Allometry of the quasi-pipe (qPipe) model for estimating tree leaf area and tree leaf mass applied to plant functional types.

The allometry of the pipe model quantifies the approximate proportionality between the tree leaf amount and the stem cross-sectional area at the crown base (ACB). It is useful for estimating and modeling carbon fixation abilities of trees but requires climbing the tree and is thus unsuitable for large-scale studies. Here, we adopted a previously proposed allometry (hereafter the quasi-pipe (qPipe) model allometry) formulating the relationship between the tree leaf amount and a surrogate of ACB, ACB_Est, calculated from tree dimensions measurable from the ground. Using published/unpublished data for 962 trees of 159 species collected between tropical rainforests and boreal forests, we established pipe and qPipe model allometries for evergreen-conifer, deciduous-conifer, evergreen-broadleaf, and deciduous-broadleaf plant functional types (PFTs). For the leaf area per tree (LA), allometric lines on a log-log plane were almost identical among the four PFTs in both models, with slopes of ~ 1. For the leaf mass per tree (LM), however, the allometric lines separated among the four PFTs in both models and had slopes greater than 1, indicating that the proportionality assumed in the pipe model held for LA but not LM. The applicability of the qPipe model in estimating the stand-scale leaf amount was further examined.

Vertical distribution of radiocesium concentrations among crown positions and year-to-year variation in four major tree species after the Fukushima Daiichi Nuclear Power Plant accident.

To evaluate the distribution of radiocesium (137Cs) among crown positions in trees after the Fukushima Daiichi Nuclear Power Plant accident, we collected foliage and branch samples from different crown positions of four major tree species (Chamaecyparis obtusa, Cryptomeria japonica, Pinus densiflora, and Quercus serrata) from 2011 to 2019 in northeast Japan. We divided the samples into current-year and more than 1-year-old groups (called old foliage and old branches), which sometimes included directly contaminated parts. The 137Cs activity concentration in dry foliage and branches was measured using a germanium semiconductor detector. There were complex differences in the relative 137Cs activity concentration among species and organ types (i.e., foliage and branches) among crown positions. The relative 137Cs activity concentration in current-year foliage was higher in the upper crowns of C. obtusa, but higher in lower crown positions in C. japonica. No differences among crown positions were observed in P. densiflora and Q. serrata. In current-year branches, the relative 137Cs concentration in Q. serrata was similar among crown positions but higher in the upper crown in P. densiflora. The concentrations in old foliage and old branches in all species tended to be higher in the lower crown. The factors causing these interspecific and organ type differences among crown positions may be related to the organ turnover rate, dilution effect due to different growth rates, and potassium distribution within the crown. No year-to-year variation was observed in most foliage and branches in all species, except for current-year branches of Q. serrata, old foliage in C. japonica and P. densiflora, and old branches in P. densiflora. Our long-term data on the interspecific and inter-organ patterns of contamination, focusing on variation among crown positions and year-to-year trends, might help to improve the estimation of 137Cs deposition and dynamics in polluted forest ecosystems.

Seasonal changes in radiocesium and potassium concentrations in current-year shoots of saplings of three tree species in Fukushima, Japan.

We studied seasonal changes in radiocesium (137Cs) activity and potassium concentrations in current-year leaves and branches of Pinus densiflora (naturally regenerated saplings), Cryptomeria japonica (planted saplings) and Quercus serrata (planted saplings and coppice shoots) in Fukushima, Japan. We collected current-year shoots from 10 individuals of each species over two growing seasons at intervals of 1-4 months, between June 2016 and December 2017. For the deciduous species Q. serrata, we also collected dead leaves that remained attached to branches in December to investigate reabsorption of 137Cs. All collected shoots were divided into leaves and branches, oven-dried, and ground; dry weights of each sample were recorded. 137Cs activity concentrations were measured using a germanium semiconductor detector. Potassium concentrations were quantified using inductively coupled plasma optical emission spectrometry (ICP-OES). Increases in dry weight were observed in both leaves and branches between May/June and August; growth then slowed considerably and virtually ceased after October. Clear seasonal changes in 137Cs activity concentrations were observed in both 2016 and 2017, regardless of tree species. Concentrations were higher in young leaves and branches during May and June, then decreased and changed relatively little from August to winter. Reduced 137Cs activity concentrations in dead leaves of Q. serrata were observed only in December 2017 (approximately 15% lower than in October). This reduction may indicate reabsorption of 137Cs in leaves prior to shedding. The changes in potassium concentrations were similar to those in 137Cs in both years. Potassium concentrations were higher in young leaves than in mature leaf and branch samples collected later in the year. A reduction of about 50% in the potassium concentrations in dead leaves of Q. serrata was also observed in December. A positive relationship between 137Cs and potassium concentrations in leaves and branches was observed in all species, except for planted Q. serrata. This relationship may indicate that 137Cs moves in tree shoots with potassium. Leaf and branch weight correlated negatively with 137Cs and potassium concentrations. Reduced concentrations may indicate dilution of these elements as a result of biomass increases over the growing season. Our results imply that irrespective of species, 137Cs exhibits seasonal variations resulting from dilution; these variations correspond with trends in potassium, with higher levels in young organs and decreased levels in older organs.

Temporal changes in the radiocesium distribution in forests over the five years after the Fukushima Daiichi Nuclear Power Plant accident.

To elucidate the temporal changes in the radiocesium distribution in forests contaminated by the Fukushima Daiichi Nuclear Power Plant accident, we monitored the 137Cs concentration and inventory within forests from 2011 to 2015 across nine plots containing variable tree species and different contamination levels. The 137Cs concentrations in needles and branches decreased exponentially at all coniferous plots, with effective ecological half-lives of 0.45-1.55 yr for needles and 0.83-1.69 yr for branches. By contrast, the 137Cs concentration in deciduous konara oak leaves did not change over the five years. The concentration of 137Cs in oak wood increased by 37-75%, whereas that in Japanese red pine decreased by 63% over the five years. In Japanese cedar and hinoki cypress, the 137Cs concentration in wood showed an increasing trend in half of the plots. The changes in 137Cs in the organic and mineral soil layers were not strongly related to the tree species or contamination level. Our multi-site, multi-species monitoring results revealed that the pattern of temporal changes in radiocesium in the 9 forest plots was similar overall; however, changes in 137Cs in needles/leaves and wood differed among tree species.

Vertical and seasonal variations in temperature responses of leaf respiration in a Chamaecyparis obtusa canopy.

Leaf respiration (R) is a major component of carbon balance in forest ecosystems. Clarifying the variability of leaf R within a canopy is essential for predicting the impact of global warming on forest productivity and the potential future function of the forest ecosystem as a carbon sink. We examined vertical and seasonal variations in short-term temperature responses of leaf R as well as environmental factors (light and mean air temperature) and physiological factors [leaf nitrogen (N), leaf mass per area (LMA), and shoot growth] in the canopy of a 10-year-old stand of hinoki cypress [Chamaecyparis obtusa (Sieb. et Zucc.) Endl.] in Kyushu, Japan. Leaf respiration rate adjusted to 20 °C (R20) exhibited evident vertical gradients in each season and was correlated with light, LMA and leaf N. In contrast, the temperature sensitivity of leaf R (Q10) did not vary vertically throughout the seasons. Seasonally, Q10 was higher in winter than in summer and was strongly negatively correlated to mean air temperature. A negative correlation of R20 with mean air temperature was also observed for each of the three canopy layers. These results clearly indicate that leaf R was able to adjust to seasonal changes in ambient temperature under field conditions and down-regulate during warmer periods. We also found that the degree of thermal acclimation did not vary with canopy position. Overall, our results suggest that vertical and seasonal variations in temperature responses of leaf R within a hinoki cypress canopy could be predicted by relatively simple parameters (light and temperature). There was an exception of extremely high R20 values in April that may have been due to the onset of shoot growth in spring. Understanding thermal acclimation and variations in leaf R within forest canopies will improve global terrestrial carbon cycle models.

Seasonal variations in the stable oxygen isotope ratio of wood cellulose reveal annual rings of trees in a Central Amazon terra firme forest.

In Amazonian non-flooded forests with a moderate dry season, many trees do not form anatomically definite annual rings. Alternative indicators of annual rings, such as the oxygen (δ(18)Owc) and carbon stable isotope ratios of wood cellulose (δ(13)Cwc), have been proposed; however, their applicability in Amazonian forests remains unclear. We examined seasonal variations in the δ(18)Owc and δ(13)Cwc of three common species (Eschweilera coriacea, Iryanthera coriacea, and Protium hebetatum) in Manaus, Brazil (Central Amazon). E. coriacea was also sampled in two other regions to determine the synchronicity of the isotopic signals among different regions. The annual cyclicity of δ(18)Owc variation was cross-checked by (14)C dating. The δ(18)Owc showed distinct seasonal variations that matched the amplitude observed in the δ(18)O of precipitation, whereas seasonal δ(13)Cwc variations were less distinct in most cases. The δ(18)Owc variation patterns were similar within and between some individual trees in Manaus. However, the δ(18)Owc patterns of E. coriacea differed by region. The ages of some samples estimated from the δ(18)Owc cycles were offset from the ages estimated by (14)C dating. In the case of E. coriacea, this phenomenon suggested that missing or wedging rings may occur frequently even in well-grown individuals. Successful cross-dating may be facilitated by establishing δ(18)Owc master chronologies at both seasonal and inter-annual scales for tree species with distinct annual rings in each region.

Characteristics of initial deposition and behavior of radiocesium in forest ecosystems of different locations and species affected by the Fukushima Daiichi Nuclear Power Plant accident.

After the Fukushima Daiichi Nuclear Power Plant (FDNPP) accident, information about stand-level spatial patterns of radiocesium initially deposited in the surrounding forests was essential for predicting the future dynamics of radiocesium and suggesting a management plan for contaminated forests. In the first summer (approximately 6 months after the accident), we separately estimated the amounts of radiocesium ((134)Cs and (137)Cs; Bq m(-2)) in the major components (trees, organic layers, and soils) in forests of three sites with different contamination levels. For a Japanese cedar (Cryptomeria japonica) forest studied at each of the three sites, the radiocesium concentration greatly differed among the components, with the needle and organic layer having the highest concentrations. For these cedar forests, the proportion of the (137)Cs stock in the aboveground tree biomass varied from 22% to 44% of the total (137)Cs stock; it was 44% in highly contaminated sites (7.0 × 10(5) Bq m(-2)) but reduced to 22% in less contaminated sites (1.1 × 10(4) Bq m(-2)). In the intermediate contaminated site (5.0-5.8 × 10(4) Bq m(-2)), 34% of radiocesium was observed in the aboveground tree biomass of the Japanese cedar stand. However, this proportion was considerably smaller (18-19%) in the nearby mixed forests of the Japanese red pine (Pinus densiflora) and deciduous broad-leaved trees. Non-negligible amounts of (134)Cs and (137)Cs were detected in both the sapwood and heartwood of all the studied tree species. This finding suggested that the uptake or translocation of radiocesium had already started within 6 months after the accident. The belowground compartments were mostly present in the organic layer and the uppermost (0-5 cm deep) mineral soil layer at all the study sites. We discussed the initial transfer process of radiocesium deposited in the forest and inferred that the type of initial deposition (i.e., dry versus wet radiocesium deposition), the amount of rainfall after the accident, and the leaf biomass by the tree species may influence differences in the spatial pattern of radiocesium by study plots. The results of the present study and further studies of the spatial pattern of radiocesium are important for modeling future radiocesium distribution in contaminated forest ecosystems.

Mixed-power scaling of whole-plant respiration from seedlings to giant trees.

The scaling of respiratory metabolism with body mass is one of the most pervasive phenomena in biology. Using a single allometric equation to characterize empirical scaling relationships and to evaluate alternative hypotheses about mechanisms has been controversial. We developed a method to directly measure respiration of 271 whole plants, spanning nine orders of magnitude in body mass, from small seedlings to large trees, and from tropical to boreal ecosystems. Our measurements include the roots, which have often been ignored. Rather than a single power-law relationship, our data are fit by a biphasic, mixed-power function. The allometric exponent varies continuously from 1 in the smallest plants to 3/4 in larger saplings and trees. Therefore, our findings support the recent findings of Reich et al. [Reich PB, Tjoelker MG, Machado JL, Oleksyn J (2006) Universal scaling of respiratory metabolism, size, and nitrogen in plants. Nature 439:457-461] and West, Brown, and Enquist [West GB, Brown JH, Enquist BJ (1997) A general model for the origin of allometric scaling laws in biology. Science 276:122 -126.]. The transition from linear to 3/4-power scaling may indicate fundamental physical and physiological constraints on the allocation of plant biomass between photosynthetic and nonphotosynthetic organs over the course of ontogenetic plant growth.