Interactions between Litter Lignin and Nitrogenitter Lignin and Soil Nitrogen Availability during Leaf Litter Decomposition in a Hawaiian Montane Forest

Previous work in a young Hawaiian forest has shown that nitrogen (N) limits aboveground net primary production (ANPP) more strongly than it does decomposition, despite low soil N availability. In this study, I determined whether (a) poor litter C quality (that is, high litter lignin) poses an overriding constraint on decomposition, preventing decomposers from responding to added N, or (b) high N levels inhibit lignin degradation, lessening the effects of added N on decomposition overall. I obtained leaf litter from one species, Metrosideros polymorpha, which dominates a range of sites in the Hawaiian Islands and whose litter lignin concentration declines with decreasing precipitation. Litter from three dry sites had lignin concentrations of 12% or less, whereas litter from two wet sites, including the study site, had lignin concentrations of more than 18%. This litter was deployed 2.5 years in a common site in control plots (receiving no added nutrients) and in N-fertilized plots. Nitrogen fertilization stimulated decomposition of the low-lignin litter types more than that of the high-lignin litter types. However, in contrast to results from temperate forests, N did not inhibit lignin decomposition. Rather, lignin decay increased with added N, suggesting that the small effect of N on decomposition at this site results from limitation of decomposition by poor C quality rather than from N inhibition of lignin decay. Even though ANPP is limited by N, decomposers are strongly limited by C quality. My results suggest that anthropogenic N deposition may increase leaf litter decomposition more in ecosystems characterized by low-lignin litter than in those characterized by high-lignin litter. Received 26 October 1999; accepted 2 June 2000.     

An Exotic Tree Alters Decomposition and Nutrient Cycling in A Hawaiian Montane Forest

We evaluated the effects of the exotic tree Fraxinus uhdei on decomposition dynamics and nutrient turnover in a montane Hawaiian rainforest. We used reciprocal transplants of litterbags between forests dominated by Fraxinus and by the native Metrosideros polymorpha to distinguish between endogenous (litter quality) and exogenous (for example, microclimate, nutrient availability, microbial and invertebrate communities) effects of Fraxinus on mass loss and nutrient dynamics of decomposing litter. Fraxinus produced greater quantities of litter that was thinner, had higher N and P concentrations, and lower concentrations of lignin and soluble polyphenols. Microbes decomposing Fraxinus litter produced fewer enzymes involved in N and P acquisition and more of those involved in cellulose degradation. Differences in litter quality and microbial activity resulted in a strong effect of litter type on rates of mass loss, whereby Fraxinus litter decomposed and released nutrients at nearly twice the rate of Metrosideros litter (k=0.82 versus 0.48), regardless of site of decomposition. Although site of decomposition had no effect on rates of litter mass loss, Fraxinus litter decomposed under a Fraxinus canopy mineralized approximately 20% less P after one year than Fraxinus litter decomposed under a Metrosideros canopy. Furthermore, Fraxinus litter decomposed under a Fraxinus canopy immobilized greater amounts of N and P in the early stages of decay, suggesting that the large amounts of N and P in Fraxinus litterfall have raised nutrient availability to decomposers in the forest floor. Greater immobilization of N and P under a Fraxinus canopy may act as a governor on rates of nutrient cycling, limiting the degree to which Fraxinus invasion accelerates N and P cycling in this system.     

Foliar Nutrient Dynamics and Resorption in Naturalized Lonicera maackii (Caprifoliaceae) Populations in Ohio,USA

Foliar phosphorus (P) and nitrogen (N) concentrations and nutrient resorption in the forest understory shrub Lonicera maackii (Rupr.) Maxim (Caprifoliaceae) were measured along contiguous topographic gradients in two southwestern Ohio forests during 1992–1994. Mean summer foliar P varied significantly among topographic positions (but not sites or years), with uplands having greatest P concentrations and bottomlands exhibiting the lowest. Unlike for P, the mean summer foliar N concentrations varied little among sites, topographic positions, and years. Mean absolute and proportional P resorption ranged from 0.48 mg/dm² (33.7%) in slope positions to 0.80 mg/dm² (53.1%) in bottomland positions. Repeated-measures analysis of variance (RMANOVA) for P resorption indicated significant topographic and year effects, a site × year and a site × year × topographic interaction. Mean absolute and proportional foliar N resorption ranged from 6.82 mg/dm² (30.7%) in bottomlands to 8.41 mg/dm² (37.3%) in slope positions. RMANOVA indicated a significant topographic effect for both absolute and proportional N resorption and a significant year effect for absolute N resorption. These significant year effects for P and N stemmed from lowest resorption of nutrients in 1993. The results for P resorption support the hypothesis that foliar resorption is greater in forested sites with lower P fertility. However, resorption rates for N did not support the hypothesis clearly, as slopes with intermediate N availability had greater N resorption rates than did N-rich bottomlands.     

Patterns of Carbon, Nitrogen and Phosphorus Dynamics in Decomposing Foliar Litter in Canadian Forests

We examined the patterns of nitrogen (N) and phosphorus (P) gain, retention or loss in ten foliar tissues in a litterbag experiment over 6 years at 18 upland forest sites in Canada, ranging from subarctic to cool temperate. N was usually retained in the decomposing litter until about 50% of the original C remained. The peak N content in the litter was observed at between 72 and 99% of the original C remaining, with C:N mass quotients between 37 and 71 (mean 55). The rate of N release from the litters was not related to the original N concentration, which may be associated with the generally narrow range (0.59–1.28% N) in the litters. P was immediately lost from all litters, except beech leaves, with critical litter C:P mass quotients for P release being in the range 700–900. The rate of P loss was inversely correlated with the original litter P concentration, which ranged from 0.02 to 0.13%. The soil underlying the litterbags influenced the pattern of N and P dynamics in the litters; there were weak correlations between the N and P remaining at 60% C remaining in the litters and the C:N and C:P quotients of the surface layer of the soil. There was a trend for higher N and P retention in the litter at sites with lower soil C:N and N:P quotients, respectively. Although there was a large variation in C:N, C:P and N:P quotients in the original litters (29–83, 369–2122 and 5–26, respectively), and some variation in the retention or loss of N and P in the early stages of decomposition, litters converged on C:N, C:P and N:P quotients of 30, 450 and 16, when the C remaining fell below 30%. These quotients are similar to that found in the surface organic matter of these ecosystems.     

Asymbiotic Nitrogen Fixation and Litter Decomposition on a Long Soil-Age Gradient in Hawaiian Montane Rain Forest1

We determined rates of decomposition and asymbiotic nitrogen fixation in the leaf litter of Cheirodendron spp. on the Hawaiian Islands. Leaf litter was collected from four sites on a long soil-age gradient (300 yr to 4.1 M yr) and decomposed at two sites that differed widely in substrate age and nutrient availability. Rates of decomposition were higher in litter decomposed at the older site, where nutrient availability was greater. A substantial amount of nitrogen and phosphorus immobilization occurred in litter decomposed at the older site, with more immobilization occurring in litter with lower initial nitrogen and phosphorus concentrations, suggesting both supply and demand controls on nutrient immobilization. Potential rates of nitrogen fixation were very low in the first 25 d (0–5 nmol acetylene/gdw/h), rose to much higher rates by 70 d (20–45 nmol), and then declined by 140 d. We found no significant difference in rates of potential nitrogen fixation between sites of decomposition, but there was a strong substrate effect, with higher rates in litter with low lignin, low nitrogen, and high phosphorus. Where significant immobilization of nitrogen occurred for decomposing Cheirodendron, nitrogen fixation could have comprised no more than 10 percent of immobilized nitrogen. Overall, rates of nitrogen fixation were dependent on the source of the decomposing substrate but not on the site of decomposition, while short-term decomposition and nutrient immobilization were strongly dependent on the site of decomposition but not as much on the source of the decomposing substrate.     

Litter Decomposition Within the Canopy and Forest Floor of Three Tree Species in a Tropical Lowland Rain forest,Costa Rica

The rain forest canopy hosts a large percentage of the world's plant biodiversity, which is maintained, in large part, by internal nutrient cycling. This is the first study to examine the effects of site (canopy, forest floor) and tree species (Dipteryx panamensis, Lecythis ampla, Hyeronima alchorneoides) on decay rates of a common substrate and in situ leaf litter in a tropical forest in Costa Rica. Decay rates were slower for both substrates within the canopy than on the forest floor. The slower rate of mass loss of the common substrate in the canopy was due to differences in microclimate between sites. Canopy litter decay rates were negatively correlated with litter lignin:P ratios, while forest floor decay rates were negatively correlated with lignin concentrations, indicating that the control of litter decay rates in the canopy is P availability while that of the forest floor is carbon quality. The slower cycling rates within the canopy are consistent with lower foliar nutrient concentrations of epiphytes compared with forest floor-rooted plants. Litter decay rates, but not common substrate decay rates, varied among tree species. The lack of variation in common substrate decay among tree species eliminated microclimatic variation as a possible cause for differences in litter decay and points to variation in litter quality, nutrient availability and decomposer community of tree species as the causal factors. The host tree contribution to canopy nutrient cycling via litter quality and inputs may influence the quality and quantity of canopy soil resources.     

Extracellular Enzyme Activities and Carbon Chemistry as Drivers of Tropical Plant Litter Decomposition

Litter quality parameters such as nitrogen and lignin content correlate with decomposition rates at coarse scales, but fine‐scale mechanisms driving litter decomposition have proven more difficult to generalize. One potentially important driver of decomposition is the activity of extracellular enzymes that catalyze the degradation of complex compounds present in litter. To address the importance of this mechanism, we collected 15 Hawaiian plant litter types and decomposed them in fertilized and control plots for up to two years. We measured litter nutrient content and carbon chemistry prior to decomposition, as well as extracellular enzyme activities, mass loss, and litter nutrient content over time. We found that water‐soluble carbon content, cellobiohydrolase activities, and polyphenol oxidase activities were significantly correlated with mass loss. Enzyme activities and decomposition rate constants both varied significantly by litter type, and fertilization increased mass loss rates in five litter types. Some litter types that decayed faster under fertilization also showed time‐dependent increases in carbon‐degrading enzyme activities, but others decayed faster independent of enzyme changes. These results suggest that extracellular enzyme activities partially determine litter decomposition rates, but high soluble carbon content may circumvent the requirement for enzyme‐catalyzed decomposition.     

Contrasting Effects of Substrate and Fertilizer Nitrogen on the Early Stages of Litter Decomposition

Commonly observed positive correlations between litter nitrogen (N) concentrations and decomposition rates suggest that N frequently limits decomposition in its early stages. However, numerous studies have found little, if any, effect of N fertilization on decomposition. I directly compared internal substrate N and externally supplied inorganic N effects on decomposition in sites varying in soil N availability. I decomposed eight substrates (with initial %N from 0–2.5) in control and N-fertilized plots at eight grassland and forest sites in central Minnesota. N fertilization increased decomposition at only two of eight sites, even though decomposition was positively related to litter N at all sites and to soil N availability across sites. The effect of externally supplied N on decomposition was independent of litter N concentration, but was greater at sites with low N availability. The inconsistent effects of substrate and externally supplied N may have arisen because decomposers use organic N preferentially as an N source; because inorganic N availability across sites or with fertilization induced changes in microbial community attributes (for example, lower C:N or greater efficiency) that reduced the response of decomposition to increased inorganic N supply; or because the positive correlation between litter N or site N availability with decomposition was spurious, caused by tight correlations between litter or site N and some other factor that truly limited decomposition. These inconsistent effects of substrate N and external N supply on decomposition suggest that the oft-observed relationship between litter N and decomposition may not indicate N limitation of decomposition.     

Inter‐specific Variation in Foliar Nutrients and Resorption of Nine Canopy‐tree Species in a Secondary Neotropical Rain Forest

The influence of environmental gradients on the foliar nutrient economy of forests has been well documented; however, we have little understanding of what drives variability among individuals within a single forest stand, especially tropical forests. We evaluated inter‐ and intra‐specific variation in nutrient resorption, foliar nutrient concentrations and physical leaf traits of nine canopy tree species within a 1‐ha secondary tropical rain forest in northeastern Costa Rica. Both nitrogen (N) and phosphorus (P) resorption efficiency (RE) and proficiency of the nine tree species varied significantly among species, but not within. Both N and P RE were significantly negatively related to leaf specific strength. Green leaf N and P concentrations were strongly negatively related to leaf mass per area, and senesced leaf nutrient concentrations were significantly positively related to green leaf nutrient concentrations. This study reveals a strong influence of physical leaf traits on foliar nutrient and resorption traits of co‐occurring species in a secondary wet tropical forest stand.     

The Effect of Water on Decomposition Dynamics in Mesic to Wet Hawaiian Montane Forests

I used a mesic to wet precipitation gradient on Maui, Hawaii, to test whether variation in rainfall regulates decomposition in tropical wet forest. Decomposition rates of leaves and roots from the dominant tree species, Metrosideros polymorpha, were measured at six sites similar in temperature regime, parent material, ecosystem age, vegetation, and topographical relief, whereas mean annual precipitation (MAP) at these six sites varied from 2200 to over 5000 mm/y. In situ decomposition rates of leaves placed on the soil surface declined by a factor of 6.4 with increased precipitation, whereas the decomposition rate of roots placed below ground declined by a factor of 2.3 across the gradient. Leaves collected from the 2200-mm site and placed at all sites on the gradient decomposed faster on the soil surface than they did below ground, whereas both above- and belowground decomposition rates of the common leaves decreased by a factor of 2.5 with increased precipitation. Of the environmental variables that changed with MAP, soil oxygen availability appeared to be the proximal factor that limited decomposition rates across the gradient, both above and below ground. When plant tissue collected from all sites across the gradient was decomposed at a common site, leaves from the wettest sites decomposed almost three times more slowly than leaves from the mesic sites. In contrast, roots from across the gradient all decomposed at a similar rate in a common site. Of tissue chemistry variables, high lignin concentration was correlated consistently with slow decomposition for roots and leaves. These results suggest that soil oxygen limitation combined with poorly decomposable leaves caused slower rates of decomposition and nutrient release with increased rainfall in these upland forests. Received 14 April 2000; Accepted 11 December 2000.     

Changes in Production and Nutrient Cycling across a Wetness Gradient within a Floodplain Forest

Floodplain forest ecosystems are highly valuable to society because of their potential for water quality improvement and vegetation productivity, among many other functions. Previous studies have indicated that hydrology influences productivity but that the relationship between hydroperiod and productivity is a complex one. Consequently, we compared multiple indexes of productivity, nutrient circulation, and hydroperiod among three communities on the Flint River floodplain, Georgia, that differed in terms of inundation frequency. We hypothesized that (a) the wettest community would have the lowest total net primary production (NPP) values because of saturated soil conditions; (b) as wetness increases, nutrient circulation in litterfall would decrease because of the hypothesized lower productivity in the wetter community; and (c) as wetness increases, internal translocation would become more efficient. The study site was partitioned into three wetness types—somewhat poorly drained (SPD), intermediate (I) and poorly drained (PD). We found that belowground biomass was greatest on the SPD, litterfall was similar for all three sites, and that woody biomass current annual increment (CAI) was greatest in the PD community. However, when the three variables were totaled for each site, the PD had the greatest NPP, thus disproving hypothesis (a). For hypothesis (b), we observed that P content in litterfall, although not significant, followed the predicted trend; nitrogen (N) content displayed the opposite pattern (PD > I > SPD). As wetness increased, internal translocation became more efficient for phosphorus (support for hypothesis [c]), but the SPD community was more efficient at retranslocating N (contradiction of hypothesis [c]). Received 19 June 2000; accepted 19 October 2000.     

红松人工林枝叶分解速度及养分动态的研究

修枝作为培育无节良材的重要经营措施已被广泛应用于生产,修落的枝叶能为林地提供大量的有机物,分解后提高林地土壤肥力。为了探明红松人工纯林枝叶分解过程中各种营养元素的变化规律,我们于1984年至1986年在草河     

Resorption efficiency of leaf nutrients in woody plants on Mt. Dongling of Beijing,North China

Aims To explore resorption efficiency of nitrogen (NRE) and phosphorus (PRE) of woody plants in relation to soil nutrient availability, climate and evolutionary history, in North China.Methods We measured concentrations of nitrogen ([N]) and phosphorus ([P]) in both full expanded mature green and senescent leaves of the same individuals for 88 woody species from 10 sites of Mt. Dongling, Beijing, China. We built a phylogenetic tree for all these species and compared NRE and PRE among life forms (trees, shrubs and woody lianas) and between functional groups (N-fixers and non-N-fixers). We then explored patterns of NRE and PRE along gradients of mean annual temperature (MAT), soil inorganic N and available P, and phylogeny using a general linear model.Important findings Mass-based NRE (NRE m) and PRE (PRE m) averaged 57.4 and 61.4%, respectively, with no significant difference among life forms or functional groups. Neither NRE m nor PRE m exhibited significant phylogenetic signals, indicating that NRE m and PRE m were not phylogenetically conserved. NRE m was not related to [N] in green leaves; PRE m was positively correlated with [P] in green leaves; however, this relationship disappeared for different groups. NRE m decreased with [N] in senescent leaves, PRE m decreased with [P] in senescent leaves, for all species combined and for trees and shrubs. NRE m decreased with soil inorganic N for all species and for shrubs; PRE m did not exhibit a significant trend with soil available P for all species or for different plant groups. Neither NRE m nor PRE m was significantly related to MAT for overall species and for species of different groups.     

Nitrogen Fixation in Bryophytes, Lichens, and Decaying Wood along a Soil-age Gradient in Hawaiian Montane Rain Forest

We determined rates of acetylene reduction and estimated total nitrogen fixation associated with bryophytes, lichens, and decaying wood in Hawaiian montane rain forest sites with underlying substrate ranging in age from 300 to 4.1 million years. Potential N fixation ranged from ca 0.2 kg/ha annually in the 300‐year‐old site to ca 1 kg/ha annually in the 150,000‐year‐old site. Rates of acetylene reduction were surprisingly uniform along the soil‐age gradient, except for high rates in symbiotic/associative fixers at the 150,000‐year‐old site and in heterotrophic fixers at the 2100‐year‐old site. Low fixation at the youngest site, where plant production is known to be N‐limited, suggests that demand for N alone does not govern N fixation. Total N fixation was highest in sites with low N:P ratios in leaves and stem wood, perhaps because epiphytic bryophytes and lichens depend on canopy leachate for mineral nutrients and because heterotrophic fixation is partly controlled by nutrient supply in the decomposing substrate; however, differences in substrate cover, rather than in fixation rates, had the largest effect on the total N input from fixation at these sites.     

Litter Decomposition and Nutrient Release in a Tropical Seasonal Rain Forest of Xishuangbanna,Southwest China1

We studied litter decomposition and nutrient release in a tropical seasonal rain forest of Xishuangbanna, Southwest China. The monthly decay rates (k) of leaf litter ranged from 0.02 to 0.21/mo, and correlated with rainfall and soil moisture. Annual k values for leaf litter (1.79/yr) averaged 4.2 times of those for coarse wood (2.5–3.5 cm in diameter). The turnover coefficients of forest floor mass (annual litterfall input/mean floor mass) were: 4.11/yr for flowers and fruits, 2.07/yr for leaves, and 1.17/yr for fine wood (≤2 cm in diameter), with resident time decreasing from fine woods (0.85 yr) to leaves (0.48 yr) and to flower and fruits (0.24 yr). Nutrient residence times in the forest floor mass were ranked as: Ca (1.0 yr) > P (0.92 yr) > Mg (0.64 yr) > N (0.36 yr) > K (0.31 yr). Our data suggest that rates of litter decomposition and nutrient release in the seasonal rain forest of Xishuangbanna are slower than those in typical lowland rain forests, but similar to those in tropical semideciduous forests.     

Nutrient Limitation in a Fire‐derived,Nitrogen‐rich Hawaiian Grassland1

Grasslands created by grass invasions into shrublands or woodlands followed by fire are now a dominant feature of many seasonally dry environments. In Hawaii Volcanoes National Park, introduced perennial grasses dominate grasslands created by fire in grass‐invaded woodlands. In a previous study, we found that net primary production in these grasslands is substantially lower than in unburned woodlands. Yet, our estimates of annual net nitrogen (N) mineralization showed higher rates in these savannas than in the unburned woodlands, rates that appear to greatly exceed annual N demand by the vegetation. We therefore hypothesized that N should not be limited to the plants growing in these sites. We tested this hypothesis with a 2‐yr fertilization experiment. At peak biomass, we found a 30 percent increase in live biomass in plots with N added and no increase in production with only phosphorus (P) added. N and P together were synergistic, suggesting that co‐limitation or P limitation becomes important when N is more available. Plants responded to added N by increasing individual leaf area and shoot length by over 50 percent. Tissue N was higher with added N; hence, biomass N was substantially higher. Tissue P concentrations declined with N addition but were elevated by P addition despite lack of a growth response to P alone. Overall, N limitation exists despite high annual rates of net N mineralization, and co‐limitation of production by P may occur when N is abundant. Here, asynchrony between plant nutrient demand and N availability may contribute to N limitation.     

福建东山短枝木麻黄小枝氮磷含量及其再吸收率季节动态

对福建东山赤山林场短枝木麻黄纯林小枝中的N和P含量、N ∶ P比、养分再吸收率的季节动态进行研究.结果表明,随着小枝的成熟和衰老,各季节N和P的含量依次降低,其中衰老小枝中N和P含量分别为(3.97±0.21)～(8.64±0.58) mg · g~(-1)和(0.04±0.03)～(0.10±0.03) mg · g~(-1),基本上表现为N和P的完全再吸收.成熟小枝N ∶ P比的季节动态介于(18.67±3.24)～(37.98±1.32)之间,均高于16,表明木麻黄的生长受到不同程度的P限制.N、P再吸收率的季节变化分别为(51.02±4.66)%～(63.00±8.61)%和(81.24 ±5.08)%～(91.78±5.84)%,P再吸收率显著高于N.N和P的再吸收率之间以及成熟叶中N、P含量与二者再吸收率之间没有显著相关性,但在衰老叶中存在显著负相关,表明养分再吸收程度越高,其再吸收率就越高.因此,木麻黄通过小枝衰老过程中的养分再吸收,减少养分的损失,从而适应贫瘠的立地条件.