Large-scale asymmetric introgression of cytoplasmic DNA reveals Holocene range displacement in a North American boreal pine complex

Jack pine (Pinus banksiana) and lodgepole pine (Pinus contorta var. latifolia) are two North American boreal hard pines that hybridize in their zone of contact in western Canada. The main objective of this study was to characterize their patterns of introgression resulting from past and recent gene flow, using cytoplasmic markers having maternal or paternal inheritance. Mitochondrial DNA (mtDNA) and chloroplast DNA (cpDNA) diversity was assessed in allopatric populations of each species and in stands from the current zone of contact containing morphological hybrids. Cluster analyses were used to identify genetic discontinuities among groups of populations. A canonical analysis was also conducted to detect putative associations among cytoplasmic DNA variation, tree morphology, and site ecological features. MtDNA introgression was extensive and asymmetric: it was detected in P. banksiana populations from the hybrid zone and from allopatric areas, but not in P. contorta populations. Very weak cpDNA introgression was observed, and only in P. banksiana populations. The mtDNA introgression pattern indicated that central Canada was first colonized by migrants from a P. contorta glacial population located west of the Rocky Mountains, before being replaced by P. banksiana migrating westward during the Holocene. In contrast, extensive pollen gene flow would have erased the cpDNA traces of this ancient presence of P. contorta. Additional evidence for this process was provided by the results of canonical analysis, which indicated that the current cpDNA background of trees reflected recent pollen gene flow from the surrounding dominant species rather than historical events that took place during the postglacial colonization.     

Gas exchange and growth responses of ectomycorrhizal Picea mariana, Picea glauca, and Pinus banksiana seedlings to NaCl and Na2SO4

Abstract: Black spruce (Picea mariana), white spruce (Picea glauca), and jack pine (Pinus banksiana) seedlings were inoculated with Hebeloma crustuliniforme or Laccaria bicolor and subjected to NaCl and Na₂SO₄ treatments. The effects of ectomycorrhizas on salt uptake, growth, gas exchange, and needle necrosis varied depending on the tree and fungal species. In jack pine seedlings, ectomycorrhizal (ECM) fungi reduced shoot and root dry weights and in the ECM white spruce, there was a small increase in dry weights. Sodium chloride treatment reduced net photosynthesis and transpiration rates in the three studied tree species. However, NaCl-treated black spruce and jack pine colonized by H. crustuliniforme maintained relatively high photosynthetic and transpiration rates and needle necrosis of NaCl-treated black spruce seedlings was reduced by the ECM fungi. Higher concentrations of Na+ were found in shoots compared with roots of the three examined conifer species. ECM fungi reduced the concentrations of Na+ mainly in the shoots and this reduction was greater in plants treated with NaCl compared with Na₂SO₄. Shoots contained generally higher concentrations of Cl^- compared with roots. In the NaCl-treated black spruce and white spruce, both ECM species significantly reduced Cl^- concentrations. Our results point to overall greater phytotoxicity of NaCl compared with Na₂SO₄ and support our earlier findings which demonstrated beneficial effects of ECM fungi for woody plants exposed to NaCl stress.     

Genetic consequences of glacial survival and postglacial colonization in Norway spruce: combined analysis of mitochondrial DNA and fossil pollen

Norway spruce (Picea abies [L.] Karst.) is a broadly distributed European conifer tree whose history has been intensively studied by means of fossil records to infer the location of full‐glacial refugia and the main routes of postglacial colonization. Here we use recently compiled fossil pollen data as a template to examine how past demographic events have influenced the species’ modern genetic diversity. Variation was assessed in the mitochondrial nad1 gene containing two minisatellite regions. Among the 369 populations (4876 trees) assayed, 28 mitochondrial variants were identified. The patterns of population subdivision superimposed on interpolated fossil pollen distributions indicate that survival in separate refugia and postglacial colonization has led to significant structuring of genetic variation in the southern range of the species. The populations in the northern range, on the other hand, showed a shallow genetic structure consistent with the fossil pollen data, suggesting that the vast northern range was colonized from a single refugium. Although the genetic diversity decreased away from the putative refugia, there were large differences between different colonization routes. In the Alps, the diversity decreased over short distances, probably as a result of population bottlenecks caused by the presence of competing tree species. In northern Europe, the diversity was maintained across large areas, corroborating fossil pollen data in suggesting that colonization took place at high population densities. The genetic diversity increased north of the Carpathians, probably as a result of admixture of expanding populations from two separate refugia.     

Climate influences on growth and reproduction of Pinus banksiana (Pinaceae) at the limit of the species distribution in eastern North America

The presence of conflicts in the allocation of resources among the different functions of an organism is a fundamental postulate of modern ecology. It is assumed that reproduction occurs at a cost because it monopolizes resources that could be used for other functions (e.g., growth). These conflicts may be particularly evident under stressful conditions, such as under low water or nutrient availability, or under severe climatic conditions. There we may expect to find strong negative relationships between an organism's growth and reproduction. We studied a population of Pinus banksiana (Pinaceae) at the northern limit of the species distribution, in subarctic Québec (Canada) where Pinus banksiana occupies nutrient-poor, sandy terraces along the Great Whale river. Serotinous cones of Pinus banksiana produced between 1969 and 1992 were sampled to estimate interannual variations in several variables representing reproduction, and to relate these to climate and tree growth. Climate appears to influence each developmental stage involved in the production of viable seeds, from the time of cone initiation to that of seed maturation. In general, reproductive variables are positively related to high temperatures during the three growing seasons required for seed production; growth is also positively correlated to summer temperatures. Consequently, investment in maturing seeds is positively associated with growth. Thus, both reproduction and growth covary with climate: during relatively warm and long growing seasons, resource allocation to both functions increases. Under these conditions, no trade-off is apparent.     

Tree species dynamics and disturbance in three Swedish boreal forest stands during the last two thousand years

Abstract. Data from three forest stands for the past 2000 yr show how the shade-intolerant species Pinus sylvestris and Betula pubescens maintain significant populations in the Swedish boreal landscape. Age structure data from a northern stand close to the range limits of Picea abies and Pinus complement a local pollen diagram, and reveal cyclic population fluctuations which can be related to periods of climatic stress and fire. Pollen data from two southern stands show that high fire frequencies in the past prevented the expansion of Picea populations. Reduction of the fire frequency during the last 200 yr has favoured Picea. A long time perspective reveals the population dynamics of long-lived species and indicates the controlling factors. Such knowledge permits assessment of the current status and likely future of forest stands.     

Origin and demographic history of the endemic Taiwan spruce (Picea morrisonicola)

Taiwan spruce (Picea morrisonicola) is a vulnerable conifer species endemic to the island of Taiwan. A warming climate and competition from subtropical tree species has limited the range of Taiwan spruce to the higher altitudes of the island. Using seeds sampled from an area in the central mountain range of Taiwan, 15 nuclear loci were sequenced in order to measure genetic variation and to assess the long‐term genetic stability of the species. Genetic diversity is low and comparable to other spruce species with limited ranges such as Picea breweriana, Picea chihuahuana, and Picea schrenkiana. Importantly, analysis using approximate Bayesian computation (ABC) provides evidence for a drastic decline in the effective population size approximately 0.3–0.5 million years ago (mya). We used simulations to show that this is unlikely to be a false‐positive result due to the limited sample used here. To investigate the phylogenetic origin of Taiwan spruce, additional sequencing was performed in the Chinese spruce Picea wilsonii and combined with previously published data for three other mainland China species, Picea purpurea, Picea likiangensis, and P. schrenkiana. Analysis of population structure revealed that P. morrisonicola clusters most closely with P. wilsonii, and coalescent analyses using the program MIMAR dated the split to 4–8 mya, coincidental to the formation of Taiwan. Considering the population decrease that occurred after the split, however, led to a much more recent origin.     

Adaptive gradients and isolation-by-distance with postglacial migration in Picea sitchensis

Fossil pollen records suggest rapid migration of tree species in response to Quaternary climate warming. Long-distance dispersal and high gene flow would facilitate rapid migration, but would initially homogenize variation among populations. However, contemporary clinal variation in adaptive traits along environmental gradients shown in many tree species suggests that local adaptation can occur during rapid migration over just a few generations in interglacial periods. In this study, we compared growth performance and pollen genetic structure among populations to investigate how populations of Sitka spruce (Picea sitchensis) have responded to local selection along the historical migration route. The results suggest strong adaptive divergence among populations (average Q(ST)=0.61), corresponding to climatic gradients. The population genetic structure, determined by microsatellite markers (R(ST)=0.09; F(ST)=0.11), was higher than previous estimates from less polymorphic genetic markers. The significant correlation between geographic and pollen haplotype genetic (R(ST)) distances (r=0.73, P<0.01) indicates that the current genetic structure has been shaped by isolation-by-distance, and has developed in relatively few generations. This suggests relatively limited gene flow among populations on a recent timescale. Gene flow from neighboring populations may have provided genetic diversity to founder populations during rapid migration in the early stages of range expansion. Increased genetic diversity subsequently enhanced the efficiency of local selection, limiting gene flow primarily to among similar environments and facilitating the evolution of adaptive clinal variation along environmental gradients.     

Phylogenetic and biogeographic implications of chloroplast DNA variation in Picea

Purified chloroplast DNA ( cp DNA) extracts from 31 species of Picea and two species of Pinus (P. sylvestris and P. cembra ) were digested with eight restriction endo-nucleases, separated by electrophoresis and scored for restriction fragment length polymorphisms. The resulting data was analyzed phenetically and cladistically. The phenetic analysis indicated lower levels of cpDNA differentiation within Picea than within Pinus and lower levels of differentiation among Eurasian than among North-American Picea species. The cladistic analysis, using Pinus sylvestris as an outgroup, suggested monophyly for Picea and resolved several monophyletic groups among the 31 species of Picea . An assessment of biogeographic events, based on the cladogram, suggests that Picea originated in North-America and that the colonization of Eurasia occurred through separate, intercontinental migrations.     

长白山过渡带红松和鱼鳞云杉径向生长对气候因子的响应

利用树轮生态学方法, 研究了长白山阔叶红松林和暗针叶林过渡带优势树种红松(Pinus koraiensis)和鱼鳞云杉(Picea jezoensis var. komarovii)的生长特征及其与气候因子的关系, 以期揭示气候响应关系的种间差异性。结果表明, 红松和鱼鳞云杉年平均径向生长量与生理年龄显著相关, 红松先于鱼鳞云杉达到最大年生长量, 且红松年平均生长速率显著高于鱼鳞云杉(p < 0.001); 红松和鱼鳞云杉对气候的响应存在差异, 红松与7月份的月平均温度和降水显著正相关, 而鱼鳞云杉与5月平均温度显著正相关, 与5月降水显著负相关。响应面分析进一步证实, 红松生长主要与生长季温度和降水相关, 而生长季初期的降水是限制鱼鳞云杉生长的主要原因。全球变暖有利于红松径向生长, 红松种群有向高海拔上升的可能。     

DNA from bird-dispersed seed and wind-disseminated pollen provides insights into postglacial colonization and population genetic structure of whitebark pine (Pinus albicaulis)

Uniparentally inherited mitochondrial (mt)DNA and chloroplast (cp)DNA microsatellites (cpSSRs) were used to examine population genetic structure and biogeographic patterns of bird-dispersed seed and wind-disseminated pollen of whitebark pine (Pinus albicaulis Engelm.). Sampling was conducted from 41 populations throughout the range of the species. Analyses provide evidence for an ancestral haplotype and two derived mtDNA haplotypes with distinct regional distributions. An abrupt contact zone between mtDNA haplotypes in the Cascade Range suggests postglacial biogeographic movements. Among three cpSSR loci, 42 haplotypes were detected within 28 cpSSR sample populations that were aggregated into six regions. Analysis of molecular variance (amova) was used to determine the hierarchical genetic structure of cpSSRs. amova and population pairwise comparisons (FST ) of cpSSR, and geographical distribution of mtDNA haplotypes provide insights into historical changes in biogeography. The genetic data suggest that whitebark pine has been intimately tied to climatic change and associated glaciation, which has led to range movements facilitated by seed dispersal by Clark's nutcracker (Nucifraga columbiana Wilson). The two hypotheses proposed to explain the genetic structure are: (i) a northward expansion into Canada and the northern Cascades in the early Holocene; and (ii) historical gene flow between Idaho and the Oregon Cascades when more continuous habitat existed in Central Oregon during the late Pleistocene. Genetic structure and insights gained from historical seed movements provide a basis on which to develop recovery plans for a species that is at risk from multiple threats.     

Variation in intra-annual wood formation, and foliage and shoot development of three major Canadian boreal tree species

? Premise of the study: In a warming climate, boreal trees may have adjusted their growth strategy (e.g., onset and coordination of growth among different organs such as stem, shoot, and foliage, within and among species) to cope with the extended growing seasons. A detailed investigation into growth of different organs during a growing season may help us assess the potential effects of climate change on tree growth in the boreal forest. ? Methods: The intra-annual growth of stem xylem, shoot tips, and foliage area of Pinus banksiana, Populus tremuloides, and Betula papyrifera was monitored in a boreal forest in Quebec, Canada during the growing season of 2007. Xylem formation was measured at weekly intervals, and shoot elongation and foliage expansion were measured three times per week from May to September. Growth indices for stem, shoot, and foliage were calculated and used to identify any climate-growth dependence. ? Key results: The time periods required for stem growth, branch extension, and foliage expansion differed among species. Of the three species, P. banksiana had the earliest budburst (20 May) yet the latest completion date of the foliage growth (2 August); P. tremuloides had the latest budburst (27 May) yet the earliest completion date of the foliage growth (10 July). Air temperature positively affected shoot extension growth of all three species. Precipitation positively influenced stem growth of the two broadleaf species, whereas growing season temperature positively impacted stem growth of P. banksiana. ? Conclusion: The results show that both the timing of growth processes and environmental dependences differ among co-occurring species, thereby leading to different adaptive capability of these boreal tree species to climate change.     

On Dispersal Efficiency of Picea Pollen

To learn the dispersal efficiency of picea pollen, surface samples was collected from four Picea forests and adjacent areas in Nei Monggol and Xinjiang Autonomous regions. In the Picea forests at least half and up to 80% of the pollen are of Picea, whereas it is 30% 30 meters away from the forests. In a distance of 500 meters from the margin of the Picea forest it could be as low as 1%. However it also scatters within a radius at least 150Km. Only when the Picea pollen content is higher than 15%, can ir indicate the growth of Picea within an area of dozens of kilometers. Wind velocity and direction as well as the topographic conditions are also important factors affecting dispersal efficiency of Picca pollen. The dispersal pattern for Pious, Abies and Picea are different.     

Competition Within Stands of Picea sitchensis and Pinus contorta

Competition was analysed in plots of Picea sitchensis and Pinuscontorta grown in 50 x 50 hexagonal arrays at 14 cm spacingto ages 7 and 5 years, respectively. Relative growth rates in height (RHGR) became positively relatedto tree heights during the year before harvest. Frequency distributionsof tree heights became negatively kurtotic with tendencies towardsleft-skew ness. By the time of harvest, dead trees were evenlydispersed over the plots. Trees with many taller neighbourshad lower RHGRs than trees with few taller neighbours, and theRHGRs of intermediate-sized trees were correlated with their‘competitive status’. Competition was confined mostlyto first-order neighbours. Large trees depressed the RHGRs ofsmaller neighbours and not vice versa; a simple test for this‘one-sided’ competition is described. Neighboursdid not need to greatly overtop a tree to depress its RHGR,they needed only to be at least as tall. Systematic trends inRHGR across the plots, attributed to site heterogeneity, decreasedwith time, and accounted for only about 10 per cent of the variationin RHGR at harvest. Competitive status accounted for 25 and38 per cent of the variation in RHGR in the Picea sitchensisand Pinus contorta plots, respectively. Picea sitchensis, Pinus contorta, competition, monoculture, self-thinning, relative growth rate, growth model     

Phylogeography of a northeast Asian spruce, Picea jezoensis, inferred from genetic variation observed in organelle DNA markers

Range-wide genetic variation of the widespread cold-temperate spruce Picea jezoensis was studied throughout northeast Asia using maternally inherited mitochondrial DNA and paternally inherited chloroplast DNA markers. This study assessed 33 natural populations including three varieties of the species in Japan, Russia, China, and South Korea. We depicted sharp suture zones in straits around Japan in the geographical distribution pattern of mitochondrial haplotypes (GST=0.901; NST=0.934). In contrast, we detected possible extensive pollen flow without seed flow across the straits around Japan during the past population history in the distribution pattern of chloroplast haplotypes (GST=0.233; NST=0.333). The analysis of isolation by distance of the species implied that by acting as a barrier for the movement of seeds and pollen, the sharp suture zones contributed considerably to the level of genetic differentiation between populations. Constructed networks of mitochondrial haplotypes allowed inference of the phylogeographical history of the species. We deduced that the disjunction with Kamchatka populations reflects range expansion and contraction to the north of the current distribution. Within Japan, we detected phylogeographically different types of P. jezoensis between Hokkaido and Honshu islands; P. jezoensis in Honshu Island may have colonized this region from the Asian continent via the Korean peninsula and the species in Hokkaido Island is likely to have spread from the Asian continent via Sakhalin through land bridges. Japanese endemism of mitochondrial haplotypes in Hokkaido and Honshu islands might have been promoted by separation of these islands from each other and from the Asian continent by the straits during the late Quaternary.     

Seasonal ionic fluctuations and annual growth rates in stands of Pinus silvestris L. and Picea abies Karst. (L.)

Seasonal variations in potassium, calcium, nitrogen, phosphorus, and magnesium contents of needles from two different stands of Pinus silvestris L. and Picea abies (L.) Karst. were determined monthly during a period of two years, and the growth rates once a year during five years. The stands were growing on sand or on clay. The supposed transport of potassium to the shoot during autumn could not be demonstrated. The nutrient status of pine and spruce growing on sand and clay was very similar, except for calcium and magnesium. There were only small differences in ion contents between pine and spruce, except for calcium. On soils with low water-holding capacity, the available water in dry years is the limiting factor affecting the growth rate of spruce but not of pine. It is concluded that the nutrient status of the leaves, the growth rates and the available water in the soil are factors that should have an important part in any discussion of fertilization of coniferous forests.     

贺兰山油松和青海云杉交错区的物种丰富度研究

对贺兰山东坡的油松(Pinus tabulaeformisCarr.)和青海云杉(Picea crassifoliaKom.)交错区进行了野外群落调查和数据分析。结果表明,该交错区跨越2 100~2 500 m海拔范围,油松林和青海云杉林的海拔分界线为2 300~2 350 m。地形是该交错区群落物种丰富度的主要影响因子,阳坡群落的物种丰富度高于阴坡。中等坡度样地的物种丰富度较低;平缓或陡峭样地的物种丰富度较高。海拔升高,物种的丰富度也有所增加。当群落中油松和青海云杉的优势度基本相等时,为典型的油松与青海云杉交错区,在该交错区内,随着油松优势度的降低和青海云杉优势度的升高,土壤条件有所改善,物种丰富度增加。     

Fire history and composition of the subalpine forest of western Colorado during the Holocene

Pollen and plant macrofossils from the Keystone Ironbog are used to document changes in species composition and the dynamics of the subalpine forest in western Colorado over the past 8000 years. Modern pollen spectra (particularly pollen influx), plant macrofossils, observations on modern species composition, and quantified densities and mean basal areas of forest trees are used to interpret the paleoecology of the forest. From 8000 to 2600 years ago the fen was surrounded by a subalpine forest. However, unlike the modern subalpine forest where Abies lasiocarpa (Hooker) Nuttall is slighlty more abundant than Picea engelmannii (Parry) Engelmann, these Holocene forests had a greater dominance of P. engelmannii , perhaps reflecting a summer wet climate like that of the modern southern Rocky Mountains and Colorado Plateau. Mesic conditions promoted a dense understory of Sphagnum moss, forbs, grasses, and shrubs which periodically burned with long (centennial) return-interval and stand-replacing fires. Populus tremuloides Michaux was the dominant successional forest tree 8000–6400 and 4400–2600 years ago, with Picea engelmannii and Abies lasiocarpa becoming reestablished within a couple hundred years. A subalpine meadow or grassland covered the fen for about 2000 years between 6400 and 4400 years ago. Over the past 2600 years a stable, non-successional Pinus contorta (Douglas) spp latifolia (Engelmann) Critchfield forest grew around the fen. This forest stand had a relatively sparse understory. The persistence of Pinus contorta at this elevation (2920 m) probably reflects a shift to drier climatic conditions, perhaps coupled with a change in fire regime to relatively frequent (decadal) surface fires. Following fire Pinus contorta became reestablished at least within 200 years, but the subalpine Picea engelmannii-Abies lasiocarpa forest never regenerated at this elevation     

Chloroplast DNA variation and postglacial recolonization of common ash (Fraxinus excelsior L.) in Europe

We used chloroplast polymerase chain reaction-restriction-fragment length polymorphism (PCR-RFLP) and chloroplast microsatellites to assess the structure of genetic variation and postglacial history across the entire natural range of the common ash (Fraxinus excelsior L.), a broad-leaved wind-pollinated and wind-dispersed European forest tree. A low level of polymorphism was observed, with only 12 haplotypes at four polymorphic microsatellites in 201 populations, and two PCR-RFLP haplotypes in a subset of 62 populations. The clear geographical pattern displayed by the five most common haplotypes was in agreement with glacial refugia for ash being located in Iberia, Italy, the eastern Alps and the Balkan Peninsula, as had been suggested from fossil pollen data. A low chloroplast DNA mutation rate, a low effective population size in glacial refugia related to ash's life history traits, as well as features of postglacial expansion were put forward to explain the low level of polymorphism. Differentiation among populations was high (GST= 0.89), reflecting poor mixing among recolonizing lineages. Therefore, the responsible factor for the highly homogeneous genetic pattern previously identified at nuclear microsatellites throughout western and central Europe (Heuertz et al. 2004) must have been efficient postglacial pollen flow. Further comparison of variation patterns at both marker systems revealed that nuclear microsatellites identified complex differentiation patterns in south-eastern Europe which remained undetected with chloroplast microsatellites. The results suggest that data from different markers should be combined in order to capture the most important genetic patterns in a species.     

树木年龄和断面积对加拿大北方林树木死亡率的影响

以加拿大北部的杨树(Populus spp.)、斑克松(Pinus banksiana)、黑云杉(Picea mariana)为对象,采用长期定位试验,对134块固定样地的活立木及枯死木进行调查,并运用线性回归的方法研究树木年龄、断面积和林分类型对3种树木死亡率的影响.结果表明:随着树龄和断面积的增加,林木的死亡率呈上升趋势.杨树在斑克松林中的死亡率较高,而在黑云杉林中死亡率较低.在黑云杉林中,树龄是影响斑克松死亡率的主要因子;而在杨树林中,断面积是影响斑克松死亡率的重要因子;不同林分类型中树龄对黑云杉死亡率的影响显著.树种组成对树种的死亡率有显著影响;树木年龄、断面积和林分类型之间的交互效应对各树种的死亡率均有显著影响;不同林分类型中同一树种的死亡状况有明显差异.     

Acclimation of respiration to temperature and CO2 in seedlings of boreal tree species in relation to plant size and relative growth rate

The role of acclimation of dark respiration to temperature and CO₂ concentration and its relationship to growth are critical in determining plant response to predicted global change. We explored temperature acclimation of respiration in seedlings of tree species of the North American boreal forest. Populus tremuloides, Betula papyrifera, Larix laricina, Pinus banksiana, and Picea mariana plants were grown from seed in controlled-environments at current and elevated concentrations of CO₂ (370 and 580 μmol mol^–1) in combination with three temperature treatments of 18/12, 24/18, and 30/24 °C (light/dark period). Specific respiration rates of roots and shoots acclimated to temperature, damping increases in rates across growth-temperature environments compared to short-term temperature responses. Compared at a standard temperature, root and shoot respiration rates were, on average, 40% lower in plants grown at the highest compared to lowest growth temperature. Broad-leaved species had a lower degree of temperature acclimation of respiration than did the conifers. Among species and treatment combinations, rates of respiration were linearly related to size and relative growth rate, and relationships were comparable among growth environments. Specific respiration rates and whole-plant respiratory CO₂ efflux as a proportion of daily net CO₂ uptake increased at higher growth temperatures, but were minimally affected by CO₂ concentration. Whole-plant specific respiration rates were two to three times higher in broad-leaved than coniferous species. However, compared to faster-growing broad-leaved species, slower-growing conifers lost a larger proportion of net daily CO₂ uptake as respiratory CO₂ efflux, especially in roots. Interspecific variation in acclimation responses of dark respiration to temperature is more important than acclimation of respiration to CO₂ enrichment in modifying tree seedling growth responses to projected increases in CO₂ concentration and temperature.