We examined 15 traits in leaves and stems related to leaf C economy and water use for 32 co-existing angiosperms at ridge sites with shallow soil in the Bonin Islands. Across species, stem density was positively correlated to leaf mass per area (LMA), leaf lifespan (LLS), and total phenolics and condensed tannins per unit leaf N (N-based), and negatively correlated to leaf osmotic potential and saturated water content in leaves. LMA and LLS were negatively correlated to photosynthetic parameters, such as area-, mass-, and N-based assimilation rates. Although stem density and leaf osmotic potential were not associated with photosynthetic parameters, they were associated with some parameters of the leaf C economy, such as LMA and LLS. In the principal component (PCA) analysis, the first three axes accounted for 74.4% of total variation. Axis 1, which explained 41.8% of the total variation, was well associated with parameters for leaf C and N economy. Similarly, axis 2, which explained 22.3% of the total variation, was associated with parameters for water use. Axis 3, which explained 10.3% of the total variation, was associated with chemical defense within leaves. Axes 1 and 2 separated functional types relatively well, i.e., creeping trees, ruderal trees, other woody plants, C(3) shrubs and forbs, palms, and CAM plants, indicating that plant functional types were characterized by similar attributes of traits related to leaf C and N economy and water use. In addition, when the plot was extended by two unrelated traits, leaf mass-based assimilation rates and stem density, it also separated these functional types. These data indicate that differences in the functional types with contrasting plant strategies can be attributed to functional integration among leaf C economy, hydraulics, and leaf longevity, and that both leaf mass-based assimilation rates and stem density are key factors reflecting the different functions of plant species.

These results indicate that species with a shorter leaf life span have higher photosynthetic productivity and are more susceptible to herbivore attack than species with a longer leaf life span. This supports the "cost-benefit hypothesis".

Background and Aims Betula ermanii, B. maximowicziana and B. platyphylla var. japonica have heterophyllous leaves (i.e. early leaves and late leaves) and are typical pioneer species in northern Japan. Chemical and physical defences against herbivores in early and late leaves of these species were studied. d Methods Two-year-old seedlings were grown under full sunlight in a single growing season. Three-week-old leaves of each seedling were harvested three times (May, July and October). Total phenolics and condensed tannin content were determined for chemical defence and leaf toughness and trichome density were assessed for physical defence. Defoliation of early leaves in May was also performed to study the contribution of early leaves to subsequent growth. d Key Results Chemical and physical defences were greater in early than late leaves in B. platyphylla and B. ermanii, whereas the reverse was true in B. maximowicziana. In contrast to its weak chemical defences, the trichome density in B. maximowicziana was very high. In B. platyphylla and B. ermanii, the relative growth rates (RGR) were greater early in the growing season. Negative effects on growth of removal of early leaves were signi®cant only in B. platyphylla. d Conclusions B. platyphylla and B. ermanii invest in defence in early rather than late leaves, since early leaves are crucial to subsequent growth. In contrast, B. maximowicziana more strongly defends its late leaves, since its RGR is maintained at the same level throughout the growing season.

No abstract

Mixed broad-leaved forests are abundantly populated by several kinds of herbivore species in northern Japan. The life of herbivores depends strongly on the quality of food leaves. Leaf quality is changing with increasing atmospheric CO 2 and nitrogen deposition. Four seral species of deciduous broad-leaved tree seedlings (alder, birch, oak and maple) were raised in all four combinations of two levels of CO 2 and two levels of nutrient, to examine foliar defense traits. To evaluate the relative defense capacity of the four tree species, we used wild silkworms (Erisan; a generalist herbivore) as a bioassay material. Except with alder, the survival rate and longevity (ML 50 ) of the silkworms were least when they were fed with leaves of seedlings raised under high CO 2 and infertile soil conditions, and longest on a diet of leaves grown in ambient CO 2 and fertile soil, especially in birch and maple. The longevity of Erisan decreased in the order birch, oak and maple. The longevity of Erisan fed with alder leaves was independent of CO 2 levels and was longer on alder seedlings grown in infertile soil conditions. Alder is an actinorrhizal plant that can fix atmospheric nitrogen in root nodules formed by the actinomycetes Frankia sp. The activity of symbiotic microbes would have been enhanced by the greater amount of photosynthates received from the host plants at high CO 2 , improving the food quality for the silkworms. All but alder, leaf chemical traits especially C/N ratio were concluded to affect ML 50 of Erisan larvae.