Aluminum,Fe, Ca,Mg, K,Mn, Cu,Zn and P in above- and belowground biomass. II. Pools and circulation in a subalpineAbies amabilis stand

Elemental concentrations of above- and belowground tissues were determined in anAbies amabilis stand in the Cascade Mountains, Washington, USA. These data were used to calculate the pools and circulation of trace elements and micronutrients on a stand level. For all elements except Al, a greater proportion (from 62 to 87%) was distributed in above- rather than belowground tissues. This contrasted sharply with the biocirculation of elements where 97% of the Al and Fe, 88% of the Cu and 67–84% of the Ca, P, and Mg of total detrital cycling was from the belowground components. Aboveground tissues, however, contributed 69% of the Zn, 65% of the K and 68% of the Mn found in annual detritus production. The proportion of total element pool circulated annually was the highest for Al (82%) and Fe (32%) followed by 13% and less for the remaining elements. Copper, Fe and Al were accumulated in root tissues, while Mn and Zn accumulated in foliage.We hypothesize that roots are an effective mechanism for avoiding Al toxicity in these subalpine ecosystems. The large root biomasses of these stands allow for high Al levels to be taken up and immobilized in roots; this is observed in the significantly higher Al accumulations in below- than aboveground tissues. The high root turnover in these stands is hypothesized to be a result of root senescence occurring in response to high Al accumulation. Furthermore, Al inputs into detritus production occur by soil horizon so that roots with high Al concentrations located in the Bhs horizon turnover and are retained within that horizon. These roots also decompose very slowly (99% decay = 456 years) due to the high Al and low Ca, Mn and Mg present in these tissues and therefore have very little impact on short-term elemental cycling.     

Structure and dynamics of an ancient montane forest in coastal British Columbia

Old-growth forests are common in the snowy, montane environments of coastal western North America. To examine dynamics of a stand containing four canopy tree species (Abies amabilis, Chamaecyparis nootkatensis, Tsuga mertensiana and T. heterophylla), we used four stem-mapped, 50 m ×50 m plots. From measurements of annual rings, we obtained ages from basal discs of 1,336 live trees, developed master chronologies for each species, reconstructed early growth rates, and delineated periods of release. The stand was ancient; individuals of all four species exceeded 900 years in age, and the oldest tree exceeded 1,400 years. The four plots differed in the timing of events, and we found no evidence of major, stand-level disturbance. Instead the stand was structured by small-scale patch dynamics, resulting from events that affected one to several trees and initiated episodes of release and relatively rapid early growth. The species differed in age structure and dynamics. A. amabilis and T. heterophylla had a classical reverse-J age structure indicative of stable populations, whereas C. nootkatensis and T. mertensiana appeared to rely on local episodes of increased recruitment, which were often separated by centuries, and were probably related to multiple-tree gaps that occurred infrequently. However, such gaps could be considered normal in the long-term history of the stand, and thus these species with their long life spans can persist. Most individuals of all four species grew extremely slowly, with trees typically spending centuries in the understory before reaching the canopy, where they were able to persist for additional centuries. Thus, the key features of this forest are the very slow dynamics dominated by small-scale events, and the slow growth of stress-tolerant trees.     

The influence of soil chemistry on fine root aluminum concentrations and root dynamics in a subalpine Spodosol,Washington State,USA

The relationship between root Al concentrations and Al fractions in the soil solution was examined in a mature Abies amabilis ecosystem in the Cascade Range of Washington State. The naturally acidic soils in these ecosystems lead to high concentrations of aqueous Al in soil solutions and contribute to the biocycling of Al by the A. amabilis/T. mertensiana stand. Root concentrations of Al were very closely related to aqueous Al³⁺ activities, but poorly correlated with total aqueous Al concentrations. The solution Al/Ca molar ratios followed a seasonal cycle with low values during the fall and high values during the spring. Ratios remained <1 throughout the year in the Oa horizon while they varied between 2 and 14 in the E and Bhs horizons. The vertical distribution of roots and the mortality of fine roots may be linked to the soil solution Al/Ca ratio. Root cation exchange capacity ranged between 180 and 225 mol g^-1 and the exchangeable Al fraction represented from 12–17% of the total Al content in the root. Evidence for solid-phase co-precipitates of Al with PO₄ and oxalate was indicated from selective dissolution of the root tissue. Sufficient quantities of PO₄ and oxalate exist in the roots to tie up 20–40% of the Al present in the roots of the Oa and E horizons, but only 9% of that present in the Bhs horizon. Species differences in the distribution of Al between the above-ground and below-ground components may be dictated by these retention processes in the fine roots.     

Anomalous secondary vascular tissue inHelminthostachys zeylanica (Ophioglossaceae)

The vascular anatomy ofHelminthostachys zeylanica was examined with special reference to anomalous secondary tissue. Primary xylem development gradually takes place centrifugally. In branched rhizomes with destroyed apices, the vascular cylinder apical to the insertion of branch traces is generally composed of primary xylem, accessory xylem, inner parenchyma of radially arranged cells, outer parenchyma of irregularly arranged cells, and partly crushed phloem, listed in order going outwards. The accessory xylem as well as the inner parenchyma ofHelminthostachys zeylanica is probably secondarily produced, partly to contribute to the branch traces, in a position corresponding to that of secondary vascular tissue developed from a normal cambium inBotrychium sensu lato. It is suggested that although a cambium is lacking inHelminthostachys zeylanica, the secondary vascular tissues are comparable between the genera. The phylogenetic implication of this tissue is discussed.     

Vesicular endophytes in roots of the Pinaceae

Vesicles and hyphae typical of vesicular-arbuscular mycorrhizae (VAM) were common in seedlings of Pseudotsuga menziesii, Abies lasiocarpa and Tsuga mertensiana growing in openings where herbaceous hosts of these fungi were common. Seedlings of A. lasiocarpa, Tsuga heterophylla, and T. mertensiana growing under closed forest canopies also had vesicles but at a much lower incidence than seedlings in the openings. The Pinaceae are generally assumed to be ectomycorrhizal, but Glomus-type colonizations occurred on the same seedlings as the ectomycorrhizae. The ecological significance of abundant VAM-type endophytes in otherwise ectomycorrhizal hosts deserves comprehensive study.     

Effects of N addition rates on the productivity ofPicea Sitchensis,Thuja plicata,andTsuga heterophylla seedlings

Seedlings ofPicea sitchensis, Thuja plicata andTsuga heterophylla were supplied N hydroponically at one of four exponentially increasing rates of addition (0.09, 0.07, 0.05, or 0.025 gN^-1 day^-1) for up to 3 months in a naturally illuminated glasshouse. Relative growth rates (RGR) were analyzed as a function of N uptake, the allocation of assimilated N to foliage (LNFR), foliar N concentrations (N_la) and met assimilation rates (NAR), which were combined to estimate N productivity (RGR per unit whole-plant N concentration). Nitrogen accumulation, biomass and N partitioning and RGR and its components varied with species in response to the different N regimes.T. heterophylla had the lowest maximum wholeplant N concentrations (wpN) and specific absorption rates for N and exhibited the least plasticity in root: shoot ratios as wpN increased from 11–21 mg g^-1. In all species, RGR increased linearly with wpN, while LNFR increased curvilinearly. Foliar N (N_la) increased linearly with wpN and NAR increased linearly with N_la. The RGRs ofT. heterophylla were highest at wpNs up to 18 mg g^-1, a result of higher foliar N use efficiencies (NAR/N_la). However, RGR increased more with wpN inT. plicata andP. sitchensis. Although LNFR increased with wpN in all species, foliar N use efficiency declined, possibly due to an increased partitioning of foliar soluble N to non-photosynthetic compounds. Thus, in each species, N productivity did not increase above intermediate levels of wpN: 14 mg g^-1 inT. heterophylla, 16 mg g^-1 inP. sitchensis and 17 mg g^-1 inT. plicata.     

The effect of aluminium on the distribution of calcium,magnesium and phosphorus in mycorrhizal and non-mycorrhizal seedlings of Eucalyptus rudis: a cryo-microanalytical study

The distribution of Al, Ca, Mg and P in the lateral roots and leaves of mycorrhizal and non-mycorrhizal seedlings of Eucalyptus rudis grown with and without Al was analysed using energy-dispersive X-ray microanalysis on a cryo-scanning electron microscope. Al accumulated in all tissues of nonmycorrhizal plants: the endodermis was not a barrier to the translocation of Al. In mycorrhizal roots, Al was concentrated within the sheath. The presence of Al reduced the levels of Ca and Mg in both mycorrhizal and non-mycorrhizal roots and shoots in comparison with control plants. The presence of mycorrhizas increased the levels of Ca and Mg in plants grown with Al in comparison with non-inoculated plants, although there was no evidence that mycorrhizas increased the levels of P in plants grown in Al-amended soils. P levels were higher in the mycorrhizal sheath of plants grown with Al than the controls.     

Tsuga shimokawaensis,a new species of permineralized conifer leaves from the Middle Miocene Shimokawa Group,Hokkaido, Japan

Silicified pinaceous leaves from the late Middle Miocene bed of Shimokawa Town, central Hokkaido, Japan are described asTsuga shimokawaensis sp. nov. Comparisons with leaves of extant species ofTsuga show that the new species is assigned to sect.Tsuga, resembling the extant speciesT. heterophylla in having usually one-cell-layered hypodermis, andT. sieboldii in having mesophyll cells between the resin canal and the hypodermis. From cladistic analyses we infer thatTsuga shimokawaensis is positioned the basal in sect.Tsuga and that sect.Tsuga comprises four species groups.     

Differential foliar responses of northern hardwoods to fertilization

A 70-year-old thinned northeastern Fagus-Betula-Acer stand in the Adirondack Mountains of northern New York was fertilized with varying combinations of N, P, K, and lime in the spring of 1976.Betula alleghaniensis Brit.,Acer saccharum Marsh.,Acer rubrum L., andFagus grandifolia Ehrh. foliage was collected in the autumn for 1974 through 1977 and analyzed for foliage areas and weights, and levels of ash, N, P, K, Ca, Mg, Mn, Na, Fe, Zn, Al, Cu, and Co. Comparisons are made within species and among treatments, expressed as concentrations on a dry weight basis. Elemental composition is examined to determine the differential foliar responses to fertilization.Contribution of State University of New York College of Environmental Science and Forestry, Syracuse, New York, 13210.The authors are Graduate Research Assistant, Director of Huntington Forest, Technical Research Assistant, and Professor of Forest Soil Science (now deceased), SUNY, respectively.     

Elemental concentrations in plant tissues as influenced by low pH soils

Summary Soils influenced by acid mine drainage (pH<5.0) are characterized by low concentrations of essential nutrients and increased solubility of heavy metals. The conditions typically reduce plant establishment and growth. However, river birch (Betula nigra L.) is commonly found along low pH streams in southeastern Ohio. The objective of this study was to determine the concentration of Al, Mn, Ca and Mg inB. nigra tissues.The results indicate Al and Mn are accumulating inB. nigra when compared to other species. Within river birch, Al concentrations are highest in roots; Mn concentrations are highest in leaves. There is not a concomitant reduction in Ca and Mg concentrations as suggested by soil levels.     

The mountain forests of british columbia and the american northwest: Floristic patterns and syntaxonomy

An overview of the montane and subalpine forests ofAbies lasiocarpa, Picea engelmannii, Abies amabilis andTsuga mertensiana of northwestern North America is provided based on 541 relevés of forest vegetation from coastal and inland British Columbia, Washington and northwestern Montana. A new classification is proposed based on an examination of the floristic patterns across these communities. Owing to their broadscale floristic similarities, coastal subalpineAbies amabilis-Tsuga mertensiana forests and inland forests ofAbies lasiocarpa, Picea engelmannii, P. glauca, Thuja plicata andTsuga heterophylla are combined into a single order,Thujetalia plicatae. The floristic patterns within this order in turn support the recognition of four alliances. These are (1)Tsugion mertensianae, including coastal and inland subalpine forests ofTsuga mertensiana, Abies amabilis, A. lasiocarpa andPicea engelmannii, with deep winter snowpacks and an often luxuriously developed shrub layer; (2)Abieti-Tsugion heterophyllae all. nov., including mesic inland montaneThuja plicata-Tsuga heterophylla andAbies lasiocarpa-Picea engelmannii forests and related southern borealAbies lasiocarpa-Picea glauca communities; (3)Gymnocarpio-Abietion lasiocarpae, including moist coastal and inland forests dominated by a fern-rich understory andOplopanax horridus; and (4)Lysichito-Chamaecyparidion nootkatensis, which includes wet coastal subalpine forests withLysichiton americanum. A total of five new associations are described, bringing the number of recognized associations within the order to twenty-five. An extensive synonymy and crosswalk to other classification systems is provided.     

Late quaternary dynamics of pollen influx at mineral lake,Washington

Pollen influx analysis at Mineral Lake, Washington, indicates that immediately south of the Puget Lobe of the Fraser Glaciation, tundra was a characteristic vegetation until 16,300 years ago. Invasion ofPinus contorta began 17,500 years B.P., and boreal climax conifers (Abies, Picea andTsuga mertensiana), 16,300, but was temporarily interrupted by the Vashon advance (14,500–14,000 yr B.P.).Pseudotsuga menziesii began to grow in population 10,750 years ago, and woodland was established within a time span of 1,000 years. Modern lowland coniferous forests began to form 7,000 years ago. Logistic analysis of pollen abundance changes show that the intrinsic growth rate,r (yr⁻¹), of pioneer species (e.g. 0.024–0.026 inPteridium aquilinum) is higher than that of climax species (e.g. 0.003 inThuja plicata).P. menziesii, a subclimax species, shows an intermediater value (0.013) between these two ecologically different taxa. The absoluter value ofP. contorta (−0.011) is only slightly lower than that ofP. menziesii, although their replacement began almost simultaneously. Thus competition between these species was intense before the inflection point ofPinus curve 10,100 years ago. At this time, forest gaps became abundantly available forPseudotsuga, as indicated by a peak of the diagnostic factor (the reciprocal of the pollen influx).     

The dynamics of old-growth Pseudotsuga forests in the western Cascade Range,Oregon, USA

Size and age structure, spatial analysis, and disturbance history were used to analyse the population structures and regeneration patterns of 8 conifer stands in the central western Cascade Range, Oregon, USA. Variation in forest structure reflected the effects of frequent (20–50 yr) low-intensity fires and treefalls, infrequent (100–200 yr) localised, intense fires, and extensive fires that resulted in stand replacement (every ca 400 yr?).The amount of canopy removed and the size of openings formed by fires and treefalls were important determinants of subsequent forest establishment. Single or several species stands of Pseudotsuga and/or Abies procera, or mixed species stands of Pseudotsuga, Abies procera, Tsuga heterophylla and Abies amabilis established in openings where intense fires had removed most of the canopy trees over several ha. Multi-tiered and multi-aged stands, often containing 400–500 yr-old Pseudotsuga and variously-sized more or less even-aged patches of younger shade tolerant Tsuga heterophylla and/or Abies amabilis, occurred where lower-intensity fires did not kill all overstorey trees or where treefalls occurred after the initial fire.Current regeneration processes are influenced by overstorey composition, the availability and size of canopy openings, and the availability of substrates suitable for regeneration. Tsuga heterophylla and Abies amabilis established under Pseudotsuga menziesii and Abies procera canopies and in small canopy openings (<400 m²) created by windfalls, but rarely under Tsuga. Down logs and stumps were favoured establishment sites for Tsuga.The disturbance regime of fires of low-to moderate-intensity, windfalls, and occasional fires that result in extensive stand replacement contrasts with the pattern of infrequent, catastrophic disturbances proposed for other areas of the Pacific Northwest. Although fires at stand establishment commonly determine much of the composition, structure, and subsequent stand development, canopy replacement by shade tolerant species occurs as the different life histories of the species are expressed in response to various disturbances differing in intensity and frequency. Such a non-equilibrium view of vegetation change is consistent with many other fire-dominated forests of the western United States.     

Identity and frequency of occurrence ofTrichoderma spp. in roots of wheat and rye-grass in Western Australia and their effect on root rot caused byGaeumannomyces graminis var.tritici

Trichoderma hamatum, T. harzianum andT. koningii were isolated from wheat and rye-grass roots from a field in Western Australia. Frequency of occurrence ofTrichoderma spp. was higher on roots subjected to washing only, for both wheat and rye-grass than the roots which were surface-sterilized with 0.6% or 1.25% NaOCl.Trichoderma spp. were recovered at a higher frequency on PDA amended with lactic acid (pH 4.5) than on PDA alone (pH 5.6) or PDA with streptomycin. In general,Trichoderma spp. were isolated at a higher frequency from roots of wheat than that of rye-grass.T. hamatum occurred at a higher frequency in rye-grass roots than in wheat, whereasT. harzianum was more common in roots of wheat than in rye-grass, especially in seedling and milky ripe stages.T. koningii was recovered at a higher frequency from roots at seedling stage of rye-grass than wheat, the reverse being true at tillering stage.T. koningii was not recovered from roots of either host in any sampling when they were surface sterilized with 1.25% NaOCl.The take-all fungus was isolated from wheat and rye-grass roots more frequently at tillering and stem extension stages than others. It was severely pathogenic to both hosts in sterilized and non-sterilized soil.Addition of lactic acid, HCl or streptomycin to PDA did not affect the growth of theTrichoderma spp. tested, but the growth was slower on Martin's medium than on other media. In generalT. harzianum andT. koningii grow faster thanT. hamatum. The growth of the three species were not different at 20 and 25°C, but at 15°c growing of all species was significantly reduced.Incorporation of lactic acid into PDA prevented the bacterial growth in all treatments. Streptomycin too reduced but to a lesser degree than lactic acid. Surface sterilization with NaOCl decreased the recovery of both bacteria and fungi. T. hamatum andT. koningii reduced the mortality of wheat and rye-grass plants inoculated with the take-all fungus in sterilized and non-sterilized soil, whereT. harzianum did not protect wheat or rye-grass from infection by the take-all fungus.     

Effect of metabolites ofAspergillus niger andTrichoderma viride on development and structure of radicle of cocoa (Theobroma cacao) seedlings

Both the volume and concentration of filtrate ofAspergillus niger andTrichoderma viride influenced the development of the radicles of cocoa seedlings. Radicles placed in Petri dishes containing filter paper moistened with 20 and 30 ml of undiluted filtrate and filtrate dilutions of 1∶1 and 1∶2 failed to develop lateral roots and eventually died. The culture filtrate ofA. niger was more repressive. Radicles in the same volume of filtrate (20 and 30 ml) of higher dilutions (1∶5 and 1∶10) developed lateral roots and survived. Radicles placed in less volume (≤10 ml) survived and produced lateral roots irrespective of concentration of filtrates. Development of the radicles and roots in the control was consistently better (P=0.05) than in filtrate solutions of eitherA. niger orT. viride. The hypocotyls of seedlings under the influence of metabolites ofA niger showed greater cambial activity and formed xylem vessels and tracheids with larger lumina.     

Effects of aluminium on growth and cation uptake in seedlings of Eucalyptus mannifera and Pinus radiata

This experiment was designed to examine the effects of aluminium (Al) on the growth of Pinus radiata (D. Don) and Eucalyptus mannifera subsp. mannifera (Mudie) seedlings in culture solutions in a glasshouse to help explain the failure of radiata pine trees on some acid, low fertility soils in Australia on which the native eucalypts flourish. Aluminium (Al) in culture solution increased the growth of roots and shoots of seedlings of both species but while growth of the eucalypt continued to increase with increases in Al to 2.222 μM, growth of the pine was largest at 370 μM Al. In addition to total root length, specific root length (length per unit dry weight), a measure of fineness of the root, increased in the eucalypt seedlings as the substrate Al increased. Growth of the shoots and roots of the pine in the absence of any added Al was extremely poor suggesting that Al, in low concentrations, may be an essential element or ameliorate some other factors in solution culture at low pH. Root and shoot concentrations of K increased with increasing Al, whilst Ca and Mg Concentrations decreased and Mn concentrations were unaffected in both species. Tissue Ca and Mg concentrations were 2 to 3 times higher in the eucalypt seedlings than the pine at all levels of added Al due to greater uptake of these elements by the eucalypt. In contrast, at the highest concentration of Al in the medium, shoot Al concentrations were lower in the cucalypt than in the pine due to a greater proportion of Al being retained in the eucalypt roots. These differences between the seedlings in terms of root growth and tissue cation concentrations may help explain the ability of eucalypt species to maintain vigorous growth on acid soils high in Al and low in Ca and P, where growth of the pines failed.     

A comparative floral and pollination biology ofTricyrtis flava maxim.,T. nana yatabe andT. ohsumiensis

The floral and pollination biology of three closely related species,Tricyrtis flava, T. nana andT. ohsumiensis, were comparatively investigated. The primary pollinator wasBombus diversus in all the three species observed, andAmegilla sp. also acted forT. ohsumiensis. The flowers ofT. flava andT. ohsumiensis bloom for two days and are protandrous. Thus autogramy seems to be prevented in these species when the larger bees forage on them, though geitonogamy may also occur. On the other hand,T. nana appears to be a primarily self-pollinating species. The flowers of this species open only during one day and are homogamy. The stigmata seem to receive much pollen of their own flower by the visit of bumblebees. Moreover, many flowers ofT. nana fruited without any visit of pollinators.T. nana has many features characterizing the autogamous derivatives.     

Comparative study of leaf carbon gain in saplings ofThujopsis dolabrata var.hondai andQuercus mongolica var.grosseserrata in a cool-temperate deciduous forest

Seasonal courses of leaf CO₂ gas exchange in a growing season were examined in saplings ofThujopsis dolabrata var.hondai andQuercus mongolica var.grosseserrata in a cool temperate deciduous forest. Between the two tree species there were no large differences in the light compensation point of leaf photosynthesis, except for the season of new leaf expansion. However, light-saturated rates of net photosynthesis were obviously high inT. dolabrata var.hondai. EvergreenT. dolabrata var.hondai saplings had large photosynthetic production in two seasons, before the emergence of new foliage and after foliage fall of the overstory deciduous trees, because of the significantly high solar radiant energy penetrating under the forest canopy during the seasons. Saplings of deciduousQ. mongolica var.grosseserrata were heavily shaded throughout the growing season by foliage of the overstory trees, which resulted in a low daily surplus production. The annual surplus production of leaves in the growing season was estimated to be 2300 mmol CO₂ m⁻² inT. dolabrata var.hondai and −100 mmol CO₂ m⁻², slightly negative, inQ. mongolica var.grosseserrata. These results supported the high survivability ofT. dolabrata var.hondai saplings and the high mortality ofQ. mongolica var.grosseserrata in the deciduous forest.     

Lupinus species differ in their requirements for iron

The effect of iron supply on the growth and nodulation ofLupinus angustifolius L. (Gungurru),Lupinus luteus L. (R-1171) andLupinus pilosus Murr. (P20957) was studied in acid solutions. Plants of the three species were grown together in the same solution and inoculated withBradyrhizobium (Lupinus) WU 425. Plants were then grown with or without applied NH₄NO₃. The lupin species differed greatly in their sensitivity to low iron concentrations in solution withL. pilosus being most tolerant andL. luteus most sensitive.L. pilosus had the highest iron concentration in tissues and had a higher ratio of iron concentration in the youngest fully expanded leaf blades (YEB) to that in roots than the other two species.L. luteus had higher iron concentrations in roots but lower iron concentration in YEB and shoots than didL. angustifolius. The requirements of internal iron for the maximal chlorophyll synthesis in YEB were 65 μg g^-1 forL. angustifolius andL. luteus, and 52 μg g^-1 forL. pilosus. In contrast to effects on growth, the three species had similar external iron requirements for nodule formation in roots and for maximal nitrogen concentrations in shoots. The results indicate that iron tolerant lupin species require lower internal and external iron supply and have a greater ability than sensitive species to translocate iron from roots to shoots.     

Laminar hydathodes inUrticaceae: Survey of tribes and anatomical observations onPilea pumila andUrtica dioica

Laminar hydathodes are known from only three dicot families. InUrticaceae they are associated with minor vein junctions in all five tribes, as surveyed from cleared leaves of 43 species in 30 genera. Only one species lacked hydathodes. Exclusively adaxial hydathodes were found in 28 genera. In tribeElatostemeae, laminar hydathodes inPilea andPellionia species are abaxial, adaxial, or on both surfaces. Guttation was observed in four species.Urtica dioica (adaxial) andPilea pumila (abaxial) were studied anatomically in detail. Hydathodes in the former have normal bundle structure but xylem gaps sometimes occur. In the latter, phloem is displaced in three previously undescribed ways: 1) ends abruptly near hydathode, 2) curves into connecting vein at adjacent junction, or 3) departs xylem, skirts hydathode independently, and rejoins adjacent xylem strand. Laminar hydathodes are a unifying character of theUrticaceae, and they also strengthen its close relationship to theMoraceae.