**3. Conclusion**

These characteristics that show high photosynthetic capacity and high concentration of nitrogen for younger leaf and short leaf longevity are corresponded with fast-growing species [25]. In general, fast-growing species shows that photosynthetic rate is decreased drastically by increase of leaf age [21, 30]. This trend is clear for evergreen oak compared with conifer species [30]. Moreover, there are a fast-growing species among same genus of *Picea*, and *Picea abies* and *Picea glauca* are considered as fast-growing species [21]. These species showed high photosynthetic rate for younger leaves; however, their high values were not maintained. Also, fast-growing species have a high rate of leaf turnover [31]. Woody species have a leaf turnover mechanism, and when old leaves are lost, leaf nitrogen is retranslocated to younger leaves [32]. *S. nipponica* showed continuous decrease of *N*mass (**Figure 6**), and its trait is probably related with retranslocation of nitrogen. *S. nipponica* may be retranslocated nitrogen from old

For the other *Sasa* species, the maximum value of *P*sat for current leaves of *S. kurilensis* was lower than other species; however, its value for 2-year-old leaves was maintained for 5 months (**Figure 3**). These traits are corresponded with slow-growing species [25]. The concrete slow-growing species are *Taxus baccata*, *Picea mariana*, and *Picea rubens* [21, 30]. The leaf longevities of these species were over 5 years, and photosynthetic rates showed high value for 6-year-old leaves [21, 30]. Also, maximum leaf longevity of *S. kurilensis* is 5 years [4], and its ecophysiological characteristics are similar with other slow-growing species. Also, slowgrowing species have a characteristic to maintain high value of PNUE for aged leaves [21, 30], and *S. kurilensis* showed high PNUE for 2-year-old leaves (**Figure 7**). This trait is related with

On other traits, slow-growing species has thick leaves [21, 25]. Leaves of *S. kurilensis* showed a low value of SLA (**Figure 4**), which was characterised by thick leaves. In general, species with a small SLA allocates nitrogen to the leaf cell wall and increases toughness of the cell [33]. This trait contributes to the extent of leaf longevity [34]. Thus, allocation of nitrogen in leaves for *S. kurilensis* is probably larger for cell wall than for protein of photosynthetic apparatus. As a result, *S. kurilensis* may make leaves with a long longevity but with a low photosynthetic rate.

*P*sat of *S. senanensis* for current leaves showed high values in August and September (**Figure 3**). In contrast, current leaves of *S. senanensis* were thick (**Figure 4**), and *N*area and *N*mass were low compared with *S. nipponica* (**Figure 5**). Thus, ecophysiological characteristics of *S. senanensis* are not similar with *S. nipponica*. In contrast, leaves of *S. senanensis* were thin (**Figure 4**) and short longevity (about 2 years, [4]) compared with *S. kurilensis*. Thus, ecophysiological characteristics of *S. senanensis* are also not similar with *S. kurilensis*. Consequently, ecophysiological characteristics of *S. senanensis* are intermediate between fast- and slow-growing species. On the remarkable characteristics of *S. senanensis*, PNUE showed the highest value for current leaves (**Figure 7**). *S. senanensis* may allocate more nitrogen to protein of photosynthesis apparatus compared with other *Sasa* species. Similar ecophysiological characteristics were

In addition, the trait of chlorophyll concentration also concerns with ecophysiological characteristics. The concentration of chlorophyll showed high values for *S. kurilensis*, especially 2-year-old leaves (**Figure 6**). In general, chlorophylls have light harvesting complex proteins

to young leaves, thus maintaining high photosynthetic capacity.

194 Bamboo - Current and Future Prospects

the maintenance of photosynthetic rate for long period.

reported for *Pinus pinea* and *Picea jezoensis* var. *hondoensis* [21, 30].

*Sasa* species regenerates at the same place with clonal development, and these traits cannot be simply classified into fast- and slow-growing species as other species. We regard the *Sasa* species as follows: *S. nipponica* is classified as a fast-growing species, whereas *S. kulinensis* are slow-growing species. Indeed, ecophysiological characteristics of *Sasa* sp. are the same as slowand fast-growing species as found in other plant species. *S. senanensis* cannot be classified as two growing types and showed intermediate characteristics between fast- and slow-growing species.

Related to the habitat of the three *Sasa* species, edaphic habitat of *S. nipponica* is considered to be the deep humus layer and A-horizon [36]. The characteristics of a fast-growing species is to have an advantage in a fertile habitat, and the growth trait of *S. nipponica* shows a rapid turnover of leaves and culms [4], which is considered to be suitable for the habitat. We conclude that ecophysiological characteristics of *S. nipponica* are adapted to fertile habitats. The distribution area of *S. nipponica* is classified as low altitudes, facing to the coast of Pacific Ocean where the summers are relatively cloudy with high humidity and the high photosynthetic performance of *S. nipponica* is kept [8]. Moreover, although the snowy period there is short, the soil freezes with cold climate [5]. *Sasa* cannot keep evergreen leaves during winter; hence, the *Sasa* species must produce new leaves from spring after the death of leaves of previous year. Its high photosynthetic rate may be compensating short leaf longevity.

In contrast, the distribution of *S. kurilensis* is hillsides and slope of valley sides where soil depth is shallow [36]. In general, these locations restrict plant growth. The leaves and culms of *S. kurilensis* can survive for several years [4], and these traits may exist to compensate for low photosynthetic productivity. *S. kurilensis* showed high concentration of chlorophyll and PNUE for 2-year-old leaves (**Figures 6** and **7**). This characteristic is suitable for conditions where resources are limited*.* Thus, we conclude that ecophysiological characteristics of *S. kurilensis* reflect the adaptability to infertile habitats. *S. kurilensis* distributes at high mountain areas in Hokkaido Island (**Figure 1**). The area of *S. kurilensis* probably corresponds with deep snow and harsh environmental conditions.

The habitat of *S. senanensis* is similar to the soil condition of *S. nipponica* [36]. Leaf longevity of *S. senanensis* is about 2 years [4], and this characteristic is probably suitable for relatively good environmental conditions, such as high soil fertility. Compared with *S. nipponica*, *N*mass and *N*area in current leaves were lower for *S. senanensis* (**Figure 5**). Thus, the nutrient requirement of *S. senanensis* is also lower than that of *S. nipponica*, and *S. senanensis* can adapt to infertile habitats or resources limited conditions. Moreover, the longevity of culm of *S. senanensis* is about 5 years, which is different from its leaves. Its culm has buds at every node (**Figure 2**), and the leaves can flush during the latter period of the culm life-span. Based on these results, we conclude that the growth characteristics of *S. senanensis* may be high flexibility, and it is also able to adapt to different nutrient and environmental conditions. In fact, the distribution area for *S. senanensis* in Hokkaido Island is the largest (**Figure 1**). The flexibility of *S. senanensis* may be enabling this species to grow in a broad distribution range.

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