Morphological analysis of early contacts between pine roots and two ectomycorrhizal Suillus strains

 Selection of ectomycorrhizal strains for application in forestry is mostly based on the evaluation of symbiotic performance in small-scale experiments. Two Suillus collinitus strains isolated from a Mediterranean and an alpine area were inoculated onto two pine tree species (Pinus pinea and P. nigra ssp. laricio var. corsicana) typical of these two environments. The early events during contact between the cell surfaces of plant and fungal partners were analysed morphologically using ultrastructural and immunocytochemical techniques. All four plant-fungus combinations led to a similar degree of mycorrhizal infection and to a similar colonization pattern. The first contact of fungal hyphae with root cap cells usually involved breakdown of the outermost electron-opaque layer of the plant cell walls. Hyphae further developed between this layer and the underlying wall strata. Ultrastructural observations revealed that S. collinitus strain J3-15-24, isolated from a Mediterranean area, induced a defence reaction in the roots of P. nigra, which grows typically in alpine areas. These observations suggest functional differences between the two fungal strains in their mycorrhizal capabilities. Accepted: 3 April 1998     

Light attenuation in a 14-year-old loblolly pine stand as influenced by fertilization and irrigation

 We examined empirical and simulated estimates of canopy light attenuation at SETRES (Southeast Tree Research and Education Site), a 2×2 factorial study of water and nutrients. Fertilized plots had significantly lower under-canopy PAR transmittance (T_C) when compared to non-fertilized plots. Light interception efficiency, as measured by the canopy cosine-corrected light extinction coefficient, G, was significantly lower in irrigated plots for all dates examined. Estimates of G ranged from a low of 0.36 in irrigated plots in September to a high of 0.64 in March for control plots. Study-wide analyses indicate that a G of 0.50 and a k (uncorrected light extinction coefficient) of 0.69 may be reasonable parameter estimates of canopy light extinction in intermediate-aged loblolly pine plantations across a range of stand conditions and seasons when site-specific data are unavailable. Simulated T_C from our version of the BIOMASS model corresponded well to the empirical estimates. Varying the vertical distribution of foliage in simulations (from 10:60:30 to 40:40:10% in the upper, middle, and lower canopy positions, respectively) resulted in only a ±7% change in total PAR intercepted, whereas varying G from 0.3 to 0.7 resulted in a 67% and 31% increase in light intercepted for control and fertilized plots, respectively. Decreased G resulted in an increased proportion of beam radiation intercepted – 63–67% of total PAR intercepted – by the middle canopy where 55–60% of the foliage was found. We hypothesize that proportionally increased productivity observed in irrigated treatments may be attributed to increased beam radiation intercepted deeper into the canopy by a greater foliage area. Received: 23 April 1998 / Accepted: 17 June 1998     

Comparative mapping in loblolly and radiata pine using RFLP and microsatellite markers

Genetic linkage maps were constructed for loblolly pine (Pinus taeda L.) and radiata pine (P. radiata D. Don) using a common set of RFLP and microsatellite markers. The map for loblolly pine combined data from two full-sib families and consisted of 20 linkage groups covering 1281 cM. The map for radiata pine had 14 linkage groups and covered 1223 cM. All of the RFLP probes readily hybridise between loblolly and radiata pine often producing similar hybridisation patterns. There were in total 60 homologous RFLP loci mapped in both species which could be used for comparative purposes. A set of 20 microsatellite markers derived from radiata pine were also assayed; however, only 9 amplified and revealed polymorphic loci in both species. Single-locus RFLP and microsatellite markers were used to match up linkage groups and compare order between species. Twelve syntenic groups were obtained each consisting of from 3 to 9 homologous loci. The order of homologous loci was colinear in most cases, suggesting no major chromosomal rearrangements in the evolution of these species. Comparative mapping between loblolly and radiata pine should facilitate genetic research in both species and provide a framework for mapping in other pine species. Received: 25 November 1998 / /Accepted: 19 December 1998     

Average effect of a mutation in lignin biosynthesis in loblolly pine

Cinnamyl alcohol dehydrogenase (CAD, E.C. 1.1.1.195) is a monolignol biosynthetic enzyme that catalyzes the final step of lignin subunit biosynthesis in higher plants. Recently, a mutant allele of the cad gene, cad-n1, encoding for the CAD enzyme, was discovered in loblolly pine. By reducing the expression of the cad gene, this mutant has a decreased lignin content and major changes in the lignin composition in wood. In this study, we found that the substitution of a wild-type allele by cad-n1 was associated with a significant effect on 2nd-year shoot elongation in a half-sib family of loblolly pine (designated family 7–1037). The average effect of cad-n1 appeared to increase with tree growth and was greater for stem radial growth than height growth. An increase of 14.1% in de-barked volume in year 4 was associated with cad-n1. Co-segregation analysis indicated that the cad locus itself might represent a gene that governs stem growth in pine. The significance of the mutation cad-n1 for tree growth and wood processing is discussed. Received: 31 December 1998 / Accepted: 30 January 1999     

Condition of Scots pine fine roots and mycorrhiza after fungicide application and low-level ozone exposure in a 2-year field experiment

 In a 2-year experiment at an open-air ozone fumigation field, the effects of fungicide application and low-level ozone exposure, single and combined, on fine root and mycorrhiza condition of Scots pine (Pinus sylvestris) seedlings were studied. Two different fungicides, copper oxychloride and propiconazole, with different modes of actions, were used. Propiconazole treatment reduced mycorrhizal infection in both years while copper oxychloride treatment and ozone exposure slightly stimulated mycorrhizal infection after the first year. Different mycorrhizal morphotypes showed different kinds of responses to the two fungicides. Light brown morphotype appeared to be the most sensitive one to propiconazole treatment. After the second year, ectendomycorrhizas disappeared in propiconazole treatment while in control treatment ectendomycorrhizas formed the majority of the light brown morphotype. The root biomass was not affected by fungicide treatments, but ozone exposure increased the total amount of short roots and the fresh weight of propiconazole treated roots. No significant differences in the concentrations of ergosterol, starch and total phenolics in pine roots between treatments were found. However, ergosterol concentration correlated positively with the mycorrhizal infection level. Both fungicides reduced the soil respiration compared to controls. At the ultrastructural level, both fungicides caused increased transparency and gradual granulation and degeneration of cytoplasm in the fungal symbiont of mycorrhizal short roots. Slightly elevated ozone did not have harmful effects on root ultrastructure. These results suggest that fungicides have deleterious effects on the quantity and quality of mycorrhizas in Scots pine roots and also side-effects on non-target soil fungi. Some of these deleterious effects were noticeable only at the ultrastructural level. Received: 23 June 1997 / Accepted: 11 December 1997     

Purification and cloning of an arabinogalactan-protein from xylem of loblolly pine

 An arabinogalactan-protein (AGP) was purified from differentiating xylem of loblolly pine (Pinus taeda L.) and the N-terminal sequence used to identify a cDNA clone. The protein, PtaAGP3, was not coded for by any previously identified AGP-like genes. Moreover, PtaAGP3 was abundantly and preferentially expressed in differentiating xylem. The encoded protein contains four domains, a signal peptide, a cleaved hydrophilic region, a region rich in serine, alanine, and proline/hydroxyproline, and a hydrophobic C-terminus. It is postulated to contain a GPI (glycosylphosphatidylinositol) anchor site. If the protein is cleaved at the putative GPI anchor site, as has been observed in other classical AGPs, all but the Ser-Ala-Pro/Hyp-rich domain may be missing from the mature protein. Xylem-specific AGPs are hypothesized to be involved in xylem development. Received: 29 July 1999 / Accepted: 19 August 1999     

Seasonal development of loblolly pine lateral roots in response to stand density and fertilization

In 1989, two levels each of stand density and fertilization treatments were factorially established in a 9-year-old loblolly pine plantation on a P-deficient Gulf Coastal Plain site in Rapides Parish, Louisiana, USA. In 1995, a second thinning was conducted on the previously thinned plots and fertilizer was re-applied to the previously fertilized plots. The morphology of new long lateral roots was evaluated at 2-week intervals in five Plexiglas rhizotrons per plot of two replications. The overall objective of this study was to evaluate the seasonal initiation of six morphological categories of long lateral roots ( 2.5 cm in length) in response to stand density and fertilization. Lateral root development exhibited a seasonal pattern with the initiation of branched lateral roots predominantly occurring in spring and summer. The initiation of non-branched lateral roots occurred throughout the year regardless of season. Stand density did not affect lateral root morphological development. However, fertilization stimulated the initiation of branched lateral roots that were greater than 1 mm in diameter.     

Identification of quantitative trait loci influencing annual height- and diameter-increment growth in loblolly pine (Pinus taeda L.)

 Quantitative trait loci (QTLs) for annual height- and diameter-increment growth were estimated in each of two three-generation loblolly pine pedigrees. Annual height-increment was measured in three consecutive years in the BASE pedigree and in four consecutive years in the QTL pedigree. Annual diameter-increment was measured only in the QTL pedigree for two consecutive years. An interval mapping-approach was used to estimate the number of QTLs, the magnitude of QTL effects, and their position on genetic linkage maps. Thirteen different height-increment and eight different diameter-increment QTLs were detected, suggesting that the these traits are, at least in part, controlled by a few genes of large effect. Little evidence was found for the expression of individual QTLs in multiple years or in multiple genetic backgrounds. These results were discussed in terms of the power of the experiment and their consequences for marker-assisted breeding. Received: 12 July 1998/Accepted: 5 August 1998     

Genotype by environment interaction for index traits that combine growth and wood density in loblolly pine

 Genotype×environment interactions of individual traits have been assessed in numerous experiments with forest trees. However, since breeding programs rarely aim at the improvement of a single trait, the impact of G×E on index or composite traits must also be assessed. In a study with 12-year-old loblolly pine families in the southeastern U.S., G×E variance was of relatively little importance compared to genetic variance for wood density but was of greater significance for several growth traits. An index that combined stem volume and wood density to improve dry weight but maintain wood density constant (restricted selection index) resulted in substantially greater G×E variance compared to either of the component traits. The interaction variance of an index trait is shown to be a function of the index coefficients and the G×E variances and covariances for its constituent traits. As a result, for some conditions it surpasses the magnitude of G×E variance for each component trait. Received: 5 September 1996 / Accepted: 25 October 1996     

Identification of QTLs influencing wood property traits in loblolly pine ( Pinus taeda L.). II. Chemical wood properties

Chemical wood property traits were analyzed for the presence of quantitative trait loci (QTLs) in a three-generation outbred pedigree of loblolly pine (Pinus taeda L.). These traits were assayed using pyrolysis molecular beam mass spectrometry and include mass spectrum peak intensities associated with carbohydrates, α-cellulose and hemicellulose sugars, and lignin. Models for projection to latent structures (PLS) were used to also estimate the chemical composition of cell walls (i.e., α-cellulose, galactan and lignin) from mass spectrum data using multivariate regression. Both earlywood and latewood fractions from the fifth annual ring were analyzed for each trait. An interval mapping approach designed for an outbred pedigree was used to estimate the number of QTLs, the magnitude of QTL effects, and their genomic position. Eight unique QTLs influencing cell wall chemistry were detected from multiple peak intensities and/or PLS estimates using the one- and two-QTL models. Significant differences in chemical contents were observed among the populations from North Carolina vs Oklahoma, and results from QTL×environment analyses suggest that QTLs interact with environmental location. QTLs should be verified in larger experiments and in different genetic and environmental backgrounds. QTL mapping will help towards eventually identifying genes having a major effect on chemical wood properties. Received: 19 January 2001 / Accepted: 31 May 2001     

Morphological and anatomical characterisation of black alder Alnus glutinosa (L.) Gaertn. ectomycorrhizas

 Ectomycorrhizal types of black alder [Alnus glutinosa (L.) Gaertn.] collected over a 3-year period within an alder forest were characterised by morphological and anatomical features. Of the total of 16 types, 14 are described for the first time in this paper. Eight identified types belong to the genera Russula, Lactarius, Naucoria, and Cortinarius, while eight further types remained unidentified. In some cases, similarities of mantle features indicate relationships to identified mycorrhizas. Mycorrhizas of Naucoria escharoides and N. subconspersa were not distinguished. Two unidentified mycorrhizal types exhibited hyphal mantle structures very similar to these Naucoria species. Within the genus Cortinarius, mycorrhizas of C. cf. helvelloides were easily distinguished from all other Cortinarius-like mycorrhizas described on Alnus, which in general showed little anatomical variation. Two further unidentified mycorrhizas, "Alnirhiza lilacina" and "A. violacea", probably also belong to Cortinarius. The ectomycorrhiza of Russula pumila was the only identified type within the genus Russula, but the unidentified type "Alnirhiza cremicolor" also likely belongs to this genus. Three Lactarius species were present in the experimental plot. Two species (L. obscuratus and L. omphaliformis) had indistinguishable mycorrhizal types, but were easily differentiated from the mycorrhizas of L. lilacinus, which caused intracellular penetration of Hartig net hyphae into epidermal and cortical cells. All other mycorrhizal types of black alder exhibited a paraepidermal Hartig net without penetration of root cells. Two unidentified mycorrhizal types "Alnirhiza atroverrucosa" and "A. cystidiobrunnea", already described from North American Alnus rubra as unnamed morphotypes, showed no similarity to identified mycorrhizas. All 16 mycorrhizal types appeared to be specific or at least typical for alders, since they have not yet been reported from other tree species. Accepted: 29 August 1997     

A procedure for the establishment of Glomus mosseae in dual culture with Ri T-DNA-transformed carrot roots

 A stepwise procedure was investigated to determine the optimal conditions for the establishment of Glomus mosseae (Nicol. & Gerd.) Gerdemann & Trappe in dual in vitro culture with Ri T-DNA-transformed roots of Daucus carota L. Glomus mosseae spores germinated best in 10 mm Tris or MES-buffered medium at pH values just above neutral. Growth of hyphae from germinated spores was much greater in the presence of Tris than MES, eg. 8 mm versus 4 mm per spore for Tris and MES, respectively, at pH 7.2. Roots exhibited a broad pH optimum for growth of 6.0–7.0 in both MES and Tris, but did not grow well above pH 7.5. In addition, purified gelling agent, gellan gum, was utilized to lower the P concentration of media. With these factors combined, mycorrhizas were successfully established in 14% of dual cultures. Accepted: 5 March 1997     

Climate-tree-growth relationships of Scots pine stands (Pinus sylvestris L.) exposed to soil dryness

 Dendroclimatological techniques were used to assess the impact of climatic factors on radial tree growth (total ring-width and latewood-width) of stunted Scots pine trees (Pinus sylvestris L.) exposed to soil dryness and nutrient deficiency on a dolomite substrate. The response of eight scattered populations representing various habitats, yet influenced by the same regional climate was investigated. Total ring-width and latewood-width were dated, standardized and several chronology statistics, which estimate the chronology signal strength and the potential climate signal in the series, were determined. Dendroclimatic analysis comprised evaluation of event/pointer years as well as response function elements. Response function analysis indicates that at most sites wide rings are significantly associated with high precipitation in April to June and cool conditions in May of the current year, and high precipitation in August to September of the preceding year. Latewood chronologies show a lower climate signal at all sites. Limited water availability causes a homogeneous relationship of radial tree growth to climate at all habitats, though site characteristics (slope magnitude, slope aspect, soil depth, vegetation cover) differ substantially. Cluster analysis of negative event/pointer years suggests that within the study area stands react to extreme climatic events depending on susceptibility to soil dryness, which is primarily determined by site topography. Received: 5 January 1998 / Accepted: 22 April 1998     

Phosphate uptake and polyphosphate metabolism of mycorrhizal and nonmycorrhizal roots of pine and of Suillus bovinus at varying external pH measured by in vivo 31P-NMR

 Comparative in vivo ³¹P-NMR analyses of mycorrhizal and nonmycorrhizal roots of Pinus sylvestris and the fungus of Suillus bovinus in pure culture were used to investigate alterations in phosphate metabolism due to changes in external pH in the range 3.5–8.5. All control samples maintained a constant pH in both cytoplasm and vacuole. Mycorrhizal roots and pure fungus, but not nonmycorrhizal roots, transformed accumulated inorganic phosphate into mobile polyphosphate with a medium chain length. Phosphate uptake rates and polyphosphate accumulation responded differently to external pH. In all cases, maximal phosphate uptake occurred at an external pH close to 5.5. At an external pH of 8.5, both roots and fungus showed a distinct lag in phosphate uptake, which was abolished when the external pH was lowered to 7.5. An irreversible effect on phosphate uptake as a consequence of variation in external pH was also observed. The central role of the fungus in regulating mycorrhizal phosphate metabolism is discussed. Accepted: 15 April 1997     

Segregation and linkage relationships of isoenzymes in shortleaf pine (Pinus echinata Mill.)

 Segregation and linkage relationships were analyzed between 28 isoenzyme loci in ten natural stands representing much of the natural range of Pinus echinata Mill. (shortleaf pine). A total of 203 possible two-locus combinations were tested. Three linkage groups were revealed in this study at a linkLOD of 4.0. The first linkage group (A) consisted of Pgi and Adh-1; Gdh, Idh, Skdh-2, G6pd-2 and Aco were mapped to the second linkage group (B); the third group (C) had 2 loci: Mdh-2 and Mdh-3. A moderate linkage between Mnr-2 and Dia-2 and weak linkages between Mnr-1 and Dia-1, and Got-2 and 6pgd-2 were also detected. The significance of these results in shortleaf pine is discussed and compared to linkage maps previously reported in other conifers, including pines. Received: 7 April 1997 / Accepted: 25 April 1997     

Nutrient uptake by intact and disturbed roots of loblolly pine seedlings

Most measurements of nutrient uptake use either hydroponic systems or soil-grown roots that have been disturbed by excavation. The first objective of this study was to test how root excavation affects nitrate uptake. Rates of NO₃^? uptake by mycorrhizal loblolly pine (Pinus taeda L.) seedlings were measured in intact sand-filled columns, hydroponics, and disturbed sand-filled columns. Total nitrate uptake in intact sand-filled columns was higher than in disturbed columns, indicating that disturbance lowers uptake. Transferring plants from the sand-filled columns to hydroponics had little effect on NO₃^? uptake beyond delaying uptake for an hour. The second objective of this study was to determine whether NH₄⁺, Ca²⁺, Mg²⁺ and K⁺ uptake could be studied using sand-filled columns, since previous studies had tested this method only for nitrate uptake. Uptake rates of NH₄⁺ and K⁺ were positive, while Ca²⁺ and Mg²⁺ uptake rates were negative in intact sand-filled columns, indicating that net efflux may occur even without physical disturbance to the root system. The sand-filled column approach has some limitations, but holds promise for conducting nutrient uptake studies with minimal disturbance to the root system.     

Identification of QTLs influencing wood property traits in loblolly pine (Pinus taeda L.). I. Physical wood properties

Physical wood property traits for loblolly pine (Pinus taeda L.) were analyzed for the presence of quantitative trait loci (QTLs) in a three-generation outbred pedigree. These traits include wood specific gravity (wsg), volume percentage of latewood (vol%), and microfibril angle (mfa). Phenotypic data were collected for rings 2–11 for wsg and vol%, and rings 3, 5 and 7 for mfa. Both earlywood and latewood were analyzed for each trait. An interval mapping approach designed for an outbred pedigree was used to estimate the number of QTLs, the magnitude of QTL effects, and their genomic position. Nine unique QTLs were detected for composite traits (average of all rings) for wsg, five for vol%, and five for mfa. The majority of these QTLs were verified by analyses of individual-ring traits. Additional QTLs for each trait were also detected by these individual-ring analyses. Most QTLs for wsg were specific to either earlywood or latewood, whereas each QTL for mfa was detected for both earlywood and latewood. Before these QTLs are utilized in a breeding program, they should be verified in larger experiments and in different genetic and environmental backgrounds. Received: 15 February 2000 / Accepted: 31 March 2000     

High-efficiency induction of soybean hairy roots and propagation of the soybean cyst nematode

Cotyledon explants of 10 soybean [Glycine max (L.) Merr.] cultivars were inoculated with Agrobacterium rhizogenes strain K599 with and without binary vectors pBI121 or pBINm-gfp5-ER possessing both neomycin phosphotransferase II (nptII) and β-glucuronidase (gus) or nptII and green fluorescent protein (gfp) genes, respectively. Hairy roots were produced from the wounded surface of 54–95% of the cotyledon explants on MXB selective medium containing 200 μg ml⁻¹ kanamycin and 500 μg ml⁻¹ carbenicillin. Putative individual transformed hairy roots were identified by cucumopine analysis and were screened for transgene incorporation using polymerase chain reaction. All of the roots tested were found to be co-transformed with T-DNA from the Ri-plasmid and the transgene from the binary vectors. Southern blot analysis confirmed the presence of the 35S-gfp5 gene in the plant genomes. Transgene expression was also confirmed by histochemical GUS assay and Western blot analysis for the GFP. Attempts to induce shoot formation from the hairy roots failed. Infection of hairy roots of the soybean cyst nematode (Heterodera glycines Ichinohe)-susceptible cultivar, Williams 82, with eggs of H. glycines race 1, resulted in the development of mature cysts about 4–5 weeks after inoculation. Thus the soybean cyst nematode could complete its entire life cycle in transformed soybean hairy-root cultures expressing GFP. This system should be ideal for testing genes that might impart resistance to soybean cyst nematode. Received: 13 July 1999 / Accepted: 8 August 1999     

Correlation between infection by ophiostomatoid fungi and the presence of subterranean termites in loblolly pine (Pinus taeda L.) roots

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