Spring ephemerals of deciduous forest are adapted to take advantage of the high-light period available in early spring. They appear shortly after snow melt and complete their aboveground growth, including fruit production, within 2 months. After they produce new buds, they senesce and enter dormancy. Dormancy is not very deep in spring ephemerals and during summer differentiation occurs in the bud of the apparently resting organ. Low soil temperatures release dormancy, and the shoots and roots then grow slowly over autumn and winter. The goal of this paper is to show how this characteristic phenology influences many aspects of spring ephemerals' physiology, and the influences these different physiological parameters have on each other. Spring ephemerals have high photosynthetic rates that allow them to rapidly accumulate carbohydrates and complete their aboveground growth in a few weeks. To sustain high photosynthetic rates in early spring, the plants must be able to absorb water efficiently at low soil temperatures and to allocate large amounts of nutrients to the shoot to compensate for lower enzymatic activity at low temperatures. Nutrients are mainly absorbed in spring, although the root system is established in autumn. This means that a large amount of both carbohydrates and nutrients is translocated from the perennial organ to the developing shoot starting in autumn through early spring. Spring ephemerals have low nutrient absorption rates, but high resorption efficiency during leaf senescence. Nevertheless, their high nutrient needs restrict them to rich forest soils. The annual growth rate of spring ephemerals is very slow and this is more likely related to the inherent slow growth rate of the perennial organ than to their short leaf life. As soon as carbohydrate reserves are replenished in spring, sink limitation apparently builds up and induces leaf senescence. A better understanding of the factors controlling the growth rate of spring ephemerals is needed before we can predict these plants' response to climatic changes.
Tissue analysis is commonly used in ecology and agronomy to portray plant nutrient signatures. Nutrient concentration data, or ionomes, belong to the compositional data class, i.e., multivariate data that are proportions of some whole, hence carrying important numerical properties. Statistics computed across raw or ordinary log-transformed nutrient data are intrinsically biased, hence possibly leading to wrong inferences. Our objective was to present a sound and robust approach based on a novel nutrient balance concept to classify plant ionomes. We analyzed leaf N, P, K, Ca, and Mg of two wild and six domesticated fruit species from Canada, Brazil, and New Zealand sampled during reproductive stages. Nutrient concentrations were (1) analyzed without transformation, (2) ordinary log-transformed as commonly but incorrectly applied in practice, (3) additive log-ratio (alr) transformed as surrogate to stoichiometric rules, and (4) converted to isometric log-ratios (ilr) arranged as sound nutrient balance variables. Raw concentration and ordinary log transformation both led to biased multivariate analysis due to redundancy between interacting nutrients. The alr- and ilr-transformed data provided unbiased discriminant analyses of plant ionomes, where wild and domesticated species formed distinct groups and the ionomes of species and cultivars were differentiated without numerical bias. The ilr nutrient balance concept is preferable to alr, because the ilr technique projects the most important interactions between nutrients into a convenient Euclidean space. This novel numerical approach allows rectifying historical biases and supervising phenotypic plasticity in plant nutrition studies.
We investigated the impact of overstory tree leaf phenology on growth rates, carbon allocation pattern, and fruit characteristics in the spring flowering species, Trillium erectum (Liliaceae). Air temperature, overstory canopy closure, and T. erectum phenology were monitored at three locations following a latitudinal gradient in Québec, Canada. Northern sugar maple trees leaf out at cooler temperatures than more southern populations, while Trillium development was initiated at the same soil temperature irrespective of the latitude. Therefore, in northern areas, the time between initiation of T. erectum leaf expansion and canopy closure was shorter than in southern areas, which left less time for northern plants to accumulate reserves before canopy closure. Differences in growth patterns were noted between T. erectum populations. From a south-north gradient, investment to reproduction, total plant biomass, and annual growth rate decreased, while specific leaf area and stem height increased, indicating shade acclimation. The length of the high light period in early spring seems to be a determinant for spring flowering plants' growth and reproduction and may explain the northern distribution limit of some of these species.
Summary• Root carbon (C) partitioning in two host plant species colonized by one of three arbuscular mycorrhizal (AM) fungal species was investigated.• Split-root systems of barley ( Hordeum vulgare ) and sugar maple ( Acer saccharum ) were inoculated on one side with one of three AM fungi. Leaves were labelled with 14 CO 2 3 wk after inoculation. Plants were harvested 24 h later and the root systems from the mycorrhizal (M) and nonmycorrhizal (NM) sides were analysed separately for 14 C.• Partitioning of 14 C between M and NM sides varied depending on the fungal and host plant species used. Gigaspora rosea showed a strong C-sink capacity with both plant species, Glomus intraradices showed a strong C-sink capacity with barley, and Glomus mosseae did not affect 14 C partitioning. The C-sink strength of the M barley roots inoculated with G. rosea or G. intraradices was linearly correlated with the degree of colonization.• The use of three AM fungal and two plant species allowed us to conclude that C-sink strength of AM fungi depends on both partners involved in the symbiosis.
We investigated in the field the carbon (C) transfer between sugar maple (Acer saccharum) saplings and the spring ephemeral Erythronium americanum via the mycelium of arbuscular mycorrhizal (AM) fungi. Sugar maple saplings and E. americanum plants were planted together in pots placed in the ground of a maple forest in 1999. Ectomycorrhizal yellow birches (Betula alleghaniensis) were added as control plants. In spring 2000, during leaf expansion of sugar maple saplings, the leaves of E. americanum were labelled with CO. Seven days after labelling, radioactivity was detected in leaves, stem and roots of sugar maples. Specific radioactivity in sugar maples was 13-fold higher than in yellow birches revealing the occurrence of a direct transfer of C between the AM plants. The quantity ofC transferred to sugar maple saplings was negatively correlated with the percentage of C allocated to the storage organ of E. americanum. A second labelling was performed in autumn 2000 on sugar maple leaves during annual growth of E. americanum roots. Radioactivity was detected in 7 of 22 E. americanum root systems and absent in yellow birches. These results suggest that AM fungi connecting different understorey species can act as reciprocal C transfer bridges between plant species in relation with the phenology of the plants involved.
Acer saccharum Marsh. (sugar maple) is one of only few arbuscular mycorrhizal trees to form extensive stands in northern temperate biomes. Recent maple decline could result from altered intensity and quality of root colonization by associated mycobionts or possible shifts in symbiotic fungal community composition following environmental stresses. In this study the effects on arbuscular mycorrhizal fungi of soil acidification, one of several proposed causal stresses underlying forest decline, and remedial liming were investigated under glasshouse conditions. Acer saccharum seedlings were grown in unsterilized, pH altered, forest soils from healthy and declining maple stands. Over a range of treatment pHs normally tolerated by A. saccharum, fungal populations and responses to pH changes differed between the two soils. The declining site with more acidic soil had an initially larger spore population but lower taxonomic diversity than the healthy site. However, liming stimulated sporulation of several taxa initially apparently absent from the declining site spore population. The quantity of colonization generally increased with pH for both sites. Five Glomus taxa and Scutellospora calospora (Nicol. & Gerd.) Walker & Sanders are added to the list of fungi known to form arbuscular mycorrhizas with A. saccharum, and the known range of Acaulospora cavernata Blaszkowski is extended from Poland to eastern North America.Résumé : Acer saccharum Marsh. (érable à sucre) est l'un des rares arbres endomycorhiziens à former des peuplements étendus dans les biomes tempérés nordiques. Suite au dépérissement des érablières, l'acidification des sols a été suggérée comme stress causal. Bien qu' A. saccharum puisse tolérer l'acidité des sols, les effets de cette acidification sur les symbioses endomycorhiziennes de l'érable ont été jusqu'à maintenant largement ignorés. L'impact du pH du sol sur le taux et la qualité de la colonisation endomycorhizienne ainsi que sur la diversité des Glomales a été étudié en serres sur des semis d'A. saccharum dans des sols provenant respectivement d'érablières en santé et en dépérissement, ajustés à différents pH. Initialement, le site en dépérissement au sol plus acide, avait un nombre plus élevé de spores malgré une diversité taxonomique réduite. Le chaulage subséquent de ce sol a stimulé la sporulation de plusieurs taxons apparemment absents au début de l'expérience. Cette étude a aussi démontré une corrélation positive du pH avec la quantité et la qualité de la colonisation intra-racinaire.
SummaryInvasive plants impose novel selection pressures on na€ ıve mutualistic interactions between native plants and their partners. As most plants critically rely on root fungal symbionts (RFSs) for soil resources, invaders that disrupt plant-RFS mutualisms can significantly depress native plant fitness. Here, we investigate the consequences of RFS mutualism disruption on native plant fitness in a glasshouse experiment with a forest invader that produces known antifungal allelochemicals.Over 5 months, we regularly applied either green leaves of the allelopathic invader Alliaria petiolata, a nonsystemic fungicide to simulate A. petiolata's effects, or green leaves of nonallelopathic Hesperis matronalis (control) to pots containing the native Maianthemum racemosum and its RFSs. We repeatedly measured M. racemosum physiology and harvested plants periodically to assess carbon allocation.Alliaria petiolata and fungicide treatment effects were indistinguishable: we observed inhibition of the RFS soil hyphal network and significant reductions in M. racemosum physiology (photosynthesis, transpiration and conductance) and allocation (carbon storage, root biomass and asexual reproduction) in both treatments relative to the control.Our findings suggest a general mechanistic hypothesis for local extinction of native species in ecosystems challenged by allelopathic invaders: RFS mutualism disruption drives carbon stress, subsequent declines in native plant vigor, and, if chronic, declines in RFS-dependent species abundance.
Global warming might improve fitness of herbaceous species in deciduous forests, mainly by advancing their spring emergence. However, other impacts of global warming such as drier soils in the summer might partly reduce the carbon gain associated with early emergence.
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