Resistance and virulence interactions between two white pine species and blister rust in a 30-year field trial

We monitored infection and mortality of full-sib families of sugar pine (SP) and western white pine (WWP) selected for different mechanisms of resistance to white pine blister rust for more than 30 years in a field test in northern California. Natural infection was enhanced by interplanting alternate host Ribes spp. among test seedlings. Parents of the families were from three geographic provinces, representing the northern Rocky Mountains of Idaho (WWP), the Cascade Range of Oregon and Washington (WWP), and the Siskiyou Mountains of California and Oregon (SP). Several families of each pine species had major gene resistance (MGR), or genes for partial resistance (PR), or both types. Under the chronic epidemic conditions that prevailed, susceptible controls of both species became infected and died within a few years. Families with MGR segregated in expected Mendelian ratios (susceptibility/resistance), but each R gene—Cr1 in sugar pine and Cr2 in western white pine—was eventually defeated by biotypes of the rust with specific virulence to it (vcr1 and vcr2, respectively). WWP families from Idaho, which lacked Cr2, performed the best overall, ranging from 44 to 64% rust-free; a few WWP families from the Cascades, both with and without Cr2, performed comparably. Families with PR were not specifically vulnerable to vcr1 or vcr2. PR was less frequent in SP than WWP, and SP families had steeper infection rate curves than WWP. Combining PR in pedigrees with MGR may prolong protection of MGR by inhibiting selection and epidemic increase of vcr genes in the rust.     

Stand structure and dynamics of wave-typeAbies sachalinensis coastal forest

Stand structure and regeneration pattern were examined inAbies sachalinensis coastal forest in northern Hokkaido. In the forest a similar phenomenon to the wave regeneration in subalpine forests has been found. Wave regeneration has been reported for montaneAbies forests in central Japan and North America. Differences and similarities between wave-type stands in this coastal forest and wave-regenerated montane forests were clarified. The shift of dead tree zone, stand structure and regeneration pattern in wave-type stands are the same as in subalpine wave-regenerated forests. High density of individuals is considered to be an internal factor which causes stand-level dieback and also enables the stands to persist in the severe environment in both forests. A difference between wave-regenerated forests andA. sachalinensis wave-type stands is the number of dead tree zones, which is only one in wave-type stands. Changes of regeneration patterns ofAbies sachalinensis with environmental gradient from seaward to inland were related to this difference.     

The chemical composition of pine foliage in relation to the population dynamics of the pine beauty moth,Panolis flammea,in Scotland

Summary This paper reports part of a study to determine why damaging outbreaks of the pine beauty moth, Panolis flammea (D & S) (Lepidoptera: Noctuidae), in Scotland are frequent on lodgepole pine but do not occur on Scots pine, and why outbreaks on lodgepole pine are mainly confined to trees growing in deep unflushed peat. The elongation of shoots and the growth of needles of Scots pine occurred later in the season than did those of lodgepole pine. The foliage of Scots pine generally had a higher level of nitrogen, and consistently had a higher level of phosporus, but had a consistently lower level of tannins than that of lodgepole pine during the period when the larvae were feeding each year. The nitrogen content of the foliage of lodgepole pine growing in an iron pan soil was generally higher than that of lodgepole pine growing in deep peat during the same period but there were no general differences in the phosphorus or tannin contents of lodgepole pine in the 2 soil types. These findings suggest that Scots pine is a more suitable host plant than lodgepole pine and that the foliage of lodgepole pine growing in deep peat is not more suitable than lodgepole pine growing in an iron pan soil. On the basis of the chemical analyses used in this study, it is concluded that the abundance of pine beauty moth in Scotland is not strongly influenced by the nutritional suitability of its host plants.     

Effect of shade stress on growth,morphology, and carbon dynamics of loblolly pine branches

Summary Shade treatments were applied to the terminal portions of branches in loblolly pine trees to test whether portions of branches were autonomous with respect to carbohydrates. The shade treatments reduced light by 50% and 72% compared with full sun conditions resulting in significant reductions in net photosynthesis. Branch growth (branch diameter and length, branch and needle biomass) decreased significantly within the shaded portion of the branch. Branch and needle morphology were also affected by shading. Lateral branches subtending terminal shoots were labelled with ¹⁴C 2 times during the growing season. No movement of ¹⁴C into the terminal portions of the branch was evident during the first growth flush. However, during the second flush of growth small but statistically significant amounts of ¹⁴C were imported into the terminal portion of the shaded branches from subtending laterals. It was concluded that loblolly pine shoots were usually autonomous with respect to carbohydrate supply, but that carbohydrate movement into the terminal shoot from subtending foliage could occur under conditions of very high stress.     

Lignosulfonate promotes the interaction between Scots pine and an ectomycorrhizal fungus Pisolithus tinctorius in vitro

Lignosulfonate (LS) is a lignin-based polymer obtained as a by-product from paper industry, which may have potential as an amendment with macronutrients. We studied effects of LS on the interaction between Scots pine (Pinus sylvestris L.) seedlings and hypocotyl cuttings and the ectomycorrhizal (ECM) fungusPisolithus tinctorius (Pers.) Coker and Couch. The experiments were performed in vitroon the MMN agar medium containing Fe–LS chelate at the concentrations of 0, 5, 10 and 25 mg/L. Inoculation with P. tinctoriusincreased root growth of the seedlings. Fe–LS enhanced P. tinctorius induced formation of lateral roots and had a dose-dependent positive effect on the establishment of mycorrhizas on the seedlings. The growth of the fungal mycelium was improved by Fe–LS, which might cause faster and more intensive contact with the roots and, thus, better root growth and mycorrhiza formation. P.tinctorius enhanced also adventitious root formation and subsequent root growth of the hypocotyl cuttings but without any synergistic effect with Fe–LS. Our study with P. tinctorius and Scots pine in vitro indicates that a low-cost by-product Fe–LS, obtained from paper industry, may be a potential tool to improve the efficiency of fungal inoculations, thus, facilitating the early interaction between an ECM fungus and host seedling.     

Sexual reproduction and crossing barriers in white pines: the case between <Emphasis Type="Italic">Pinus lambertiana</Emphasis> (sugar pine) and <Emphasis Type="Italic">P. monticola</Emphasis> (western white pine)

The sexual reproductive process in Pinus lambertiana has not been completely described, and previous attempts to generate hybrids with Pinus monticola and other North American pines have not been successful. The nature of incompatibility barriers between P. lambertiana and P. monticola is unknown. This needs to be understood if strategies are to be developed to overcome the said barriers. In this paper, development on interspecific crosses is compared with that on intraspecific crosses on the same parent trees. Pollen grains of both species germinated on the nucellus of both species within a week after pollination. Seed cone receptivity in P. lambertiana came approximately 2 weeks after receptivity in P. monticola, and this delay was perpetuated throughout ovule development in the first year of the reproductive process. Development of the second-year seed cones proceeded more gradually in P. lambertiana. However, seed cones reached maturity only for P. monticola x P. lambertiana. In both crosses, the barriers to hybridization occurred during the second year of the reproductive process. With the P. lambertiana as the seed parent, it was manifested through the failure of the megaspores at the free-nuclear stage to resume development. When P. monticola was used as the seed parent, the male and female gametes failed to fuse. Our results clearly show that the barriers to hybridization in these species occur before or at fertilization. However, the exact mechanisms behind these are still unknown. Based on the results of this study, we present several strategies to bypass the developmental barriers and possibly produce hybrid progenies.     

Recovery from winter depression of photosynthesis in pine and spruce

Summary Recovery from winter depression of photosynthesis was studied in Pinus sylvestris, Pinus conforta and Picea abies by means of chlorophyll fluorescence and gas exchange measurements. During the winter 1986–1987 the fluorescence yield was low and no variable fluorescence was detectable before the end of March. In the field recovery of variable fluorescence/maximum fluorescence (F_v/F_m) during spring was slow for all three species studied. The temperature dependence of recovery was confirmed from measurements of the potential rate of recovery of F_v/F_m at different temperatures in the laboratory. At 20° C, F_v/F_m increased from 0.1 to 0.8 within 3 days. Recovery of F_v/F_m was paralleled by an increase in apparent photon yield. No significant differences could be demonstrated between the studied tree species in potential rate of recovery in the laboratory or in actual recovery in the field.     

Gas exchange and canopy structure of 9-year-old loblolly pine,pitch pine and pitch x loblolly hybrids

Summary Seasonal gas exchange and canopy structure were compared among 9-year-old loblolly pine (Pinus taeda L.), pitch pine (Pinus rigida Mill.), and pitch x loblolly hybrids (Pinus rigida x taeda) growing in an F2 plantation located in Critz, Va., USA. Leaf net photosynthesis, conductance, internal CO₂ concentration (c_i), water use efficiency (WUE; photosynthesis/conductance), dark respiration and the ratio of net photosynthesis/respiration did not vary among or within the three taxa. Significant differences in volume production, crown length, total crown leaf surface area and the silhouette area of shade shoots among the taxa were observed. The loblolly-South Carolina source had greater volume and crown surface area than the pitch pine, and the hybrid taxa were intermediate between the two. Although the silhouette area ratio of shade foliage varied among taxa, it was not related to volume. A strong relationship between total leaf surface area and volume was observed. Leaf conductance, c_i, WUE and leaf water potential were the physiological parameters significantly and positively correlated with volume. This study suggests that the amount of needle surface in the canopy is more important in early stand volume growth than the leaf carbon exchange rate and the degree of needle self-shading in the lower canopy.     

Variation of specific leaf area and upscaling to leaf area index in mature Scots pine

Reliable and objective estimations of specific leaf area (SLA) and leaf area index (LAI) are essential for accurate estimates of the canopy carbon gain of trees. The variation in SLA with needle age and position in the crown was investigated for a 73-year-old Scots pine (Pinus sylvestris L.) stand in the Belgian Campine region. Allometric equations describing the projected needle area of the entire crown were developed, and used to estimate stand needle area. SLA (cm² g⁻¹) as significantly influenced by the position in the crown and by needle age (current-year versus 1-year-old needles). SLA increased significantly from the top to the bottom of the crown, and was significantly higher near the interior of the crown as compared to the crown edge. SLA of current-year needles was significantly higher than that of 1-year-old needles. Allometric relationships of projected needle area with different tree characteristics showed that stem diameter at breast height (DBH), tree height and crown depth were reliable predictors of projected needle area at the tree level. The allometric relationships between DBH and projected needle area at the tree level were used to predict stand-level needle area and estimate LAI. The LAI was 1.06 (m² m⁻²) for current-year needles and 0.47 for 1-year-old needles, yielding a total stand LAI of 1.53.     

云龙天池国家级保护区云南松种群年龄结构及动态分析

云南松是中国西南地区的特有树种。云南松林是暖温性针叶林的典型代表,主要分布于滇中高原。云南松种群是否可以自我更新,将影响滇中地区的植被动态。明晰未受干扰的云南松种群的结构和动态是研究云南松林可持续发展的重要基础。云南省云龙天池自然保护区内的云南松林是较为原始的森林。以云龙天池国家级自然保护区云南松天然种群为研究对象,分析了云南松天然种群静态生命表、存活曲线、生存函数曲线和谱分析。结果表明：云龙天池国家级自然保护区的云南松年龄结构为倒J型,种群年龄结构数量变化动态指数V_pi为28.29%,存在外部干扰时侯的种群年龄结构动态指数V′_pi为1.77%,种群动态指数V_pi和V′_pi都大于0,种群在外部环境干扰下承担的风险概率P_max为6.25%,以上结果表明云南松种群为增长型种群结构,具有一定的抗干扰能力。云南松种群的存活曲线为Deevey-Ⅱ型,生存分析发现生存率、累计死亡率、死亡密度和危险率在Ⅰ—Ⅶ龄级(胸径<35 cm)变化较大,Ⅶ—XVI龄级(胸径35—80 cm)趋于稳...     

Fungi associated with the southern pine beetle: avoidance of induced defense response in loblolly pine

Summary Loblolly pines (Pinus taeda) are rapidly killed by colonizing southern pine beetle (Dendroctonus frontalis). The female beetles carry two species of fungi (Ceratocystis minor var. barrasii and an unnamed basidiomycete) within a mycangium. The insects are also frequently associated with a blue-staining form of C. minor. These fungi are inoculated into the tree during colonization. The tree has an induced defensive response that involves resin soaking and necrosis of affected tissue isolating the invading organlsms. The blue-staining fungus stimulates formation of this response in the tree, but the two mycangial fungi do not. These results suggest that the beetles are closely associated with two highly pathogenic fungi that do not stimulate one of the critical components of tree defense.     

不同林龄和密度对马尾松人工林凋落叶养分变化的影响

该文选择广西南宁市横县镇龙林场的4种林龄(幼龄林、中龄林、成熟林和过熟林)和4种密度(低密度林、中低密度林、中高密度林和高密度林)马尾松人工林共8种林分作为研究对象,分析了未破碎和破碎两个不同降解阶段的凋落叶C、N、P含量及其生态化学计量学特征。结果表明:(1)不同林龄中,凋落叶初始C、N含量在过熟林和成熟林中较高,P含量没有显著变化,且C∶N比值和C∶P比值从幼龄林到成熟林逐渐升高,说明较高林龄马尾松对N和P重吸收较低,而较低林龄马尾松对N和P重吸收较强,需要较大。(2)不同密度林中,随着林木密度的增加,凋落叶初始C含量逐渐升高,N含量无显著变化,P含量降低;高密度林凋落叶的初始C∶P比值和N∶P比值较高,说明高种植密度下马尾松可能对N和P养分的需求较大,P重吸收较强。(3)不同林龄和不同密度马尾松林的破碎凋落叶C含量、C∶N比值、C∶P比值和N∶P比值比未破碎凋落叶的低,N和P含量较高,说明凋落物在降解过程中出现N和P养分的富集现象。(4)中林龄和较高种植密度的马尾松破碎凋落叶与未破碎凋落物的C含量差值最大,C∶N比值和C∶P比值较低,说明这两种林分的凋落叶C的降解速率可能较大。上...     

Fungal gene expression in early symbiotic interactions between Laccaria bicolor and red pine

Ectomycorrhizas are mutualistic symbiotic organs formed by interaction between plant roots and fungi. Mycorrhizal initiation, development and functional maintenance involve morphological changes that are mediated by activation and suppression of several fungal and plant genes. During the pre-infection stage, a harmonized cross-talk takes place between the symbionts, to determine their compatibility. Upon mutual recognition, the symbionts initiate further physiological and morphological changes essential for the formation of the symbiotic organ. In order to understand the molecular mechanisms underlying these events, we developed an interaction-specific cDNA library from Laccaria bicolor that represents fungal genes regulated by its interaction with Pinus resinosa roots. Membrane array analyses of these cDNAs suggested that a wide variety of genes are involved in the pre-infection stage processes.     

Influence of induced water stress in ponderosa pine on pine sawflies

Summary Three levels of water stress were induced on pole-size ponderosa pine (Pinus ponderosa) to determine the influence of plant moisture stress on oviposition, survival, and growth of two species of pine sawfly (Neodiprion fulviceps and N. autumnalis). It was found that water stress affected oviposition and subsequent egg survival but not larval development or survival. Stress had a negative effect on early season oviposition (N. fulviceps) and a positive effect on late season oviposition (N. autumnalis). Egg hatch was different between species and years and among treatment levels. Larval development, feeding, and survival were not affected by water stress. Overall, the effect of stress was not sufficient to explain population outbreaks of sawflies. Several hypotheses are presented as possible explanations for the observed experimental results.     

Complex population genetic structure in the endemic Canary Island pine revealed using chloroplast microsatellite markers

The Canary archipelago, located on the northwestern Atlantic coast of Africa, is comprised of seven islands aligned from east to west, plus seven minor islets. All the islands were formed by volcanic eruptions and their geological history is well documented providing a historical framework to study colonization events. The Canary Island pine (Pinus canariensis C. Sm.), nowadays restricted to the westernmost Canary Islands (Gran Canaria, Tenerife, La Gomera, La Palma and El Hierro), is considered an old (Lower Cretaceous) relic from an ancient Mediterranean evolutionary centre. Twenty seven chloroplast haplotypes were found in Canary Island pine but only one of them was common to all populations. The distribution of haplotypic variation in P. canariensis suggested the colonization of western Canary Islands from a single continental source located close to the Mediterranean Basin. Present-day populations of Canary Island pine retain levels of genetic diversity equivalent to those found in Mediterranean continental pine species, Pinus pinaster and Pinus halepensis. A hierarchical analysis of variance (AMOVA) showed high differentiation among populations within islands (approximately 19%) but no differentiation among islands. Simple differentiation models such as isolation by distance or stepping-stone colonization from older to younger islands were rejected based on product-moment correlations between pairwise genetic distances and both geographic distances and population-age divergences. However, the distribution of cpSSR diversity within the islands of Tenerife and Gran Canaria pointed towards the importance of the role played by regional Pliocene and Quaternary volcanic activity and long-distance gene flow in shaping the population genetic structure of the Canary Island pine. Therefore, conservation strategies at the population level are strongly recommended for this species.Communicated by D.B. NealeA. Gómez and S.C. González-Martínez as joint authors     

Exotic pine plantations and indigenous Lepidoptera in Australia

Temperate regions of Australia have extensive commercial plantations of Monterey pine, Pinus radiata D. Don. Replacement of indigenous forests by P. radiata is likely to have significant effects on assemblages of native Lepidoptera, and has been considered a major threat to native fauna through displacing native species. However, many species of Lepidoptera, including ennomine geometrid moths in the genus Chlenias Guenèe, have adopted P. radiata as a larval host. Comparisons were made of oviposition preferences and nutritional ecology of Chlenias auctaria Guenée on P. radiata and two native host plants [Acacia mearnsii De Wild. (Mimosaceae) and Eucalyptus obliqua LHérit (Myrtaceae)]. Females showed no significant oviposition preference for any of the three hosts. Growth of sixth instar larvae was significantly less on P. radiata than on the native hosts. Pupal weights were significantly lower, suggesting that the fitness of moths reared on P. radiata will be significantly reduced. The life history strategy of C. auctaria, which includes dispersal of first instar larvae by ballooning, may predispose this species to feed on poor quality hosts, and this may be common to other polyphagous species found feeding on P. radiata in southern hemisphere plantations. The impact of P. radiata may have a long term effect on lepidopteran communities, beyond the simple replacement of indigenous host plants leading to extirpations of feeding specialists.     

Inbreeding and conservation genetics in whitebark pine

Whitebark pine (Pinus albicaulisEngelm.) is threatened across its native rangeby an exotic fungal pathogen introduced withinthe last century. Mortality has beenextensive, and projected potential range shiftsbased on impending climate change have revealedfurther pressures to survival and adaptationfor this long-lived, high-elevation conifer. Quantifying genetic variation and the matingsystem of whitebark pine in its northern rangeprovides a basis for effective conservationmeasures. Isozyme analysis of vegetative budtissue revealed high expected heterozygosity(0.262), moderate population differentiation(F_ST = 0.061) and highly significantcorrelations between observed heterozygosityand geographic variables (R² = 0.36,latitude; R² = 0.30 longitude), supportingthe hypothesis that this species recolonizedits current northern range following glacialretreat from several refugia in the Washingtonand Oregon Cascades and in the northernRockies. Mating system analysis based onsimultaneous isozyme analyses of embryo andhaploid megagametophyte tissues foundrelatively high levels of consanguineous matingand selfing for a conifer (t _m =0.73) within populations. Avian seeddistribution by the Clark's nutcracker (Nucifragia columbiana Wilson) appears to bethe overriding factor influencing geneticpatterns: being a mutualistic seed disperser,caches comprised of related seeds develop intoclumped stands with strong family substructure. While it is a critical wildlife habitatcomponent, lack of commercial utilization hasmade in situ adaptation the primaryconservation focus. Encouraging regenerationsuccess and nutcracker caching by maintainingnatural fire regimes will provide anecosystem-based conservation solution; however,in the Rocky Mountains between 52° N and47° N, disease-resistant individualsshould be located and propagated in order toensure long-term survival of the species inhigh pathogen hazard areas.     

The relationship between fire history and an exotic fungal disease in a deciduous forest

Exotic diseases have fundamentally altered the structure and function of forest ecosystems. Controlling exotic diseases across large expanses of forest has proven difficult, but fire may reduce the levels of diseases that are sensitive to environmental conditions. We examined Cornus florida populations in burned and unburned Quercus–Carya stands to determine if burning prior to anthracnose infection has reduced the impacts of an exotic fungal disease, dogwood anthracnose, caused by Discula destructiva. We hypothesized that fire has altered stand structure and created open conditions less conducive to dogwood anthracnose. We compared C. florida density, C. florida health, and species composition and density among four sampling categories: unburned stands, and stands that had burned once, twice, and 3 times over a 20-year period (late 1960s to late 1980s). Double burn stands contained the greatest density of C. florida stems (770 stems ha⁻¹) followed by triple burn stands (233 stems ha⁻¹), single burn stands (225 stems ha⁻¹) and unburned stands (70 stems ha⁻¹; P < 0.01). We observed less crown dieback in small C. florida trees (<5 cm diameter at breast height) in burned stands than in unburned stands (P < 0.05). Indicator species analysis showed that burning favored species historically associated with Quercus–Carya forests and excluded species associated with secondary succession following nearly a century of fire suppression. Our results suggest that fire may mitigate the decline of C. florida populations under attack by an exotic pathogen by altering forest structure and composition. Further, our results suggest that the burns we sampled have had an overall restorative effect on forest communities and were within the fire return interval of the historic fire regime. Consequently, prescribed fire may offer a management tool to reduce the impacts of fungal disease in forest ecosystems that developed under historic burning regimes.     

Hurricane-induced modification of nitrogen and phosphorus resorption in an aspen clone: an example of diffuse disturbance

Summary On 27 September 1985, Hurricane Gloria intersected the Rhode Island land mass and disrupted normal autumnal element resorption in 20 ramets of a trembling aspen clone (Populus tremuloides). No mechanical injury to the plants was observed, but in contrast to undamaged aspens, the wind/salt-damaged aspens did not withdraw nitrogen (N) or phosphorus (P) from senescing leaves. There was actually 17% more N in abscising leaves than in presenescent, pre-hurricane leaves.     

Evidence for recombination and segregation of virulence to pine in a hybrid cross between Gibberella circinata and G. subglutinans

Two species associated with the Gibberella fujikuroi species complex, G. circinata (the cause of pitch canker in pines) and G. subglutinans (avirulent on pine), were found to have limited interfertility in hybrid crosses. MAT idiomorphs, polymorphisms in the histone H3 gene, vegetative compatibility, and virulence phenotypes were used to verify recombination. The MAT idiomorphs appeared to be assorting independently, but the histone H3 haplotype was disproportionately represented by that of the G. subglutinans parent. Ninety-eight percent (45/46) of the progeny tested were vegetatively incompatible with both parents. All F₁ progeny were avirulent to pine, but a wide range of virulence was restored through a backcross to the virulent parent (G. circinata). Attempts at hybrid crosses using other isolate combinations were rarely successful (1/26). This limited interfertility supports retention of G. circinata and G. subglutinans as separate species, but offers opportunities to characterize the inheritance of virulence to pine.