A high-density genetic linkage map of a black spruce (Picea mariana) × red spruce (Picea rubens) interspecific hybrid

Genetic maps provide an important genomic resource of basic and applied significance. Spruce (Picea) has a very large genome size (between 0.85 × 1010 and 2.4 × 1010 bp; 8.5-24.0 pg/1C, a mean of 17.7 pg/1C ). We have constructed a near-saturated genetic linkage map for an interspecific backcross (BC1) hybrid of black spruce (BS; Picea mariana (Mill.) B.S.P.) and red spruce (RS; Picea rubens Sarg.), using selectively amplified microsatellite polymorphic loci (SAMPL) markers. A total of 2284 SAMPL markers were resolved using 31 SAMPL-MseI selective nucleotide primer combinations. Of these, 1216 SAMPL markers showing Mendelian segregation were mapped, whereas 1068 (46.8%) SAMPL fragments showed segregation distortion at α = 0.05. Maternal, paternal, and consensus maps consistently coalesced into 12 linkage groups, corresponding to the haploid chromosome number (1n = 1x = 12) of 12 in the genus Picea. The maternal BS map consisted of 814 markers distributed over 12 linkage groups, covering 1670 cM, with a mean map distance of 2.1 cM between adjacent markers. The paternal BS × RS map consisted of 773 markers distributed over 12 linkage groups, covering 1563 cM, with a mean map distance of 2.0 cM between adjacent markers. The consensus interspecific hybrid BC1 map consisted of 1216 markers distributed over 12 linkage groups, covering 1865 cM (98% genome coverage), with a mean map distance of 1.5 cM between adjacent markers. The genetic map reported here provides an important genomic resource in Picea, Pinaceae, and conifers.     

Appearence and localization of a β-glucosidase hydrolyzing coniferin in spruce (Picea abies) seedlings

-Glucosidase activity for coniferin (coniferyl alcohol -D-glucoside) is not present in spruce (Picea abies L. Karst.) seeds but appears in the young seedlings. Lignification starts at ca. day 9 of germination in the vascular bundles. An antiserum against glucosidase 1 isolated from spruce seedlings (Marcinowski and Grisebach, Eur J. Biochem. 1978) was employed for the localization of the enzyme in cross sections of hypocotyls using immunofluorescent technique. The results indicate that at this stage of development the glucosidase is localized at the inner layer of the secondary cell wall. Glucosidase activity was present in all cells of the investigated hypocotyl tissue.     

Triggering and predisposing factors in the “Red” decline syndrome of Norway spruce (Picea abies)

Summary Analysis of 62 mature Norway spruce (Picea abies provenance Viborg) trees growing in a Danish plantation was undertaken along with analysis of their nutrient contents (N, P, K, Ca, Mg, Fe, Mn, Cu, Zn, B and Na), in each of the three youngest needle age classes, from branches of four exposure directions near the tree top. The aim was to investigate if one among the studied possible predisposing factors was also a triggering factor in the 1989 outbreak of the Red Norway spruce decline in Denmark. Neither nutrient imbalance or deficiency, nor excessive N-deposition or salt-stress were indicated as triggering factors in 1989. The Red syndrome, noticeable for the bright red colour of the current-year needles, was found to be an extension of the European type Novel Decline. Red syndrome is similar to previously reported phenomena of top-dying and sub top-dying, in that it had fewer needle age classes and significantly higher contents of mobile cations (and Ca) in the younger needle classes. Tree ring analysis suggested that the Red syndrome was initiated in the early 1980s, when the trees experienced adverse climatic conditions. Because of this long-term development of the Red Norway spruce decline syndrome, it is concluded that a triggering factor is of minor importance relative to the multitude of predisposing factors.     

Emission of hydrogen sulfide by twigs of coniferes — acomparison of Norway spruce (Picea abies (L.) Karst.), Scotch pine (Pinus sylvestris L.) and Blue spruce (Picea pungens Engelm.)

The emission of reduced volatile sulfur compounds from twigs of Norway spruce (Picea abies (L.) Karst.) was measured in the field by cryosampling and gaschromatographic analysis. Trees were growing in the Erzgebirge (E-Germany) at Oberbärenburg and at the Kahleberg and at a third stand in NW-Bavaria (S-Germany). Emission rates were also measured for Scotch pine (Pinus sylvestris L.) and Blue spruce (Picea pungens Engelm.) at the Kahleberg. Twigs still attached to the trees were enclosed in a flow-through gas exchange cuvette. H₂S was detected as the predominant reduced sulfur compound emitted from the twigs. The mean H₂S emission rate from twigs of Norway spruce varied between 0.04 pmol kg^-1 dw s^-1 at Würzburg and 6.21 pmol kg^-1 dw s^-1 at the Kahleberg. Comparing different species at the Kahleberg, the mean H₂S emission rate was almost the same from twigs of Norway spruce (6.2 pmol kg^-1 dw s^-1) and Blue Spruce trees (5.9 pmol kg^-1 dw s^-1) but it was approximately 18 times higher for Scotch pine (110 pmol kg^-1 dw s^-1). The percentage of SO₂-exclusion via H₂S-emission of the tree species investigated at the Kahleberg is calculated on the basis of data on SO₂ fluxes. It is very small for Norway spruce and Blue spruce. However, for Scotch pine, H₂S emission contributes about 10% to the detoxification of SO₂.     

Contrasting ozone × pathogen interaction as mediated through competition between juvenile European beech (Fagus sylvatica) and Norway spruce (Picea abies)

Based on the growth-differentiation balance theory (GDB) and the influence of tropospheric ozone (O₃) on plants, we hypothesized that pre-conditioning with elevated O₃ reduces adverse effects of the root rot pathogen Phytophthora citricola Sawada. To this end a 2-year phytotron study with juvenile European beech (Fagus sylvatica L.) and (Picea abies [L.] Karst.) grown in mixture was performed. The hypothesis was tested on phenological, leaf and root morphological as well as physiological aspects of plant performance. Contrasting with spruce, elevated O₃ limited leaf and root biomass development, photosynthetic performance and N uptake of beech. The growth limitation by O₃ conveyed increased resistance in beech against the pathogen. Conversely, spruce displayed enhanced susceptibility in the combined O₃/P. citricola treatment. The hypothesis was supported in the case of beech rather than spruce. Nevertheless, conclusions support GDB regarding the trade-off between growth and stress defense, although compliance appears to be species-specific.     

Does soil acidity reduce subsoil rooting in Norway spruce (Picea abies)?

Increasing evidence suggests that forest soils in central and northern Europe as well as in North America have been significantly acidified by acid deposition during the last decades. The present investigation was undertaken to examine the effect of soil acidity on rooting patterns of 40-year-old Norway spruce trees by comparing fine and coarse roots among four stands which differed in soil acidity and Mg (and Ca) nutrition. The coarse root systems of four to five 40-year-old Norway spruce trees per stand were manually excavated. The sum of cross sectional area (CSA) at 60 cm soil depth and below of all vertical coarse roots, as a measure of vertical rooting intensity, was strongly reduced with increasing subsoil acidity of the stands. This pattern was confirmed when 5 additional acidic sites were included in the analysis. Fine root biomass in the mineral soil estimated by repeated soil coring was strongly reduced in the heavily acidified stands, but increased in the humic layer. Using ingrowth cores and a screen technique, we showed that the higher root biomass in the humic layer of the more acidic stands was a result of higher root production. Thus, reduced fine root biomass and coarse root CSA in deeper soil layers coincided with increased root growth in the humic layer. Root mineral analysis showed Ca/Al ratios decreased with decreasing base saturation in the deeper mineral soil (20–40 cm). In the top mineral soil, only minor differences were observed among stands. In general, low Ca/Al ratios coincided with low fine root biomass. Calcium/aluminum ratios determined in cortical cell walls using X-ray microanalysis showed a similar pattern as Ca/Al ratios based on analysis of whole fine roots, although the amplitude of changes among the stands was much greater. Aluminum concentrations and Ca/Al ratios in cortical cell walls were at levels found to inhibit root growth of spruce seedlings in laboratory experiments. The data support the idea that Al (or Ca/Al ratios) and acid deposition-induced Mg (and possibly Ca) deficiency are important factors influencing root growth and distribution in acidic forest soils. Changes in carbon partitioning within the root system may contribute to a reduction in deep root growth.     

Can spruce beetle (Dendroctonus rufipennis Kirky) pheromone trap catches or stand conditions predict Engelmann spruce (Picea engelmannii Parry ex Engelm.) tree mortality in Colorado?

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Genetic structure of a rare European conifer, Serbian spruce (Picea omorika (Panč.) Purk.)

Genetic variation in 13 populations of a Balkan endemic, Serbian spruce (Picea omorika (Panč.) Purk.), was investigated using 16 isozyme loci. Serbian spruce is characterized by low levels of genetic variation (average proportion of polymorphic loci was 20.9% and average expected heterozygosity was 0.067). In most populations, a significant surplus of heterozygotes was observed, indicating a strong selection against inbreds. In the largest populations, fixation indices were positive, probably due to within-population differentiation and the associated Wahlund effect. Despite a strong overall differentiation (F_ST=0.261), no geographical trends in the genetic variation could be identified. Genetic drift caused by small effective population sizes and a strong fragmentation is the most plausible explanation for such variation patterns. A Bayesian analysis of population structure revealed the existence of two clusters, which are supposed to be possible remnants of ancient differentiation within a large range of the predecessor of Serbian spruce, Picea omoricoides Weber. Ex situ conservation measures, namely establishing seed orchards from trees of mixed origin, are proposed as a necessary complement to in situ protection of natural stands.     

A comparison of genomic selection models across time in interior spruce (Picea engelmannii × glauca) using unordered SNP imputation methods

Genomic selection (GS) potentially offers an unparalleled advantage over traditional pedigree-based selection (TS) methods by reducing the time commitment required to carry out a single cycle of tree improvement. This quality is particularly appealing to tree breeders, where lengthy improvement cycles are the norm. We explored the prospect of implementing GS for interior spruce (Picea engelmannii × glauca) utilizing a genotyped population of 769 trees belonging to 25 open-pollinated families. A series of repeated tree height measurements through ages 3–40 years permitted the testing of GS methods temporally. The genotyping-by-sequencing (GBS) platform was used for single nucleotide polymorphism (SNP) discovery in conjunction with three unordered imputation methods applied to a data set with 60% missing information. Further, three diverse GS models were evaluated based on predictive accuracy (PA), and their marker effects. Moderate levels of PA (0.31–0.55) were observed and were of sufficient capacity to deliver improved selection response over TS. Additionally, PA varied substantially through time accordingly with spatial competition among trees. As expected, temporal PA was well correlated with age-age genetic correlation (r=0.99), and decreased substantially with increasing difference in age between the training and validation populations (0.04–0.47). Moreover, our imputation comparisons indicate that k-nearest neighbor and singular value decomposition yielded a greater number of SNPs and gave higher predictive accuracies than imputing with the mean. Furthermore, the ridge regression (rrBLUP) and BayesCπ (BCπ) models both yielded equal, and better PA than the generalized ridge regression heteroscedastic effect model for the traits evaluated.     

Age-dependent tree-ring growth responses of Schrenk spruce (Picea schrenkiana) to climate—A case study in the Tianshan Mountain,China

For both its climatic and ecological importance, Schrenk spruce (Picea schrenkiana) is a crucial tree species living at mid-altitude on the western area of the Tianshan Mountains. It plays a key role on understanding climatic change in the Tianshan Mountains in the past 500 years. However, whether the relationship between tree growth and limiting climate factors is stable over time is still not well-known. In this study, standard and residual chronologies of four 100-year age classes (AC1 < 110a, 110a < AC2 < 210a, 210a < AC3 < 310a and AC4 > 310a) were established for detecting divergence in climate–growth relationships as well as comparing low-frequency and high-frequency variations. The results show that climate can account for a high amount of variance in tree-ring width and higher climate sensitivity was detected in younger trees. Younger trees (<210a) exhibit significantly negative growth responses to mean monthly air temperature of previous June and positive relationship with total monthly precipitation of current April and May, while mean monthly air temperature of current March may inhibit growth of older trees (>210a). Tree-ring chronology statistics and response function reveal that the age-growth patterns are non-monotonic. Our results together with previous studies demonstrate that the age effects on tree-ring growth–climate response is attributed to a combination of genetic characteristics and site microclimate, which suggests that it is necessary to consider both age-dependent and species-specific climate responses when using tree-ring measurements as a proxy for valid climate reconstructions.     

Callose in roots of Norway spruce (Picea abies (L.) Karst.) is a sensitive parameter for aluminium supply at a forest site (Höglwald)

Under controlled environmental conditions in nutrient solution experiments induction of non-constitutive callose in roots has been shown to be a symptom of aluminium (Al) toxicity. In the present study roots of Norway spruce were sampled from a forest site where soil conditions had been modified by acidic irrigation and liming (Höglwald Experiment in Bavaria, Germany). A significant positive relationship was found between the callose content in short roots and the Al concentration in the soil solution, particularly if free Al, rather than total concentrations of soluble Al, were used for prediction. At the same sites root growth of Norway spruce was not affected by free Al concentrations in the range of 2.5 to 199 µM Al. The results show that also under field conditions a positive relationship between Al supply and callose content can be established. In Norway spruce callose content in roots is a much more sensitive parameter for Al supply than root growth.     

γ-Aminobutyric acid (GABA) homeostasis regulates pollen germination and polarized growth in Picea wilsonii

γ-Aminobutyric acid (GABA) is a four-carbon non-protein amino acid found in a wide range of organisms. Recently, GABA accumulation has been shown to play a role in the stress response and cell growth in angiosperms. However, the effect of GABA deficiency on pollen tube development remains unclear. Here, we demonstrated that specific concentrations of exogenous GABA stimulated pollen tube growth in Picea wilsonii, while an overdose suppressed pollen tube elongation. The germination percentage of pollen grains and morphological variations in pollen tubes responded in a dose-dependent manner to treatment with 3-mercaptopropionic acid (3-MP), a glutamate decarboxylase inhibitor, while the inhibitory effects could be recovered in calcium-containing medium supplemented with GABA. Using immunofluorescence labeling, we found that the actin cables were disorganized in 3-MP treated cells, followed by the transition of endo/exocytosis activating sites from the apex to the whole tube shank. In addition, variations in the deposition of cell wall components were detected upon labeling with JIM5, JIM7, and aniline blue. Our results demonstrated that calcium-dependent GABA signaling regulates pollen germination and polarized tube growth in P. wilsonii by affecting actin filament patterns, vesicle trafficking, and the configuration and distribution of cell wall components.     

Relations of cell wall bound peroxidases,phenols and lignin in needles of Serbian spruce Picea omorika (Pančić) Purkynĕ in the natural habitat

The habitat of Picea omorika (Pančić) Purkynĕ, Serbian spruce, is characterized by extremely variable environmental conditions. Relations between the activities of cell wall bound peroxidase (POD) and contents of lignin and cell wall bound phenols were studied in needles of Serbian spruce, in the natural habitat on Mountain Tara. We intended to see, by using Redundancy Analysis, if the yearly peaks of activities of cell wall bound enzymes correspond to the cell wall bound phenols and lignin contents, and whether this relation contributes to the adaptation of Serbian spruce to severe habitat. The highest lignin content was found in spring and it was in high positive correlation with the activities of five covalent POD isoforms. The contents of cell wall bound phenols were lowest in spring and higher in summer and autumn, being in high positive correlation with ionic POD isoforms. The results indicate that relation found between ionic POD isoforms and cell wall bound phenols, as well as between covalent isoforms and lignin may be related to the environmental conditions.     

马衔山青杄(Picea wilsonii)林线性演替初探

植物群落演替的研究已有一百多年的历史了。虽然有许多学者对各种植物群落的各类演替进行了详细地研究,但基本上是描述性和解释性的。近代生态学发展的一个基本特征,就是由定性转为定量,由离散地解释性描述转为系统地动态描述。植物群落演替的研究更应该如此。从定量的动态描述和对演替进行预测的角度出发,我们曾把演替归为系统;从演替过程的角度又分成线性系统和非线性系统两种类     

Distribution of sulphur forms in soils from beech and spruce forests of Mont Lozère (France)

From a quantitative inventory of sulphur forms and sulphur budget, the relation between the distribution of the various sulphur forms and the sulphate fluxes in three soil profiles has been addressed. These profiles are located in two forested watersheds at Mont Lozère. One has been sampled in a beech forest and the other two in a spruce forest and in a harvested plot of this spruce forest, respectively. The mean annual input-output budgets showed a sulphur immobilization in the soil cover of the three plots. In the preserved spruce forest plot, because of larger dry depositions, the sulphur immobilization is much greater than in other plots and occurs essentially in the B horizons. In the other two profiles, the dominant immobilization occurs in the parent material.The total sulphur content is very high in the forest floor reaching 2065 g S g^-1 in the litter of one of the soils under spruce. In the organo-mineral horizons of soils under spruce, the total sulphur content decreases with depth and ranges from 310 to 520 g S g^-1 in the A horizons to 100–200 g S g^-1 in the parent material. In the profile under beech, the total sulphur content is lower except in the parent material. In all cases, the organic sulphur is the major part of sulphur often representing more than 90% of total sulphur. In organo-mineral soil horizons of the spruce forest, the part of the sulphateesters is more important than in the soil of the beech forest, probably related to the different nature of the microbial activity in the spruce forest. In contrast, the humification processes are more efficient in the soil under beech, which can be due to the greater input of organic sulphur by litterfall. It appears that the dominant organic sulphur form varies as a function of microbial ecology and sulphate flux. The maximum of the inorganic sulphate is located at the base of the B horizons in the soil of the spruce forest and in the parent material of the soil under beech. In these horizons, the high content of inorganic sulphate can be related to the higher amounts of amorphous Fe and Al phases.     

管涔山青扦(Picea wilsoni)天然林年龄结构及其动态的研究

对种群年龄结构的研究表明,虽经人为频繁干扰,管涔山青扦天然林仍表现出异龄林结构特征,立木年龄范围超过一个龄级期,根据年龄结构特征值可分为相对同龄林、相对异龄林和异龄林３种类型。林下新一代种群的数量和结构受林冠郁闭度和结构的影响。具垂直郁闭型林冠的异龄林,林下更新数量充足,幼苗幼树年龄结构合理;而水平郁闭型林冠,不利于新一代种群的发生和发展。青扦种群年龄结构受种群发生和自疏两个过程的控制,林下种群的发生以小规模林冠空隙干扰下的连续更新为主。青扦华北落叶松混交林,在其共同适生范围内是某种干扰格局控制下的稳定群落     

Morphogenesis in Sitka spruce (<Emphasis Type="Italic">Picea sitchensis</Emphasis> (Bong.) Carr.) bud cultures – tree maturation and explants from epicormic shoots

The adventitious bud forming ability of Sitka spruce (Picea sitchensis (Bong.) Carr.) buds in vitro was found to be dependent upon the age of tree from which the explants were taken. Bud formation declined exponentially with increasing tree age when 1.0 and 10 μM 6-benzylaminopurine (BA) were used to induce adventitious buds. When less BA was used (0.1 μM) bud production was much lower with all ages of tree and no mathematical relationship between declining bud production and tree age was found. By a tree age of 6 years bud-forming ability had declined severely. Even the few buds that developed on older tree tissues failed to elongate into shoots, became necrotic and eventually died, indicating that adventitious bud induction in this species is not a rejuvenative process. Callusing of bud explants also declined with increasing tree age when 0.1 μM BA was used whilst very little callusing occurred at the higher cytokinin concentrations (1.0 and 10 μM BA). Tissue necrosis in vitro increased with tree age, whilst the ability of BA to retard necrosis declined with increasing tree age. Buds from epicormic shoots, formed on the lower trunks of 20-year-old trees when these were exposed to light, were not significantly better at forming adventitious buds in vitro than buds taken from the lower branches of the crown.     

管涔山青扦(Picea wilsoni)天然林年龄结构及其动态的研究

地种群年龄的研究表明,虽经人为频繁干扰,管涔肝扦天然林仍表现出异龄林结构特征,立木年龄范围超过一个级期,根据年龄结构特征值可分为相对龄林、相对异龄林和异龄林３种类型林下新一代种群的数量和结构受林冠郁闭度和结构的影响。具垂直郁闭型林冠的异龄林,林下更新数量充足,幼苗幼树年龄结构合理;而服闭型林冠,不利于一代种群的发生和发展,青扦种群年龄结构受种群发生和自疏两个过程的控制,林下种群的发生以小规模林冠空     

Variability of antioxidant enzyme activity and isoenzyme profile in needles of Serbian spruce (Picea omorika (Panč.) Purkinye)

Variations were studied of the activity and isoenzyme patterns of soluble peroxidase, catalase, catechol oxidase and superoxide dismutase, in needles of the Balkan endemic conifer Serbian spruce, Picea omorika (Pan?.) Purkinye. The samples were collected from the natural habitat of the species, Mt. Tara. Seasonal changes were found to affect enzymatic activities and isoenzyme profiles. Total protein content was significantly lower in the summer than in other seasons. Several isoforms of peroxidase, catechol oxidase and superoxide dismutase (SOD), as well as two catalase isoenzymes were detected. The number of peroxidase isoenzymes was greatest during the vegetative season. Catalase and catechol oxidase peaked in summer and spring, respectively. Total SOD and Mn-SOD activities were significantly higher in the winter samples than the summer ones.     

青海云杉(Picea crassifolia)种群格局的分形特征及其影响因素

应用计盒维数、信息维数和关联维数对祁连山主要森林类型青海云杉(Picea crassifolia)种群空间格局进行的研究表明:青海云杉种群格局计盒维数在0.992～1.938之间,平均值为1.716,信息维数在0.996～1.860之间,平均值为1.632, 关联维数在1.418～1.759之间,平均值为1.568.计盒维数与样地海拔和土壤水分含量呈显著的负相关,与样地坡向呈显著的正相关,信息维数与样地海拔呈显著负相关而与坡向呈显著的正相关,关联维数与样地坡度呈显著的负相关,主成分分析表明,海拔和土壤含水量是影响青海云杉种群格局分形维数第一主成分,坡向是第二主成分,坡度是第三主成分,这3个主成分累计解释率为87.6%.