 glabrous1 - L light - LHC I light harvesting complex of Photosystem I - LHC II light harvesting complex of Photosystem II - M2 second generation of mutagenized seed - M3 third generation of mutagenized seed - vir virescent - WT wildtype