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PubMed2004

An ancient bottleneck in the Lost Pines of central Texas.

The retreating edge hypothesis for species responding to climate change predicts severe bottlenecks and eventual extinction. The disjunct Lost Pines population at the westernmost edge of the widespread Pinus taeda range is well suited for testing this prediction. The occurrence of one or more genetic bottlenecks in the Lost Pines population was tested using 34 nuclear microsatellite markers and a control sample from the larger, more continuous east Texas P. taeda forests. The Lost Pines population has undergone drastic contractions in effective population size between 3000 and 30 000 years bp. These results were supported by: (i) detection of transient heterozygosity excess, (ii) a mode-shift indicator of allele frequencies, and (iii) a ratio of allele number to allele size range. No bottleneck was detected for the east Texas control using any of the three methods. The distribution of allele frequencies was skewed for the Lost Pines population compared to the control, indicating a loss of rare alleles. However, allelic diversity was similar between the Lost Pines population and its east Texas control; the mean allele number per locus was 5.29 and 5.38, respectively. It is proposed that the Lost Pines population was the western refugium for P. taeda during Pleistocene glaciation and that East Texas P. taeda forests descended from the bottlenecked Lost Pines population.

Climate↗

PubMed2004

Spatial and population genetic structure of microsatellites in white pine.

We evaluated the population genetic structure of seven microsatellite loci for old growth and second growth populations of eastern white pine (Pinus strobus). From each population, located within Hartwick Pines State Park, Grayling, Michigan, USA, 120-122 contiguous trees were sampled for genetic analysis. Within each population, genetic diversity was high and inbreeding low. When comparing these populations, there is a significant, but small (less than 1%), genetic divergence between populations. Spatial distance between populations or timber harvest at the second growth site were reasonable explanations for the observed minor differences in allele frequencies between populations. Spatial autocorrelation analysis suggested that, for the old growth population, weak positive structuring at 15 m fits the isolation by distance model for a neighbourhood size of about 100 individuals. In comparison, genotypes were randomly distributed in the second growth population. Thus, logging may have decreased spatial structuring at the second growth site, suggesting that management practices may be used to alter natural spatial patterns. In addition, the amount of autocorrelation in the old growth population appears to be lower for some of the microsatellites, suggesting higher numbers of rare alleles and that higher mutation rates may have directly affected spatial statistics by reducing structure.

Forestry↗

PubMed2005

Chloroplast microsatellites: measures of genetic diversity and the effect of homoplasy.

Chloroplast microsatellites have been widely used in population genetic studies of conifers in recent years. However, their haplotype configurations suggest that they could have high levels of homoplasy, thus limiting the power of these molecular markers. A coalescent-based computer simulation was used to explore the influence of homoplasy on measures of genetic diversity based on chloroplast microsatellites. The conditions of the simulation were defined to fit isolated populations originating from the colonization of one single haplotype into an area left available after a glacial retreat. Simulated data were compared with empirical data available from the literature for a species of Pinus that has expanded north after the Last Glacial Maximum. In the evaluation of genetic diversity, homoplasy was found to have little influence on Nei's unbiased haplotype diversity (H(E)) while Goldstein's genetic distance estimates (D2sh) were much more affected. The effect of the number of chloroplast microsatellite loci for evaluation of genetic diversity is also discussed.

Computer Simulation↗

PubMed2006

Chloroplast microsatellites reveal colonization and metapopulation dynamics in the Canary Island pine.

Chloroplast microsatellites are becoming increasingly popular markers for population genetic studies in plants, but there has been little focus on their potential for demographic inference. In this work the utility of chloroplast microsatellites for the study of population expansions was explored. First, we investigated the power of mismatch distribution analysis and the F(S) test with coalescent simulations of different demographic scenarios. We then applied these methods to empirical data obtained for the Canary Island pine (Pinus canariensis). The results of the simulations showed that chloroplast microsatellites are sensitive to sudden population growth. The power of the F(S) test and accuracy of demographic parameter estimates, such as the time of expansion, were reduced proportionally to the level of homoplasy within the data. The analysis of Canary Island pine chloroplast microsatellite data indicated population expansions for almost all sample localities. Demographic expansions at the island level can be explained by the colonization of the archipelago by the pine, while population expansions of different ages in different localities within an island could be the result of local extinctions and recolonization dynamics. Comparable mitochondrial DNA sequence data from a parasite of P. canariensis, the weevil Brachyderes rugatus, supports this scenario, suggesting a key role for volcanism in the evolution of pine forest communities in the Canary Islands.

DNA, Chloroplast↗

PubMed2006

Needle cell elongation and maturation timing derived from pine needle cellulose delta18O.

Estimates of the timing of Pinus arizonica Engelm. needle development in 1998 and 1999 were derived from the leaf-cellulose delta18O of weekly growth increments. Significant correlations were noted between time series of local humidity and leaf-cellulose delta18O for needles growing near Tucson, Arizona. Correlations with temperature were also significant, but much lower, suggesting these variations in cellulose delta18O were determined mostly by changes in humidity. The timing of all significant correlations lags the timing of the appearance of the new needle growth, and is interpreted as indicating 16-23 d were required for cell enlargement in 1998 and 13-17 d in 1999. Similarly, properties of the environmental time series, when significantly correlated, are interpreted as indicating the duration of cellulose deposition (7-27 d in 1998, 13-21 d in 1999). Variations in stable-isotope back diffusion (the Péclet effect) and the synthesis of cellulose using stored photosynthate are discussed as explanations for departures from a Craig and Gordon-type model of leaf water delta18O. The Péclet effect, use of stored photosynthate, and variations in the growing-season source-water delta18O, probably confound the development of a high-resolution paleohumidity proxy from subfossil needle cellulose delta18O in this region.

Cellulose↗

PubMed2006

Diurnal variation of the delta 13C of pine needle respired CO2 evolved in darkness.

The delta 13C of pine needle CO2 evolved in darkness (delta 13Cr) for slash pine trees (Pinus elliottii) was determined by placing recently collected pine needles in darkness and collecting respired CO2 over a short time period (<15 min). Delta 13Cr measurements were made over several 24 h periods to test the hypothesis that significant variation in delta 13Cr would be observed during a diurnal cycle. The delta 13Cr measurements from the 24 h time series trials showed a consistent midday 13C-enrichment (5-10 per thousand) relative to bulk biomass. The delta 13Cr values became more 13C-depleted at night and following shading, and approached bulk-biomass delta 13C values by dawn. The effect of night-time respired 13C-enriched CO2 on the delta 13C value of the remaining assimilate is shown to be minimal (13C depleted by 0.22 per thousand) under field conditions for P. elliottii needles.

Carbon Dioxide↗

PubMed2005

The geographic selection mosaic for squirrels, crossbills and Aleppo pine.

The interactions between many species are structured in a geographic mosaic of populations among which selection is divergent. Here we tested the hypothesis that such a geographic selection mosaic arises for common crossbills (Loxia curvirostra) feeding on seeds in the cones of Aleppo pine (Pinus halepensis) because of geographic variation in the occurrence of European red squirrels (Sciurus vulgaris). On the Iberian Peninsula, Sciurus exerted directional selection favouring larger cones with larger scales, which has caused cones there to be larger than in the Balearic Islands where Sciurus are absent. Moreover, cones on the Iberian Peninsula are so large that they are apparently little used by the relatively small-billed crossbills on the Peninsula; selection by Sciurus seems to have made the cones so difficult to feed on that crossbills rely mostly on the seeds of other conifers. Where crossbills are present but Sciurus are absent (Mallorca Island), cones were smaller as a result of relaxation of selection by Sciurus. However, cones on Mallorca had proportionally thicker scales in comparison to where both Sciurus and crossbills are absent (Ibiza Island), presumably as an adaptation against crossbill predation. Here crossbills specialize on Aleppo pine, have relatively large bills and have apparently coevolved in an arms race with Aleppo pine. These results suggest that Sciurus has influenced both the geographic selection mosaics for crossbills and conifers and the adaptive radiation of crossbills in Eurasia much like Tamiasciurus has done in the North America.

Animals↗

PubMed2005

A role for habitat area in the geographic mosaic of coevolution between red crossbills and lodgepole pine.

Understanding how resource abundance limits adaptive evolution and influences species interactions is an important step towards developing insight into the role of microevolutionary processes in establishing macroevolutionary patterns. We examined how variation in resource abundance (forest area of lodgepole pine Pinus contorta ssp. latifolia) influenced patterns of co-adaptation and coevolution between red crossbill (Loxia curvirostra complex) and lodgepole pine populations. First, we found that crossbill abundance increased logarithmically as forest area increased in mountain ranges lacking a preemptive competitor (pine squirrels Tamiasciurus hudsonicus). Second, seed defences against predation by crossbills increased with increases in crossbill density, suggesting that seed defences have likely evolved in proportion to the intensity of selection that crossbills exert. Third, the average bill size of crossbill populations increased with increasing seed defences, which implies that crossbill offenses increased with increases in seed defences. The large bill size on the largest range is the result of coevolution with lodgepole pine with this crossbill population perhaps speciating. Local adaptation of crossbill populations on smaller ranges, however, is more likely the result of resident crossbills representing a subset of the potential colonists (phenotypic sorting) than of local evolution. In the smallest range, migration and possibly more frequent extinction likely impede local adaptation and may result in maladaptation.

Adaptation, Biological↗

PubMed2006

Replicated population divergence caused by localized coevolution? A test of three hypotheses in the red crossbill-lodgepole pine system.

Several lines of evidence support the hypothesis that local populations of red crossbills (Loxia curvirostra complex) enter into a predator-prey arms race with lodgepole pine (Pinus contorta latifolia) in the absence of competing pine squirrels (Tamiasciurus hudsonicus). Nevertheless, the alternative hypotheses that neutral evolution or factors other than squirrels have caused crossbill population differentiation have not been thoroughly tested. We compared crossbill and pine cone morphology between island populations where squirrels are absent or present, and mainland sites where squirrels are present, in order to distinguish among these hypotheses. All comparisons supported an effect of squirrel absence, not island status, on crossbill and cone morphology. Hence our results provide further evidence that strong localized coevolutionary interactions in a geographic mosaic have driven adaptive population differentiation. In addition, vocal differentiation of crossbills was related to the absence of squirrels, but not to island status. As morphological and vocal differentiation is correlated with reproductive isolation in crossbills, the geographic mosaic of coevolution also seems to promote ecological speciation.

Adaptation, Physiological↗

PubMed2006

Ectomycorrhizal colonization slows root decomposition: the post-mortem fungal legacy.

The amount of carbon plants allocate to mycorrhizal symbionts exceeds that emitted by human activity annually. Senescent ectomycorrhizal roots represent a large input of carbon into soils, but their fate remains unknown. Here, we present the surprising result that, despite much higher nitrogen concentrations, roots colonized by ectomycorrhizal (EM) fungi lost only one-third as much carbon as non-mycorrhizal roots after 2 years of decomposition in a piñon pine (Pinus edulis) woodland. Experimentally excluding live mycorrhizal hyphae from litter, we found that live mycorrhizal hyphae may alter nitrogen dynamics, but the afterlife (litter-mediated) effects of EM fungi outweigh the influences of live fungi on root decomposition. Our findings indicate that a shift in plant allocation to mycorrhizal fungi could promote carbon accumulation in soil by this pathway. Furthermore, EM litters could directly contribute to the process of stable soil organic matter formation, a mechanism that has eluded soil scientists.

Carbon↗

PubMed2005

Pine microsatellite markers allow roots and ectomycorrhizas to be linked to individual trees.

Linking roots and ectomycorrhizas (EcM) to individual host trees in the field is required to test whether individual trees support different ectomycorrhizal communities. Here we describe a method that identifies the source of EcM roots by PCR of polymorphic pine nuclear microsatellite loci using fluorescently labelled primers and high-throughput fragment analysis. ITS-PCR can also be performed on the same EcM DNA extract for fungal identification. The method was tested on five neighbouring Scots pine (Pinus sylvestris var scotica) trees in native woodland. Successful host tree identification from DNA extracts of EcM root tips was achieved for 93% of all root fragments recovered from soil cores. It was estimated that each individual mature pine sampled was colonised by between 15 and 19 EcM fungi. The most abundant fungal species were found on all five trees, and within the constraints of the sampling scheme, no differences between trees in EcM fungal community structure or composition were detected.

DNA Fingerprinting↗

PubMed2005

Three loblolly pine CesA genes expressed in developing xylem are orthologous to secondary cell wall CesA genes of angiosperms.

Specific plant cellulose synthases (CesA), encoded by a multigene family, are necessary for secondary wall synthesis in vascular tissues and are critical to wood production. We obtained full-length clones for the three CesAs that are highly expressed in developing xylem and examined their phylogenetic relationships and expression patterns in loblolly pine tissues. Full-length CesA clones were isolated from cDNA of developing loblolly pine (Pinus taeda) xylem and phylogenetic inferences made from plant CesA protein sequences. Expression of the three genes was examined by Northern blot analysis and semiquantitative RT-PCR. Each of three PtCesA genes is orthologous to one of the three angiosperm secondary cell wall CesAs. The PtCesAs are coexpressed in tissues of loblolly pine with tissues undergoing secondary cell wall biosynthesis showing the highest levels of expression. Phylogenetic and expression analyses suggest that functional roles for these loblolly pine CesAs are analogous to those of orthologs in angiosperm taxa. Based upon evidence from this and other studies, we suggest division of seed plant CesA genes into six major paralogous groups, each containing orthologs from various taxa. Available evidence suggests that paralogous CesA genes and their distinct functional roles evolved before the divergence of gymnosperm and angiosperm lineages.

Cell Wall↗

PubMed2005

Nucleotide variation in genes involved in wood formation in two pine species.

Nucleotide diversity in eight genes related to wood formation was investigated in two pine species, Pinus pinaster and P. radiata. The nucleotide diversity patterns observed and their properties were compared between the two species according to the specific characteristics of the samples analysed. A lower diversity was observed in P. radiata compared with P. pinaster. In particular, for two genes (Pp1, a glycin-rich protein homolog and CesA3, a cellulose synthase) the magnitude of the reduction of diversity potentially indicates the action of nonneutral factors. For both, particular patterns of nucleotide diversity were observed in P. pinaster (high genetic differentiation for Pp1 and close to zero differentiation associated with positive Tajima's D-value for CesA3). In addition, KORRIGAN, a gene involved in cellulose-hemicellulose assembly, demonstrated a negative Tajima's D-value in P. radiata accompanied by a high genetic differentiation in P. pinaster. The consistency of the results obtained at the nucleotide level, together with the physiological roles of the genes analysed, indicate their potential susceptibility to artificial and/or natural selection.

Base Sequence↗

PubMed2006

Rapid nitrogen transfer from ectomycorrhizal pines to adjacent ectomycorrhizal and arbuscular mycorrhizal plants in a California oak woodland.

Nitrogen transfer among plants in a California oak woodland was examined in a pulse-labeling study using 15N. The study was designed to examine N movement among plants that were mycorrhizal with ectomycorrhizas (EM), arbuscular mycorrhizas (AM), or both. Isotopically enriched N (K15NO3-) was applied to gray pine (Pinus sabiniana) foliage (donor) and traced to neighboring gray pine, blue oak (Quercus douglasii), buckbrush (Ceanothus cuneatus) and herbaceous annuals (Cynosurus echinatus, Torilis arvensis and Trifolium hirtum). After 2 wk, needles of 15N-treated pines and foliage from nearby annuals were similarly enriched, but little 15N had appeared in nontreated (receiver) pine needles, oak leaves or buckbrush foliage. After 4 wk foliar and root samples from pine, oak, buckbrush and annuals were significantly 15N-enriched, regardless of the type of mycorrhizal association. The rate of transfer during the first and second 2-wk periods was similar, and suggests that 15N could continue to be mobilized over longer times.

Apiaceae↗

PubMed2006

Foliage influences forced convection heat transfer in conifer branches and buds.

Conifer foliage structures affect branch and bud temperature by altering the development and convective resistance of the thermal boundary layer. This paper examines foliage effects on forced convection in branches and buds of Picea glauca (Moench) Voss and Pinus contorta Dougl. Ex. Loud., two species that represent the range of variation in foliage structure among conifers. Forced convection is characterized by a power law relating Nusselt (heat transfer) and Reynolds (boundary layer development) numbers. Data were collected in a laminar flow wind tunnel for free stream velocities of 0.16-6.95 m s(-1). Scaling parameters were compared against literature values for silver cast branch replicas, a bed of real foliage, cylinders, and tube banks. Foliage structures reduced Nusselt numbers (heat transfer) relative to cylinders, which are typically used to approximate leafless branches and buds. Significantly different scaling relationships were observed for all foliage structures considered. Forced convection scaling relationships varied with foliage structure. The scaling relationships reported here account for variation within populations of branches and buds and can be used to characterize forced convection in a forest canopy.

Convection↗

PubMed2006

Species composition of an ectomycorrhizal fungal community along a local nutrient gradient in a boreal forest.

Soil abiotic factors are considered to be important in determining the distribution of ectomycorrhizal (ECM) fungal species; however, there are few field data to support this. Here, we relate ECM species distributions to changes in soil chemistry along a short (90-m), natural nutrient gradient. The ECM community was characterized, using morphological and molecular techniques, in soil samples collected at 10-m intervals. There were pronounced changes in ECM fungal community structure along the transect, with many taxa showing discrete distributions. Although there was a change of host from Pinus to Picea along the gradient, host-specific fungi did not account for the observed change in community structure. Ordination analyses showed that community structure was strongly correlated with soil characteristics, in particular extractable ammonium and base saturation. However, autocorrelation among soil parameters makes it difficult to isolate the effects of individual parameters. The distinctive changes in soil and vegetation along the transect used in this study provided an exceptional opportunity to examine the local-scale impact of natural spatial heterogeneity on an ECM fungal community.

Biodiversity↗

PubMed2006

Below-ground interactions with arbuscular mycorrhizal shrubs decrease the performance of pinyon pine and the abundance of its ectomycorrhizas.

Few studies have examined how below-ground interactions among plants affect the abundance and community composition of symbiotic mycorrhizal fungi. Here, we combined observations during drought with a removal experiment to examine the effects of below-ground interactions with arbuscular mycorrhizal (AM) shrubs on the growth of pinyon pines (Pinus edulis), and the abundance and community composition of their ectomycorrhizal (EM) fungi. Shrub density was negatively correlated with pinyon above- and below-ground growth and explained 75% of the variation in EM colonization. Consistent with competitive release, pinyon fine-root biomass, shoot length and needle length increased with shrub removal. EM colonization also doubled following shrub removal. EM communities did not respond to shrub removal, perhaps because of their strikingly low diversity. These results suggest that below-ground competition with AM shrubs negatively impacted both pinyons and EM fungi. Similar competitive effects may be observed in other ecosystems given that drought frequency and severity are predicted to increase for many land interiors.

Ecosystem↗

PubMed2006

Measuring external mycelia production of ectomycorrhizal fungi in the field: the soil matrix matters.

Assessing mycorrhizal fungi production in field settings has been hindered by the inability to measure external mycelia. Recently, external mycelia production was measured in the field using a novel in-growth core technique with acid-washed sand as the in-growth matrix. Here, we tested the assumption that external mycelia production in acid-washed sand is representative of that in native soil. External mycelia production was estimated as the difference in fungal growth between closed (allowing only saprotrophic fungal production) and open (allowing mycorrhizal and saprotrophic fungal production) cores using a factorial design of soil matrices (acid-washed sand vs native) and fertilization treatments (control vs nitrogen (N)) in a longleaf pine (Pinus palustris) plantation. In native soils, the ectomycorrhizal to saprotrophic fungal biomass signal was strong and consistent facilitating the assessment of external mycelia production, which was 300% higher than corresponding rates in acid-washed sand and inversely correlated with soil N. These results demonstrate the efficacy and importance of using native soil as the in-growth matrix to measure ectomycorrhizal fungi external mycelia production in field settings.

Ergosterol↗