James H. Fownes

Age-related decline in forest productivity : Pattern and process

Publisher Summary This chapter reviews the evidence for the pattern of growth decline with age and discusses the evidence for the mechanisms that may be responsible. It begins with an overview of the proposed mechanisms. The chapter also presents a framework for understanding the changes in stand productivity with age, because many of the proposed mechanisms are linked and affect carbon allocation. The available information on the importance of various mechanisms behind growth decline, in the context of the stand carbon cycle is presented. The common patterns of a decline in stand leaf area and leaf photosynthetic capacity suggest a new model of carbon balance with stand development. In this model, photosynthesis and above-ground dry-matter production increase with canopy development. After the forest reaches a maximum leaf area, photosynthesis and dry-matter production decline as leaf area, photosynthetic capacity, and photosynthesis also decline. The model assumes that allocation to respiration and below ground to roots and symbionts is a constant fraction of assimilation over the life of a forest stand.

Changes in Soil Phosphorus Fractions and Ecosystem Dynamics across a Long Chronosequence in Hawaii

We tested the Walker and Syers (1976) conceptual model of soil development and its ecological implications by analyzing changes in soil P, vegetation, and other ecosystem properties on a soil chronosequence with six sites ranging in age from 300 yr to 4.1 x 10 6 yr. Climate, dominant vegetation, slope, and parent material of all of the sites were similar. As fractions of total P, the various pools of soil phosphorus behaved very much as predicted by Walker and Syers. HCl-extractable P (presumably primary mineral phosphates) comprised 82% of total P at the 300-yr-old site, and then decreased to 1% at the 20,000-yr-old site. Organic phosphorus increased from the youngest site to a maximum at the 150000 yr site, and then declined to the 4.1 x 10 6 yr site. Occluded (residual) P increased steadily with soil age. In contrast to the Walker and Syers model, we found the highest total P at the 150000-yr-old site, rather than at the onset of soil development, and we found that the non-occluded, inorganic P fraction persisted through to the oldest chronosequence site. Total soil N and C increased substantially from early to middle soil development in parallel with organic P, and then declined through to the oldest site. Readily available soil P, NH 4 + , and NO 3 - were measured using anion and cation exchange resin bags. P availability increased and decreased unimodally across the chronosequence. NH 4 + and NO 3 - pools increased through early soil development, but did not systematically decline late in soil development. In situ decomposition rates of Metrosideros polymorpha litter were highest at two intermediate-aged sites with high soil fertility (20000 yr and 150000 yr), and lowest at the less-fertile beginning (300 yr) and endpoint (4.1 x 10 6 yr) of the chronosequence. M. polymorpha leaves collected from these same four sites, and decomposed in a common site, suggested that leaves from intermediate-aged sites were inherently more decomposable than those from the youngest and oldest sites. Both litter tissue quality and the soil environment appeared to influence rates of decomposition directly. The highest mean soil N 2 O emissions (809 μg.m -2 .d -1 ) were measured at the 20 000-yr-old site, which also had the highest soil nitrogen fertility status. Plant communities at all six chronosequence sites were dominated primarily by M. polymorpha, and to a lesser extent by several other genera of trees and shrubs. There were, however, differences in overall vegetation community composition among the sites. Using a detrended correspondence analysis (DECORANA), we found that a high proportion of species variance among the sites (eigenvalue = 0.71) can be explained by site age and thus soil developmental stage. Overall, long-term soil development across the chronosequence largely coincides with the conceptual model of Walker and Syers (1976). How P is distributed among different organic and inorganic fractions at a given stage of soil development provides a useful context for evaluating contemporary cycling of P and other nutrients, and for determining how changes in P availability might affect diverse ecosystem processes.

Nitrogen mineralization from leaves and litter of tropical plants : relationship to nitrogen, lignin and soluble polyphenol concentrations

Abstract Rates and patterns of nitrogen mineralization from decomposing plant materials are known to be affected by initial concentrations of N, lignin and soluble polyphenols, but published results show differences in which properties correlated best with N release. We incubated soil with fresh leaves and litter from 12 species commonly used in tropical agroforestry systems, including both legumes and non-legumes, to estimate how release of mineral N was affected by chemical composition. After 16 weeks, net accumulation of mineral N in incubations ranged from 81% of initial N in fresh Sesbania sesban to net depletion equivalent to 70% of initial N in Casuarina equisetifolia litter. Fresh legume leaves generally had net N accumulation, whereas legume litter and non-legume fresh leaves and litter had net depletion. At all sampling dates, the percent of leaf N accumulated was most highly correlated with initial N concentration. Phosphorus and the ratios lignin:N and (lignin + polyphenol):N also strongly correlated with N accumulation, although these indices themselves were correlated with N. We conclude that differences among previous studies in best chemical predictors of mineralization rates arose from relatively small ranges in chemical composition of materials used. Over our wide range of materials, initial soluble polyphenols were secondary to initial N, which explained most of the variation in N accumulation or depletion.

AN EXPERIMENTAL TEST OF THE CAUSES OF FOREST GROWTH DECLINE WITH STAND AGE

The decline in aboveground wood production after canopy closure in even-aged forest stands is a common pattern in forests, but clear evidence for the mechanism causing the decline is lacking. The problem is fundamental to forest biology, commercial forestry (the decline sets the rotation age), and to carbon storage in forests. We tested three hypotheses about mechanisms causing the decline in wood growth by quantifying the complete carbon budget of developing stands for over six years (a full rotation) in replicated plantations of Eucalyptus saligna near Pepeekeo, Hawaii. Our first hypothesis was that gross primary production (GPP) does not decline with stand age, and that the decline in wood growth results from a shifft in partitioning from wood production to respiration (as tree biomass accumulates), total belowground carbon allocation (as a result of declining soil nutrient supply), or some combination of these or other sinks. An alternative hypothesis was that GPP declines with stand age and that the decline in aboveground wood production is proportional to the decline in GPP. A decline in GPP could be driven be reduced canopy leaf area and photosynthetic capacity resulting from increasing nutrient limitation, increased abrasion between tree canopies, lower turgor pressure to drive foliar expansion, or hydraulic limitation of water flux as tree height increases. A final hypothesis was a combination of the first two: GPP declines, but the decline in wood production is disproportionately larger because partitioning shifts as well. We measured the entire annual carbon budget (aboveground production and respiration, total belowground carbon allocation [TBCA], and GPP) from 0.5 years after seedling planting through 6 1/2 years (when trees were ~25m tall). The replicated plots included two densities of trees (1111 trees/ha and 10 000 trees/ha) to vary the ratio of canopy leaf mass to wood mass in the individual trees, and three fertilization regimes (minimal, intensive, and minimal followed by intensive after three years) to assess the role of nutrition in shaping the decline in GPP and aboveground wood production. The forest closed its canopy in 1-2 years, with peak aboveground wood production, coinciding with canopy closure, of 1.2-1.8 kg C.m-2yr-1. Aboveground wood production declined from 1.4 kg C.m-2yr-1 at age 2 to 0.60 kg C.m-2yr-1 at age 6. Hypothesis 1 failed: GPP declined from 5.0 kg C.m-2yr-1 at age 2 to 3.2 kg C.m-2yr-1 at age 6. Aboveground woody respiration declined from 0.66 kg C.m-2yr-1 at age 2 to 0.22 kg C.m-2yr-1 at age 6 and TBCA declined from 1.9 kg C.m-2yr-1 at age 2 to 1.4 kg C.m-2yr-1 at age 6. Our data supported hypothesis 3: the decline in aboveground wood production (42% of peak) was proportionally greater than the decline in canopy photosynthesis (64% of peak). The fraction of GPP partitioned to belowground allocation and foliar respiration increased with stand age and contributed to the decline in aboveground wood production. The decline in GPP was not caused by nutrient limitation, a decline in leaf area or in photsynthetic capacity, or (from a related study on the same site) by hydraulic limitation. Nutrition did interact with the decline in GPP and aboveground wood production, because treatments with high nutritient availablity declined more slowly than did our control treatment, which was fertilized only during stand establishment.

Production and Resource Use Efficiencies in N- and P-Limited Tropical Forests: A Comparison of Responses to Long-term Fertilization

At two sites at the extreme ends of a soil development chronosequence in Hawaii, we investigated whether forest responses to fertilization on young soils were similar to those on highly weathered soils and whether the initial responses were maintained after 6–11 years of fertilization. Aboveground net primary production (ANPP) was increased by nitrogen (N) application at the 300-year-old site and phosphorus (P) application at the 4.1-million-year-old site, thus confirming earlier results and their designations as N- and P-limited forests. Along with ANPP, application of the limiting element consistently increased leaf area index (LAI), radiation conversion efficiency (RCE), and foliar and litter nutrient concentrations. Fertilization did not consistently alter N or P retranslocation from senescent leaves at either site, but a comparison with other sites on the chronosequence and with a common-garden study suggests that there is a genetic basis for low foliar and litter nutrients and higher retranslocation at infertile sites vs more fertile sites. N limitation appears to be expressed as limitation to carbon gain, with long leaf lifespans and high leaf mass per area. P limitation results in high P-use efficiency and disproportionally large increases in P uptake after fertilization; a comparison with other studies indicates large investments in acquiring and storing P. Although the general responses of ANPP, LAI, and RCE were similar for the two sites, other aspects of nutrient use differ in relation to the physiological and biogeochemical roles of the two elements.

Phosphorus limitation of forest leaf area and net primary production on a highly weathered soil

We tested the hypothesis that P was the nutrient limiting net primary production of a nativeMetrosideros polymorpha forest on a highly weathered montane tropical soil in Hawaii. A factorial experiment used all combinations of three fertilizer treatments: nitrogen (N), phosphorus (P) and a mix of other essential nutrients (OE), consisting primarily of mineral derived cations and excluding N and P. P addition, but not N or OE, increased leaf area index within 12 months, foliar P concentration measured at 18 months, and stem diameter increment within 18 months. Stem growth at 18 months was even greater when trees fertilized with P also received the OE treatment. N and P additions increased leaf litterfall and N and P in combination further increased litterfall. The sequence of responses suggests that increased available P promoted an increase in photosynthetic area which led to increased wood production. P was the essential element most limiting to primary production on old volcanic soil in contrast to the N limitation found on young volcanic soils.

Effects of chemical composition on nitrogen mineralization from green manures of seven tropical leguminous trees.

Green manures from seven tropical leguminous trees were incubated with soil to determine the rates and controls of net nitrogen release. Fresh green manure (leaves and succulent twigs) was mixed with moist soil and incubated in polyethylene bags. Net N mineralization from green manures was estimated by the accumulation of extractable ammonium and nitrate minus the accumulation in soil alone. Patterns of N mineralization were complex, differed among species, and at 12 weeks ranged from 10 to 65 percent of original green-manure N. Cumulative net N mineralization was negatively correlated with initial soluble polyphenol content in the early phases of decomposition (1 through 8 weeks) and with initial lignin content in later phases (4 through 12 weeks). Neither initial percent N nor lignin: N ratio were strongly correlated with N mineralization. The best chemical index of N release was the initial polyphenol: N ratio. This study confirms previous findings that N mineralization from tropical legumes is controlled more by soluble polyphenols than by lignin or N content.

Ecophysiology of exotic and native shrubs in Southern Wisconsin. II: Annual growth and carbon gain

SummaryIn this study we compared the aboveground growth rates of two exotic shrubs (Rhamnus cathartica and Lonicera X bella) and two native shrubs (Cornus racemosa and Prunus serotina) that are important in southern Wisconsin hardwood forests. For all species except P. serotina, aboveground growth rates in an open habitat were greater than in an understory environment. Growth rates differed among species in the open habitat and were significantly correlated with woody production per unit leaf area. All species had greater leaf area per unit wood biomass in the understory than in the open habitat. A comparison of above-ground growth and annual carbon gain suggests much greater respiratory costs in the open habitat, especially for P. serotina. The data from this study were used to examine mechanisms of species response to different light availabilities. We found that the species that increased their production per unit leaf area in response to increased light did not increase their leaf area per unit wood biomass in response to low light, and vice versa. Production of proportionately high leaf area may be important for the growth of C. racemosa in low light.

Within-System Element Cycles, Input-Output Budgets, and Nutrient Limitation

Widely used conceptual models for controls on nutrient cycling and input-outputs budgets of forest ecosystems suggest that: (1) nutrient losses from ecosystems originate in the available nutrient pool in soil; (2) nutrients that limit plant production are retained tightly within those systems; (3) this retention leads to accumulation of the limiting nutrient(s), eventually to the point at which it no longer limits production; and (4) losses of nutrient(s) thereafter should reflect rates of nutrient input, rather than biotic demand In this chapter, we explore mechanisms that could constrain the accumulation of a limiting nutrient, and therefore could allow nutrient limitation to persist indefinitely. Possible mechanisms include episodic disturbance-related nutrient losses, closed element cycles, and losses of nutrients from sources other than the available inorganic pool of nutrients in soil. For the last mechanism, both a simple and a more complex model are used to show that losses of dissolved organic forms of a nutrient could constrain nutrient accumulation and permit nutrient limitation to persist indefintely. Emissions of nitrogen (N) trace gases produced during nitrification could have a similar effect. To the extent that losses of nutrients by these and related pathways are important, anthropogenic inputs of nutrients (particularly N) could alter forest ecosystems substantially, to an extent greater than standard conceptual models would allow.

Forest productivity and efficiency of resource use across a chronosequence of tropical montane soils.

ABSTRACT We tested the hypothesis that plants adjust to nutrient availability by altering carbon allocation patterns and nutrient-use efficiency (NUE = net primary production [NPP] per unit nutrient uptake), but are constrained by a trade-off between NUE and light-use efficiency (ε= NPP per unit intercepted light). NPP, NUE and ε were measured in montane Metrosiderospolymorpha forest across a 4.1 x 106 yr space for time substitution chronosequence in which available soil N and P pools change with site age. Although the range of N and P availability across sites was broad, there was little difference in NPP between sites, and in contrast to theories of carbon allocation relative to limiting resources, we found no consistent relationships in production allocation to leaves, fine roots or wood. However, canopy nutrient pools and fluxes were correlated with the mass of fine roots per unit soil volume and there was a weak but positive correlation of NPP with LAI. Patterns of ε and NUE across the soil developmental sequence were opposite to each other. ε increased as nutrient availability and nutrient turnover increased, while NUE decreased in response to the same influences but reached its highest values where either N or P availability and turnover of both N and P were low. A negative correlation between ε and NUE supports the hypothesis that a trade-off exists between ε and leaf characteristics affecting NUE.