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.     

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.     

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.     

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.     

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     

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.     

<Emphasis Type="Italic">Hedychium gardnerianum</Emphasis> Invasion into Hawaiian Montane Rainforest: Interactions Among Litter Quality,Decomposition Rate,and Soil Nitrogen Availability

Few studies have examined the invasion of understory species into closed-canopy forests and, despite inter-specific differences in litter quality and quantity between understory and dominant canopy trees, the influence of understory invasions on soil nitrogen (N) cycling remains unknown. This paper examines litter quality and decomposition of kahili ginger (Hedychium gardnerianum), an invasive understory herb, to determine the influence of this species on N cycling in a Hawaiian montane rainforest. To examine the potential feedback between increased soil N availability and litter decomposition, litter from the invasive ginger, a native tree, and native tree fern was collected from unfertilized and fertilized plots and decomposed in a reciprocal transplant design. Hedychium litter decomposed faster than litter from the two native species. Across species, decomposition rates were negatively correlated with litter lignin content. Despite rapid decomposition rates of Hedychium litter, soil nitrogen availability and rates of net mineralization in the soil were similar in invaded and uninvaded plots. Nitrogen cycling at this site may be more strongly influenced by native species, which contribute the most to overall stand biomass. A negative effect of fertilization on the decomposition of Hedychium litter suggests that a negative feedback between litter quality and soil N availability may exist over longer timescales.     

Physiological and morphological variation in Metrosideros polymorpha, a dominant Hawaiian tree species, along an altitudinal gradient: the role of phenotypic plasticity

Metrosideros polymorpha, a dominant tree species in Hawaiian ecosystems, occupies a wide range of habitats. Complementary field and common-garden studies of M. polymorpha populations were conducted across an altitudinal gradient at two different substrate ages to ascertain if the large phenotypic variation of this species is determined by genetic differences or by phenotypic modifications resulting from environmental conditions. Several characteristics, including ecophysiological behavior and anatomical features, were largely induced by the environment. However, other characteristics, particularly leaf morphology, appeared to be mainly determined by genetic background. Common garden plants exhibited higher average rates of net assimilation (5.8 μmol CO₂ m⁻² s⁻¹) and higher average stomatal conductance (0.18 mol H₂O m⁻² s⁻¹) than their field counterparts (3.0 μmol CO₂ m⁻² s⁻¹, and 0.13 mol H₂O m⁻² s⁻¹ respectively). Foliar δ¹³C of most common-garden plants was similar among sites of origin with an average value of −26.9‰. In contrast, mean values of foliar δ¹³C in field plants increased substantially from −29.5‰ at low elevation to −24.8‰ at high elevation. Leaf mass per unit area increased significantly as a function of elevation in both field and common garden plants; however, the range of values was much narrower in common garden plants (211–308 g m⁻² for common garden versus 107–407 g m⁻² for field plants). Nitrogen content measured on a leaf area basis in common garden plants ranged from 1.4 g m⁻² to 2.4 g m⁻² and from 0.8 g m⁻² to 2.5 g m⁻² in field plants. Photosynthetic nitrogen use efficiency (PNUE) decreased 50% with increasing elevation in field plants and only 20% in plants from young substrates in the common garden. This was a result of higher rates of net CO₂ assimilation in the common garden plants. Leaf tissue and cell layer thickness, and degree of leaf pubescence increased significantly with elevation in field plants, whereas in common garden plants, variation with elevation of origin was much narrower, or was entirely absent. Morphological characteristics such as leaf size, petiole length, and internode length decreased with increasing elevation in the field and were retained when grown in the common garden, suggesting a potential genetic basis for these traits. The combination of environmentally induced variability in physiological and anatomical characteristics and genetically determined variation in morphological traits allows Hawaiian M. polymorpha to attain and dominate an extremely wide ecological distribution not observed in other tree species. Received: 12 March 1997 / Accepted: 27 August 1997     

Foliar Nitrogen Dynamics and Nitrogen Resorption of a Sandy Shrub Salix gordejevii in Northern China

Resorption of nitrogen (N) from senescing leaves is an important conservation mechanism that allows plants to use the same N repeatedly. Seasonal variations in leaf nitrogen of mature green and senescing leaves and N resorption in Salix gordejevii Chang, a sandy shrub in northern China, were studied. Our objective was to compare N resorption of this Salix species that successfully occupy different habitats (shifting sandland, fixed sandland and lowland) with differences in soil N availability and moisture. Nitrogen concentrations in green and senescing leaves were higher in June and July. N resorption efficiency (percentage reduction of N between green and senescing leaves) was highest at shifting sandland, intermediate at fixed sandland, and lowest at lowland. There was a clear seasonal variation in N-resorption efficiency, with a lower value at the early growing season and a higher value during summer. N resorption efficiency was lower at the sites with higher soil N availability, suggesting that the efficiency of the resorption process is determined by the availability of the nutrient in the soil. Resorption from senescing leaves may play an important role in the nitrogen dynamics of sandy plants and reduce the nitrogen requirements for plant growth. We conclude that N resorption from senescing leaves in S. gordejevii was correlated to soil characteristics and higher N resorption on poor soils is a phenotypic adjustment by this species to maximize N-use at low availability.     

文冠果叶片养分元素含量的动态变化及再吸收特性

以文冠果叶片为试材,运用原子吸收光谱法分析了叶片养分元素含量的季节动态变化和再吸收效率。结果表明：N、P、K均呈下降趋势,是“稀释效应”和养分再吸收导致;Mg呈“单峰”曲线走势,与Mg的生理功能有关;Fe、Mn呈“V”字型走势,Cu呈“W”型,Zn呈“N”型,与树体吸收特性和不同的物候期有关。总体来看,养分元素含量顺序是：N>P>K>Mg>Fe>Zn>Mn>Cu,且不同时期又有所不同。大量元素之间存在显著的相关关系;微量元素间相关关系不显著（Fe、Zn除外）,Fe与N显著负相关,与拮抗作用有关。C/N呈升高趋势,差异显著;N/P呈降低趋势,差别不大。养分再吸收效率由大到小的顺序是：Mg>N>K>P,差异显著。微量元素由于移动性较差,不能被再吸收。N、P、K、Mg养分再吸收效率反映了文冠果较高的养分保存能力和养分利用效率。     

Litter Decomposition in a California Annual Grassland: Interactions Between Photodegradation and Litter Layer Thickness

In annual grasslands that experience a mediterranean-type climate, the synchrony between plant senescence and peak solar radiation over summer results in high litter sun exposure. We examined the decomposition of both shaded and sun-exposed litter over summer and inferred the effects of photodegradation from changes in mass loss and litter chemistry. The carry-over effects of summer litter exposure on wet season decomposition were also assessed, and the attenuation of photodegradation with litter layer thickness was used to estimate the proportion of grass litter lignin susceptible to photodegradation under different treatments of a factorial global change experiment. Over summer, mass loss from grass and forb litter exposed to ambient sunlight ranged from 8% to 10%, whereas lignin decreased in grass litter by approximately 20%. After one year of decomposition, mass losses from grass leaves exposed to sunlight over summer were more than double the mass losses from summer-shaded leaves. When shade litter layer thickness was varied, mass losses over summer for all treatments were also approximately 8%; however, lignin decreased significantly only in the low shade treatments (0–64 g m⁻² of shade litter). Aboveground production of annual grasses nearly quadrupled in response to the combined effects of N addition, elevated atmospheric CO₂, increased precipitation and warming. The estimated proportion of grass litter lignin experiencing full photodegradation ranged from 100% under ambient conditions to 31–62% in plots receiving the combined global change treatments. These results reveal an important role of sun exposure over summer in accelerating litter decomposition in these grasslands and provide evidence that future changes in the quantity of litter deposition may modulate the influence of photodegradation integrated across the litter layer.     

Decomposition of roots in loblolly pine: Effects of nutrient and water availability and root size class on mass loss and nutrient dynamics

The decomposition of plant-derived organic matter exerts strong control over the cycling of carbon and nutrients in terrestrial ecosystems and may be significantly altered by increased precipitation and nitrogen deposition associated with global change. It was the goal of this study to quantify the rate of belowground decomposition in an intact loblolly pine forest, and determine how this was affected by increased availability of water and nitrogen. A randomized complete-block factorial of irrigation and fertilization treatments was installed in an 8 yr old loblolly pine plantation in Scotland county, North Carolina. Fresh root samples of three size classes were buried in fiberglass mesh bags in January, 1994 and recovered at two-month intervals for two years. Samples were analyzed for percent mass remaining and contents of macro-nutrients. Roots decomposed in a two stage process: early in the incubation mass loss was correlated to size class and nutrient concentrations, but this correlation disappeared later in the incubation when rates of mass loss converged for all size classes. Decomposition was seldom affected by the irrigation and fertilization treatments, due to the buffering capacity of soil moisture and complex ecosystem-level responses to fertilization. Net mineralization of N, P, K, Ca, and Mg occurred in the smaller size classes of roots providing a source of these nutrients to the aggrading plantation for an estimated 2 to 15 years. The largest size class of roots was a sink for N, Ca, and Mg for the duration of this study, and was a source of P and K for an estimated 20 and 4 years, respectively. It is concluded that in moist temperate ecosystems belowground decomposition will be less affected by the projected increases in moisture and nutrient availability than will decomposition of the forest floor due to the buffering capacity of the soil. Further, small roots provide important sources of macro-nutrients for several decades to aggrading forests after large-scale disturbances such as harvesting of aboveground biomass.     

Changes in Asymbiotic, Heterotrophic Nitrogen Fixation on Leaf Litter of Metrosideros polymorpha with Long-Term Ecosystem Development in Hawaii

We measured nitrogenase activity (acetylene reduction) of asymbiotic, heterotrophic, nitrogen-fixing bacteria on leaf litter from the tree Metrosideros polymorpha collected from six sites on the Hawaiian archipelago. At all sites M. polymorpha was the dominant tree, and its litter was the most abundant on the forest floor. The sites spanned a soil chronosequence of 300 to 4.1 million y. We estimated potential nitrogen fixation associated with this leaf litter to be highest at the youngest site (1.25 kg ha^-1 y^-1), declining to between 0.05 and 0.22 kg ha^-1 y^-1 at the oldest four sites on the chronosequence. To investigate how the availability of weathered elements influences N fixation rates at different stages of soil development, we sampled M. polymorpha leaf litter from complete, factorial fertilization experiments located at the 300-y, 20,000-y and 4.1 million–y sites. At the youngest and oldest sites, nitrogenase activity on leaf litter increased significantly in the plots fertilized with phosphorus and “total” (all nutrients except N and P); no significant increases in nitrogenase activity were measured in leaf litter from treatments at the middle-aged site. The results suggest that the highest rates of N fixation are sustained during the “building” or early phase of ecosystem development when N is accumulating and inputs of geologically cycled (lithophilic) nutrients from weathering are substantial. Received 4 February 1999; accepted 29 March 2000.