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.     

Variation in mesophyll anatomy and photosynthetic capacity during leaf development in a deciduous mesophyte fruit tree (<Emphasis Type="Italic">Prunus persica</Emphasis>) and an evergreen sclerophyllous Mediterranean shrub (<Emphasis Type="Italic">Olea europaea</Emphasis>)

The relative importance that biomechanical and biochemical leaf traits have on photosynthetic capacity would depend on a complex interaction of internal architecture and physiological differences. Changes in photosynthetic capacity on a leaf area basis and anatomical properties during leaf development were studied in a deciduous tree, Prunus persica, and an evergreen shrub, Olea europaea. Photosynthetic capacity increased as leaves approached full expansion. Internal CO₂ transfer conductance (g _i) correlated with photosynthetic capacity, although, differences between species were only partially explained through structural and anatomical traits of leaves. Expanding leaves preserved a close functional balance in the allocation of resources of photosynthetic component processes. Stomata developed more rapidly in olive than in peach. Mesophyll thickness doubled from initial through final stages of development when it was twice as thick in olive as in peach. The surface area of mesophyll cells exposed to intercellular air spaces per unit leaf area tended to decrease with increasing leaf expansion, whereas, the fraction of mesophyll volume occupied by the intercellular air spaces increased strongly. In the sclerophyllous olive, structural protection of mesophyll cells had priority over efficiency of photochemical mechanisms with respect to the broad-leaved peach. The photosynthetic capacity of these woody plants during leaf development relied greatly on mesophyll properties, more than on leaf mass per area ratio (LMA) or nitrogen (N) allocation. Age-dependent changes in diffusion conductance and photosynthetic capacity affected photosynthetic relationships of peach versus olive foliage, evergreen leaves maturing functionally and structurally a bit earlier than deciduous leaves in the course of adaptation for xeromorphy.     

Characterization of temperature‐sensitive mutants reveals a role for receptor‐like kinase SCRAMBLED/STRUBBELIG in coordinating cell proliferation and differentiation during Arabidopsis leaf development

The balance between cell proliferation and cell differentiation is essential for leaf patterning. However, identification of the factors coordinating leaf patterning and cell growth behavior is challenging. Here, we characterized a temperature‐sensitive Arabidopsis mutant with leaf blade and venation defects. We mapped the mutation to the sub‐2 allele of the SCRAMBLED/STRUBBELIG (SCM/SUB) receptor‐like kinase gene whose functions in leaf development have not been demonstrated. The sub‐2 mutant displayed impaired blade development, asymmetric leaf shape and altered venation patterning under high ambient temperature (30°C), but these defects were less pronounced at normal growth temperature (22°C). Loss of SCM/SUB function results in reduced cell proliferation and abnormal cell expansion, as well as altered auxin patterning. SCM/SUB is initially expressed throughout leaf primordia and becomes restricted to the vascular cells, coinciding with its roles in early leaf patterning and venation formation. Furthermore, constitutive expression of the SCM/SUB gene also restricts organ growth by inhibiting the transition from cell proliferation to expansion. We propose the existence of a SCM/SUB‐mediated developmental stage‐specific signal for leaf patterning, and highlight the importance of the balance between cell proliferation and differentiation for leaf morphogenesis.