Kenichi Yazaki

Coordination between leaf and stem traits related to leaf carbon gain and hydraulics across 32 drought-tolerant angiosperms

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, C3 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.

Seasonal variations of gas exchange and water relations in deciduous and evergreen trees in monsoonal dry forests of Thailand

This study compared leaf gas exchange, leaf hydraulic conductance, twig hydraulic conductivity and leaf osmotic potential at full turgor between two drought-deciduous trees, Vitex peduncularis Wall. and Xylia xylocarpa (Roxb.) W. Theob., and two evergreen trees, Hopea ferrea Lanessan and Syzygium cumini (L.) Skeels, at the uppermost canopies in tropical dry forests in Thailand. The aims were to examine (i) whether leaf and twig hydraulic properties differ in relation to leaf phenology and (ii) whether xylem cavitation is a determinant of leaf shedding during the dry season. The variations in almost all hydraulic traits were more dependent on species than on leaf phenology. Evergreen Hopea exhibited the lowest leaf-area-specific twig hydraulic conductivity (leaf-area-specific K(twig)), lamina hydraulic conductance (K(lamina)) and leaf osmotic potential at full turgor (Ψ(o)) among species, whereas evergreen Syzygium exhibited the highest leaf-area-specific K(twig), K(lamina) and Ψ(o). Deciduous Xylia had the highest sapwood-area-specific K(twig), along with the lowest Huber value (sapwood area/leaf area). More negative osmotic Ψ(o) and leaf osmotic adjustment during the dry season were found in deciduous Vitex and evergreen Hopea, accompanied by low sapwood-area-specific K(twig). Regarding seasonal changes in hydraulics, no remarkable decrease in K(lamina) and K(twig) was found during the dry season in any species. Results suggest that leaf shedding during the dry season is not always associated with extensive xylem cavitation.

Response to dehydration and irrigation in invasive and native saplings: osmotic adjustment versus leaf shedding

To clarify the mechanism underlying successful invasion by tree species into xeric sites on Japans Bonin Islands, we compared the water use of an alien species, Psidium cattleianum, which is rapidly expanding on ridge sites with shallow soil, with that of a native species, Trema orientalis. We hypothesized that there is a trade-off between leaf shedding with low cavitation resistance (frequent xylem cavitation plus refilling ability) and leaf osmotic adjustment with high cavitation resistance (cessation of xylem cavitation plus canopy leaf retention), indicating contrasting strategies for drought tolerance and water use in semi-arid regions. We examined leaf turnover, leaf gas exchange, leaf water potential and water distribution in stem xylem conduits using cryo-scanning electron microscopy for the saplings of both species under three cycles of artificial drought and sudden pulse irrigation. Invasive P. cattleianum saplings were highly resistant to cavitation in stem xylem conduits, retained their leaves and exhibited effective leaf osmotic adjustment under the drought treatment. In contrast, native T. orientalis saplings exhibited xylem cavitation, conspicuous leaf shedding and less effective leaf osmotic adjustment under the drought treatment. Leaf gas exchange rate recovered more rapidly in P. cattleianum saplings than in T. orientalis saplings immediately following pulse irrigation after a period without irrigation, especially in the first drought cycle. Embolized conduits in T. orientalis were refilled by pulse irrigation, and leaf gas exchange rate recovered following refilling. The two tree species showed contrasting strategies for drought tolerance and water use along a trade-off axis. Cavitation avoidance and effective leaf osmotic adjustment in P. cattleianum saplings under drought conditions partially support their survival at the xeric ridge sites on the Bonin Islands. Our results help to explain the success of P. cattleianum in its invasion of a sub-arid environment.

Cutting stems before relaxing xylem tension induces artefacts in Vitis coignetiae, as evidenced by magnetic resonance imaging

It was recently reported that cutting artefacts occur in some species when branches under tension are cut, even under water. We used non-destructive magnetic resonance imaging (MRI) to investigate the change in xylem water distribution at the cellular level in Vitis coignetiae standing stems before and after relaxing tension. Less than 3% of vessels were cavitated when stems under tension were cut under water at a position shorter than the maximum vessel length (MVL) from the MRI point, in three of four plants. The vessel contents remained at their original status, and cutting artefact vessel cavitation declined to <1% when stems were cut at a position farther than the MVL from the MRI point. Water infiltration into the originally cavitated vessels after cutting the stem, i.e. vessel refilling, was found in <1% of vessels independent of cutting position on three of nine plants. The results indicate that both vessel cavitation and refilling occur in xylem tissue under tension following stem cutting, but its frequency is quite small, and artefacts can be minimized altogether if the distance between the monitoring position and the cutting point is longer than the MVL.

Growth, Net Photosynthetic Rate, Nutrient Status and Secondary Xylem Anatomical Characteristics of Fagus Crenata Seedlings Grown in Brown Forest Soil Acidified with H2SO4 Solution

Dry matter production, net photosynthetic rate, leaf nutrient status and trunk anatomical characteristics of Fagus crenata seedlings grown in brown forest soil acidified by adding H2SO4 solution were investigated. The soil acidification leaded to decreased (Ca+Mg+K)/Al molar ratio in the soil solution. Dry mass per plant of the seedlings grown in the soil treated with H+ at 120 mg·L−1 was significantly reduced compared with the control value at 0 mg·L−1. When net photosynthetic rate was reduced in the seedlings grown in the soil treated with H+ at 120 mg·L−1, the carboxylation efficiency and maximum net photosynthetic rate at saturated CO2-concentration were lower than the control values. The addition of H+ to the soil at 120 mg·L−1iinduced a reduction in the concentration of Ca in the leaf. By contrast, the concentration of Al in the leaf was increased with increasing the amount of H+ added to the soil. The annual ring formed in the seedlings grown in the soil treated with H+ at 120 mg·L−1 was significantly narrower than that at 0 (control), 10, 30, 60 or 90 mg·L−1. Based on the results obtained in the present study, we conclude that Fagus crenata is relatively sensitive to a reduction in the (Ca+Mg+K)/Al molar ratio of soil solution compared with Picea abies.

Recovery of Physiological Traits in Saplings of Invasive Bischofia Tree Compared with Three Species Native to the Bonin Islands under Successive Drought and Irrigation Cycles.

Partial leaf shedding induced by hydraulic failure under prolonged drought can prevent excess water consumption, resulting in delayed recovery of carbon productivity following rainfall. To understand the manner of water use of invasive species in oceanic island forests under a fluctuating water regime, leaf shedding, multiple physiological traits, and the progress of embolism in the stem xylem under repeated drought-irrigation cycles were examined in the potted saplings of an invasive species, Bischofia javanica Blume, and three endemic native species, Schima mertensiana (Sieb. Et Zucc,) Koitz., Hibiscus glaber Matsum, and Distylium lepidotum Nakai, from the Bonin Islands, Japan. The progress of xylem embolism was observed by cryo-scanning electron microscopy. The samples exhibited different processes of water saving and drought tolerance based on the different combinations of partial leaf shedding involved in embolized conduits following repeated de-rehydration. Predawn leaf water potential largely decreased with each successive drought-irrigation cycle for all tree species, except for B. javanica. B. javanica shed leaves conspicuously under drought and showed responsive stomatal conductance to VPD, which contributed to recover leaf gas exchange in the remaining leaves, following a restored water supply. In contrast, native tree species did not completely recover photosynthetic rates during the repeated drought-irrigation cycles. H. glaber and D. lepidotum preserved water in vessels and adjusted leaf osmotic rates but did not actively shed leaves. S. mertensiana exhibited partial leaf shedding during the first cycle with an osmotic adjustment, but they showed less responsive stomatal conductance to VPD. Our data indicate that invasive B. javanica saplings can effectively use water supplied suddenly under drought conditions. We predict that fluctuating precipitation in the future may change tree distributions even in mesic or moist sites in the Bonin Islands.

Seasonal ozone uptake by a warm-temperate mixed deciduous and evergreen broadleaf forest in western Japan estimated by the Penman-Monteith approach combined with a photosynthesis-dependent stomatal model.

Canopy-level stomatal conductance over a warm-temperate mixed deciduous and evergreen broadleaf forest in Japan was estimated by the Penman-Monteith approach, as compensated by a semi-empirical photosynthesis-dependent stomatal model, where photosynthesis, relative humidity, and CO2 concentration were assumed to regulate stomatal conductance. This approach, using eddy covariance data and routine meteorological observations at a flux tower site, permits the continuous estimation of canopy-level O3 uptake, even when the Penman-Monteith approach is unavailable (i.e. in case of direct evaporation from soil or wet leaves). Distortion was observed between the AOT40 exposure index and O3 uptake through stomata, as AOT40 peaked in April, but with O3 uptake occurring in July. Thus, leaf pre-maturation in the predominant deciduous broadleaf tree species (Quercus serrata) might suppress O3 uptake in springtime, even when the highest O3 concentrations were observed.

The dynamics of carbon stored in xylem sapwood to drought-induced hydraulic stress in mature trees

Climate-induced forest die-off is widespread in multiple biomes, strongly affecting the species composition, function and primary production in forest ecosystems. Hydraulic failure and carbon starvation in xylem sapwood are major hypotheses to explain drought-induced tree mortality. Because it is difficult to obtain enough field observations on drought-induced mortality in adult trees, the current understanding of the physiological mechanisms for tree die-offs is still controversial. However, the simultaneous examination of water and carbon uses throughout dehydration and rehydration processes in adult trees will contribute to clarify the roles of hydraulic failure and carbon starvation in tree wilting. Here we show the processes of the percent loss of hydraulic conductivity (PLC) and the content of nonstructural carbohydrates (NSCs) of distal branches in woody plants with contrasting water use strategy. Starch was converted to soluble sugar during PLC progression under drought, and the hydraulic conductivity recovered following water supply. The conversion of NSCs is strongly associated with PLC variations during dehydration and rehydration processes, indicating that stored carbon contributes to tree survival under drought; further carbon starvation can advance hydraulic failure. We predict that even slow-progressing drought degrades forest ecosystems via carbon starvation, causing more frequent catastrophic forest die-offs than the present projection.

Mesophyll conductance in leaves of Japanese white birch (Betula platyphylla var. japonica) seedlings grown under elevated CO2 concentration and low N availability

To test the hypothesis that mesophyll conductance (gm ) would be reduced by leaf starch accumulation in plants grown under elevated CO2 concentration [CO2 ], we investigated gm in seedlings of Japanese white birch grown under ambient and elevated [CO2 ] with an adequate and limited nitrogen supply using simultaneous gas exchange and chlorophyll fluorescence measurements. Both elevated [CO2 ] and limited nitrogen supply decreased area-based leaf N accompanied with a decrease in the maximum rate of Rubisco carboxylation (Vc,max ) on a CO2 concentration at chloroplast stroma (Cc ) basis. Conversely, only seedlings grown at elevated [CO2 ] under limited nitrogen supply had significantly higher leaf starch content with significantly lower gm among the treatment combinations. Based on a leaf anatomical analysis using microscopic photographs, however, there were no significant difference in the area of chloroplast surfaces facing intercellular space per unit leaf area among treatment combinations. Thicker cell walls were suggested in plants grown under limited N by increases in leaf mass per area subtracting non-structural carbohydrates. These results suggest that starch accumulation and/or thicker cell walls in the leaves grown at elevated [CO2 ] under limited N supply might hinder CO2 diffusion in chloroplasts and cell walls, which would be an additional cause of photosynthetic downregulation as well as a reduction in Rubisco activity related to the reduced leaf N under elevated [CO2 ].

Responses of gas-exchange rates and water relations to annual fluctuations of weather in three species of urban street trees

The frequency of extreme weather has been rising in recent years. A 3-year study of street trees was undertaken in Tokyo to determine whether: (i) street trees suffer from severe water stress in unusually hot summer; (ii) species respond differently to such climatic fluctuations; and (iii) street trees are also affected by nitrogen (N) deficiency, photoinhibition and aerosol pollution. During the study period (2010-12), midsummers of 2010 and 2012 were unusually hot (2.4-2.8 °C higher maximum temperature than the long-term mean) and dry (6-56% precipitation of the mean). In all species, street trees exhibited substantially decreased photosynthetic rate in the extremely hot summer in 2012 compared with the average summer in 2011. However, because of a more conservative stomatal regulation (stomatal closure at higher leaf water potential) in the hot summer, apparent symptoms of hydraulic failure were not observed in street trees even in 2012. Compared with Prunus × yedoensis and Zelkova serrata, Ginkgo biloba, a gymnosperm, was high in stomatal conductance and midday leaf water potential even under street conditions in the unusually hot summer, suggesting that the species had higher drought resistance than the other species and was less susceptible to urban street conditions. This lower susceptibility might be ascribed to the combination of higher soil-to-leaf hydraulic conductance and more conservative water use. Aside from meteorological conditions, N deficiency affected street trees significantly, whereas photoinhibition and aerosol pollution had little effect. The internal CO2 and δ(13)C suggested that both water and N limited the net photosynthetic rate of street trees simultaneously, but water was more limiting. From these results, we concluded that the potential risk of hydraulic failure caused by climatic extremes could be low in urban street trees in temperate regions. However, the size of the safety margin might be different between species.