Simulation of nutrition of forests under stress using FORCYTE-11

Forest ecosystems are being subjected to an increasing variety of stresses for which we do not yet have rotation-length experience. Where we lack such experience, we cannot make experience-based predictions of the long-term effects of these stresses. While we are accumulating such experience, computer models can be used to make interim knowledge-based predictions. Most scientific knowledge has been produced by reductionist, disciplinary, process-based research. Such knowledge is a vital component of any explanation of natural or human-induced landscape phenomena, but cannot, in its reductionist, disciplinary form, provide an adequate basis on its own for long-term predictions about these phenomena. Such predictions require the development of computer models of ecosystem form and function based on the integration of knowledge from eco-physiology, autecology, community ecology, soil science, and climatology into ecosystem-level models that accurately describe the function and temporal dynamics of forest ecosystems. The ability of the ecosystem-level forest management simulation model FORCYTE-11 (FORest nutrientCycling andYieldTrendEvaluator) to simulate forest stresses is described briefly. The question of how to model other stresses, such as air pollution, acid rain, climate change, soil compaction and erosion, and moisture competition is discussed, and the outline of a new model (FORECAST:FORestry andEnvironmentalChangeASsessmenT) is presented.     

Increase in the abundance of cladocera at pH 5.1 in experimentally-acidified lake 223, Experimental Lakes Area,Ontario

Lake 223 in the Experimental Lakes Area, northwestern Ontario was experimentally-acidified with H₂SO₄ from 1976 to the present. Cladocerans increased in absolute and relative abundance and in total biomass when the pH of Lake 223 was lowered from 6.7 to 5.1. Comparing the cladoceran community at pH 6.7 (1974) with that at pH 5.1 (1981 to 1983), Daphnia galeata mendotae disappeared, Diaphanosoma birgei became rare, Holopedium gibberum increased in abundance and Daphnia catawba, recorded for the first time in 1980, became abundant. Bosmina longirostris became more abundant. Except for the decline in D. birgei, the changes in cladoceran species composition with acidification are consistent with known responses of these species to low pH. The increase in importance of cladocerans in 1981 to 1983 was at the expense of copepods. It was hypothesized to be caused by the decline in predation by small fish during 1981 to 1983 as acidification reduced minnow populations and inhibited reproduction in trout and sucker. The loss of the population of the invertebrate predator, the opposum shrimp, Mysis relicta, in 1979 had little effect on cladoceran biomass.     

Fish species distribution in relation to lake acidity in Ontario

Survey data from approximately 2,900 Ontario lakes were used to examine the distribution of 12 fish species with respect to lake pH. Yellow perch (Perca flavescens), white sucker (Catostomus commersoni), brook trout (Salvelinus fontinalis), pumpkinseed (Lepomis gibbosus), fathead minnow (Pimephales promelas) and redbelly dace (Phoxinus eos) proved to be tolerant of low pH (pH <6.4). Lake trout (Salvelinus namaycush), common shiner (Notropis cornutus), blacknose shiner (Notropis heterolepis), lake whitefish (Coregonus clupeaformis), walleye (Stizostedion vitreum vitreum) and northern pike (Esox lucius) showed limited distribution in low pH lakes (pH <6.4). The limited distribution of lake whitefish and northern pike likely relates to zoogeographic factors rather than their intolerance to low pH. Lake trout and common shiner occur in lakes susceptible to, and receiving high loadings of acidic deposition. However, without historical data, we cannot determine if their decreased distribution in low pH systems is due to intolerance to low pH.     

Mercury in plants,soil and atmosphere near a chlor-alkali complex

Data of atmospheric Hg concentrations measured near the chlor-alkali complex of Rosignano Solvay (Livorno, Italy) show that the impact of the industry on the terrestrial environment is restricted to a close area around; background values (3 to 5 ng m^?3) are reached within a radius of 4 to 5 km. Hg levels in plants (Poa sp. Pinus nigra and Solidago sp.) and in soil show the highest values in the sampling stations near the electrolytic cells.     

Population dynamics of yellow perch (Perca flavescens) and pumpkinseed (Lepomis gibbosus) in two acid-stressed lakes

Fish population parameters: abundance, age-class composition, and size-at-age, which respond to environmental stresses, were investigated in two lakes showing chemical and biological evidence of acidification. Plastic Lake, Muskoka District of Ontario, shows spring pH depression and A1 toxicity in surface waters, with an associated annual fish kill. In Crosson Lake, pH 5.2 to 5.3, there has been a dramatic decline in abundance of the white sucker (Catostomus commersoni). In Platic Lake the abundance of pumpkinseed (Lepomis gibbosus) has declined and that of rock bass (Ambloplites rupestris) has increased. In Crosson Lake size-at-age has increased in yellow perch (Perca flavescens), pumpkinseed abundance has increased but with no change in growth rate.     

Role of air pollution in forest decline in Eastern North America

Evidence for and against three general mechanisms by which air pollution stress may cause or contribute to forest decline in eastern North America is examined. These mechanisms are (1) soil acidification, cation nutrient depletion, and subsequent Al toxicity, (2) direct and indirect effects of gaseous pollutants on the physiology and growth of forest trees; and (3) excess nitrogen deposition and subsequent soil acidification or physiological injury. Recent studies have revealed reductions in base saturation in some sites (both polluted and pristine), but the consequences of these changes to forest health and nutrition are not resolved. Aluminum toxicity may contribute to forest decline in red spruce (Picearubens L.) in high-elevation sites, but the data are contradictory. In forested landscapes in which regional decline (reduced tree growth and/or dieback) is reported, O₃ is the only gaseous pollutant documented as a contributing factor in the eastern North America. Whereas this secondary pollutant exhibits a regional distribution and occurs at potentially phytotoxic levels, a linkage between O₃ and forest decline is substantive only for loblolly pine (Pinustaeda Sarg.) in the southeast. Studies in high elevations do not support the hypothesis that O₃ or hydrogen peroxide directly affects the growth of red spruce or fraser fir (Abiesfraseri Poir.).However, preliminary data link chronic-level O₃ exposure with indirect effects, principally changes in drought tolerance in low-elevations (e.g., P. taeda) nd winter hardiness in high elevations (e.g., P. rubens).The emerging data on the role of air pollution as an environmental stress indicate that indirect effects (i.e., responses in which pollution alters the plant's ability to compete for limited resources or withstand other environmental stresses) are more important than direct effects (e.g., foliar necrotic lesions). Nitrogen saturation has been offered as an hypothesis for P. rubens decline, but no experimental data exist supporting this hypothesis.     

Comparison of the macrophyte communities of a clearwater and a brownwater oligotrophic lake in Kejimkujik National Park,Nova Scotia

Beaverskin and Pebbleloggitch Lakes are small, proximate, acidic, oligotrophic, headwater lakes located in southwestern Nova Scotia. Beaverskin Lake has clear water, its euphotic zone is deep, and it has extensive beds of macrophytic vegetation that cover most of its bottom to a depth of 6.5 m. In contrast, the water of Pebbleloggitch Lake is highly colored by dissolved humic substances, and macrophytes are restricted to a narrow littoral fringe in depths of less than about 1.8 m. The most widespread macrophyte community in Beaverskin Lake occurs at depths of 2.0–6.5 m and is dominated by Sphagnum macrophyllum and Utricularia vulgaris, while the second most prominent community is littoral and is dominated by Eriocaulon septangulare, Lobelia dortmanna, and Eleocharis acicularis. The most prominent community in Pebbleloggitch Lake is dominated by the floating-leaved Nuphar variegatum rooted as deep as 1.8 m, while communities dominated by E. septangulare and Sphagnum spp. occur in water less than about 0.7 m deep.     

The influence of mycorrhizal associations on paper birch and jack pine seedlings when exposed to elevated copper,nickel or aluminum

Acid deposition may adversely affect northern forest ecosystems by increasing the concentration of metals in the soil solution. This study investigates the effects of ectomycorrhizal fungi on paper birch and jack pine seedlings exposed to elevated Cu, Ni, or Al in sand culture. One of four mycorrhizal fungi,Scleroderma flavidum, was able to reduce Ni toxicity to the birch seedlings. It did this by reducing transport of Ni to the stems. None of the fungi affected Cu toxicity in birch. In separate experiments, jack pine seedlings were exposed to combinations of Al and Ca. Infection withRhizopogon rubescens increased seedling susceptibility to A1. Seedlings inoculated withSuillus tomentosus showed a greater growth stimulation by Ca than uninoculated jack pines. Thus, for both tree species, the mycorrhizal association could alter the response of seedlings to high concentrations of certain metals, although this varied with fungal species.     

Fertilizer application in aid of plantation establishment in the savanna areas of Nigeria

Fertilizer trials were conducted to stimulate tree establishment and growth in the savanna region of Nigeria. The main nutrients investigated were N, P, K and B while the tree species included Azadirachta indica (Neem), Acacia auriculiformis, Eucalyptus spp., and Pinus spp. Borate application reduced the incidence of die-back of eucalypts and it also increased height growth. There was positive response to phosphate application by all species; phosphate reduced mortality considerably in pines on some sites. Response to N fertilizer by the species varied and related to type of N fertilizer. Urea was often found injurious to pines. There was strong interaction between N and P and usually response to N was not obtained on P-deficient sites. In all the cases where it was applied, there was no response to K. On more arid sites with poorly buffered soils, mineral fertilizers alone were insufficient to produce satisfactory tree growth. Application of animal manure on such sites increased survival and growth of Azadirachta indica, E. camaldulensis and A. auriculiformis.     

Biological indicators of lake acidification

Indicator taxa are identified, based on both synoptic surveys and whole lake acidification experiments, for lake acidification in the pH 6.0 to 5.0 range. Acidobiontic diatoms (e.g Asterionella ralfsii, Fragillaria acidobiontica, etc.), periphyton (Mougeotia and related species), macroinvertebrates (e.g. Hyalella azteca, Orconectes sp., etc.), leeches, and cyprinid fishes (e.g. Pimephales promelas, Notropis cornutus, etc.) are identified as target organisms during early phases of lake acidification.     

The effect of simulated acid rain on feather mosses and lichens of the boreal forest

Permanent field plots containing a dominant ground cover of feather moss (Pleurozium schreberi) and the forage lichen, (Cladina), were established in mature, boreal forest jack pine stands to monitor the effects of simulated acid precipitation. For a five-year period commencing in 1981, bimonthly sprays (pH range 2.5 to 5.6) were given throughout the growing season. The feather moss wefts were extremely sensitive to simulated rains of pH 2.5 and 3.0; but loss of cover and frond blackening were also observed at pH 3.5. The pH 2.5 treatment killed almost all of thePleurozium, while the cover remaining in the pH 3.0 treatment after 5 years was reduced by 44%. In laboratory studies designed to compare the effects of H₂S0₄, HN03 and a 2:1 mixture of both, microcosms sprayed with H₂S0₄ alone (pH 3.0) were more significantly affected than fronds treated with HN0₃ alone or pH 5.6 sprays of any ratio. Although less sensitive thanPleurozium, field-sprayed lichens were also visibly damaged. At pHs less than 3.5,C. stellaris andC. rangiferina had reduced podetial height and dry weight; whileC. mitis was affected by a combination of the acid rain treatment and other associated factors. While ambient rains of pH 4.2 may not in themselves be harmful to the boreal ground flora, it is apparent that the feather mosses and lichens, lacking a cuticle and true roots, are very sensitive to occasional, extremely acidic rain events.     

Foliar sensitivity of eight eastern hardwood tree species to ozone

As part of a larger 3-yr study, container-grown seedlings of black cherry (Prunus serotina) red maple (Acer rubrum), red oak (Ouercus rubra), sweetgum (Liquidambar styraciflua), white ash (Fraxinus americana), white oak (Ouercus alba), yellow-poplar (Liriodendron tulipifera), and yellow birch (Betula allegheniensis) were exposed to 0,0.075, or 0.15 μL L-¹ O₃ in laboratory controlled-environment chambers for 6 hr d^?1 on 2 consecutive days for 12 weeks. On the third consecutive day of each week, plants were treated for 45 min with precipitation at pH 3.0 or 4.2. The only significant foliar symptoms were induced by the O₃ treatments, and the severity of symptoms was not influenced by precipitation pH. The most common symptom was a dark, adaxial stipple which was most severe on the oldest leaves. Equations were developed to express the influence of leaf position on percent leaf injury following 4, 8, and 12 weeks of treatment. Based on percent leaf tissue showing stipple and defoliation following exposure to 0.15 μL L^?1 O₃, the most sensitive species to O₃ was black cherry, followed by sweetgum, yellow-poplar, white ash, red maple and yellow birch. Red oak and white oak foliage did not exhibit stipple.     

The carbon cycle and global forest ecosystem

Attempts to account for the fluxes by quantifying C sources and sinks have provided evidence of a missing C sink (Detwiler and Hall, 1988), which may be located somewhere in the temperate region of the northern hemisphere (Tanset al., 1990). Until recently, most estimates have concluded that the temperate forest is a small C source. Two recent papers (Sedjo, 1992; Kauppiet al., 1992) provided evidence that the temperate forests are substantial C sinks. This paper combines these earlier findings on temeperate forest carbon sequestration with a new estimate of the annual C releases due to tropical deforestation, 1.7 Gt, which is obtained using the FAO estimates of the rate of deforestation in the tropics over the decade of the 1980s and conservative estimates of C releases associated with this deforestation. Finally, to this is added the crude estimate of C export by the global river system found in Hallet al. (1992). Applying these estimates of the C sink function of both temperate and tropical forests to Detwiler and Hall's alternative C budgets largely eliminates the “missing C” hypothesized by Detwiler and Hall, and Tanset al.     

Neutralization of acidic raindrops on leaves of agricultural crop and boreal forest species

The abilities of foliage of selected agricultural crop and native boreal forest species to neutralize acidic raindrops were compared. The species differed widely in their responses. Neutralization was influenced to a large extent by leaf wettability and was poorly related with species' susceptibility to foliar injury from acid rain sprayings. Little neutralization of pH 3.0 droplets occurred on very waxy leaves, e.g. cabbage (Brassica oleracea L.), due to the small contact area between the leaf surface and raindrops. In contrast, on sunflower (Helianthus annuus L.) and radish (Raphanus sativus L.) leaves, which are pubescent and easily wettable, neutralization was considerable. For all agricultural crop species examined, the pH of droplets drying on cotyledons was consistently higher than on the leaves. The pH values of raindrops were also higher when the foliage was injured by the acid rain, probably due to leakage of cellular contents. Among boreal forest species examined, bunchberry (Cornus canadensis L.) was particularly good at neutralizing natural acid rain, increasing the pH from 3.9 to 6.6 after 9 hr of foliar contact, while the response of other boreal species ranged from a final pH of 4.8 to 5.7 under the same conditions. Simulated raindrops on wild sarsaparilla (Aralia nudicaulis L.) were never neutralized but increased in acidity as they evaporated. Chemical analyses of droplets collected from foliage showed calcium (Ca) and potassium (K) to he the major cations entering the neutralized droplets. Neutralization of acidic raindrops appears to occur through two processes: solubilization of alkaline dusts and exudates on the leaf surface, and ion exchange removal of H⁺ by the foliage.     

Fertilizer application in aid of plantation establishment in the savanna areas of Nigeria

Fertilizer trials were conducted to stimulate tree establishment and growth in the savanna region of Nigeria. The main nutrients investigated were N, P, K and B while the tree species includedAzadirachta indica (Neem),Acacia auriculiformis, Eucalyptus spp., andPinus spp. Borate application reduced the incidence of die-back of eucalypts and it also increased height growth. There was positive response to phosphate application by all species; phosphate reduced mortality considerably in pines on some sites. Response to N fertilizer by the species varied and related to type of N fertilizer. Urea was often found injurious to pines. There was strong interaction between N and P and usually response to N was not obtained on P-deficient sites. In all the cases where it was applied, there was no response to K. On more arid sites with poorly buffered soils, mineral fertilizers alone were insufficient to produce satisfactory tree growth. Application of animal manure on such sites increased survival and growth ofAzadirachta indica, E. camaldulensis andA. auriculiformis.     

The impact of acidification on aquatic biota in North America: A comparison of field and laboratory results

Acidification of lakes and streams can lead to adverse effects on aquatic biota, and the recognition of widespread atmospheric deposition of strong acids in eastern North America has led to considerable concern over its existing and potential impact on th flora and fauna of sensitive (i.e. low alkalinity) systems. While most of the concern has been directed toward the impact on the fisheries resource, it is well documented that all levels of biota are affected (Almer et al., 1974; Likens et al., 1983; Dillon et al., 1984), and full appreciation of the impact of acidification requires examination of all living components of the aquatic ecosystem. Although extensive work has already been conducted in tho Scandinavian countries, there are numerous differences in species between those countries and North America, and a sound appraisal of the threat to North American biota requires comparable work for this continent. Large scale surveys and monitoring programs are under development or are in the early phases of execution by various institutions, but much has already been learned from laboratory and field acidification experiments, and from comparisons of acidic and non-acidic lakes and streams. The purpose of this paper is to compare the results of these studies for several dominant species from several major groups of organisms. We hope to focus attention on the nature and quality of information about the sensitivity of organisms with known or apparent significant ecological roles, identify gaps or weaknesses, and provide insight into the merits and limitations of the various methods of study.     

Fish species distribution in relation to water chemistry in selected Maine lakes

We examined the possible effects of acidity on the number of fish species in 22 lakes in Maine, ranging in pH from 4.4 to 7.0 (mean values). We caught no fish in three lakes with pH <5.0, but collected 1 to 9 species in the remaining 19 lakes (pH 5.4 to 7.0). Brook trout (Salvelinus fontinalis), golden shiners (Notemigonus crysoleucas), and white suckers (Catostomus commersoni) were ubiquitous, but no common shiners (Notropis cornutus) or creek chubs (Semotilus atromaculatus) were collected from lakes with pH below 6.0 and 5.9, respectively. The fishless lakes differed from the others primarily in water chemistry variables related to acidity, i.e., pH, Al, and concentration of divalent cations. Among lakes that contained fish, the factors related to the number of species collected were lake surface area and maximum depth, which may be related to habitat quantity and diversity. Cluster analysis identified two distinct fish species groups — depauperate and cyprinid-sucker — but multiple comparison analysis failed to relate any measured chemical or physical variable to these two groups, probably because water chemistry was suitable for reproduction by these species.     

Natural changes in the planktonic rotifera of a small acid lake near Sudbury,Ontario following water quality improvements

Swan Lake is a small, acidic, metal-contaminated lake located near Sudbury, Ontario. During the past 8 yr, the lake has experienced a substantial increase in pH, together with significant reductions in the concentrations of heavy metals and base cations. These changes were observed only after acid and metal emissions from Sudbury area smelters were reduced. The composition of the planktonic Rotifera in the lake has changed concurrent with improvements in water chemistry. Dominance of the rotifer community by the acidophile Keratella taurocephala has been sharply reduced in recent years, while significant increases in the densities of Polyarthra spp., Chromogaster ovalis, Conochiloides natans and Trichocerca similis have been observed.     

Patterns of plankton species,pH and associated water chemistry in Nova Scotia lakes

In order to assess the patterns of distribution of plankton species in relation to acidity, water chemistry, and physical characteristics, twenty lakes in Nova Scotia were selected for examination, with pH ranging from 3.5 to 7.6. Correlation and multiple regression analyses revealed associations between plankton and chemical variables. Patterns of plankton abundance and diversity were noted with respect to both pH and nutrient status, with occasionally conflicting effects of these two factors. Phytoplankton abundance was most closely associated with nutrient status, while zooplankton abundance was related to measures of primary production. pH tended to affect plankton diversity more than abundance or standing crop. Several lakes were outliers from overall patterns, with several possible explanations. Cluster and factor analyses were used to identify associations of plankton species, and to relate these associations to lake characteristics. Certain species were noted with relatively wide distribution, but which never occurred at low pH (<4.6). These were Chroococcus limneticus, Asterionella formosa, Cyclotella meneghiniana, Pinnularia braunii, and Surirella robusta for phytoplankton, and Polyphemus pediculus, Diaptomus oregonensis, and Tropocyclops prasinus for zooplankton.