Genetic analysis of inbreeding depression in plus tree 850.55 of Pinus radiata D. Don. II. Genetics of viability genes

 Most coniferous species exhibit severe inbreeding depression. Selfed individuals usually have decreased viability, reduced vigour and morphological defects. The number of filled seeds after selfing Pinus radiata plus tree 850.55 was 48% that of the outcrossing, and 26.1% of the selfed seedlings died at an early stage. The segregation of 172 markers (covering 56% of the genome) in selfed progenies of radiata pine plus tree 850.55 was studied. Based on the segregation ratio of the markers, genes associated with inbreeding depression on viability were identified (P<0.05). Using the Expectation/Conditional Maximization (ECM) algorithm, we estimated the location, degree of dominance and selection coefficient of viability genes. Nine viability genes were discovered. Seven of them appeared to be dominant and one partially dominant (degree of dominance=0.4). The other gene was overdominant or pseudo-overdominant, with selection coefficients for the two homozygotes of 0.4 and 0.42, respectively. Of the genes showing dominance or partial dominance, seven were sub-lethal with selection coefficients ranging from 0.55 to 0.79; one gene (SDPr), which was responsible for seedling death within the first month following germination, was lethal. Received: 13 December 1998 / Accepted: 28 December 1998     

Genetic analysis of inbreeding depression in plus tree 850.55 of Pinus radiata D. Don. I. Genetic map with distorted markers

 A genetic map of Pinus radiata plus tree 850.55 was constructed using megagametophytes of S₁ seeds. The map contained 19 linkage groups, with 168 RAPD and four microsatellite markers. The total map length was 1116.7 cM (Kosambi’s function) and was estimated to cover 56% of the genome. Of the 172 markers, 59 (34%) were distorted from the expected 1 : 1 ratio in megagametophytes (P<0.05). We show that if the distortion is caused by a single viability gene or by sampling error, the estimate of recombination frequency in megagametophytes of selfed seeds would not be affected. Received: 20 April 1998 / Accepted: 13 July 1998     

Inbreeding in Pinus radiata. I. The effect of inbreeding on growth, survival and variance

 The effects of inbreeding on growth, survival and variance in a 12-year-old radiata pine trial were studied in five populations each inbred to one of five different levels: outcross (F=0), half-sib (F=0.125), full-sib (F=0.25), selfing (S₁, F=0.5), and two-generations of selfing (S₂, F=0.75). These five populations were derived from a founder population of eight clones. Inbreeding reduced diameter, growth, and survival but increased the variance for diameter. Inbreeding depression at F=0.125, 0.25, 0.5, and 0.75 was 5%, 6%, 15%, and 19% respectively for DBH; −3%, 1%, 7%, and 11% respectively, for survival. The standard deviation for diameter increased by 10%, 10%, 30%, and 25% respectively for F=0.125, 0.25, 0.5, and 0.75 and, similarly, the coefficient of variation increased by 17%, 16%, 53%, and 55% respectively. There were significant differences among the eight founder clones in their response to inbreeding. The best clone in the trial showed no inbreeding depression. Overall, inbreeding depression was found to be linearly related to the inbreeding coefficient F with no significant quadratic effects for any trait at any population level. However, two individual clones had a quadratic relationship with F for DBH and one clone had a similar relationship for survival. A significant correlation (r=0.96) between S₂ and the breeding values of founder clones was observed while the correlation (r=0.58) between S₁ and breeding values was insignificant. The low inbreeding depression in radiata pine relative to other conifers may indicate that historical purging of detrimental alleles through small geographic populations, a higher degree of population subdivision, and the relative high fecundity of inbred progenies has rendered radiata pine an ideal species to use inbreeding as a breeding tool. Received: 10 March 1998 / Accepted: 19 May 1998     

Evaluation of AFLP for genetic mapping in Pinus radiata D. Don

Efficient construction of reasonable density genetic linkage maps is an essential component of QTL detection programmes. The AFLP technique has been used to produce genetic linkage maps in a range of species. We have developed protocols to generate reproducible AFLP profiles in Pinus radiata and have evaluated the inheritance and informativeness of AFLP markers in this important timber species. The large genome size of P. radiata necessitated increased levels of selection at both the pre-amplification and selective amplification steps of the AFLP protocol to generate reproducible AFLP profiles. Once optimised ca. 41.3 scorable AFLP bands were resolvable through denaturing gels, of which 48.4% were polymorphic in a screen of eight unrelated trees. This level of polymorphism is ca. three times higher than with RAPD markers. The total number of bands and the number of polymorphismic bands per PCR were ca. halved when AFLPs were electrophoresed on non-denaturing gels and stained with ethidium bromide. Using the protocols developed, AFLP is an efficient method for generating the DNA markers required for genetic linkage map construction in P. radiata. This revised version was published online in June 2006 with corrections to the Cover Date.     

Stable transformation and regeneration of transgenic plants of Pinus radiata D. Don

A biolistic particle delivery system was used to genetically transform embryogenic tissue of Pinus radiata. The introduced DNA contained a uidA reporter gene under the control of either the tandem CaMV 35S or the artificial Emu promoter, and the npt II selectable marker controlled by the CaMV 35S promoter. The average number of stable, geneticin-resistant lines recovered was 0.5 per 200 mg fresh weight bombarded tissue. Expression of the uidA reporter gene was detected histochemically and fluorimetrically in transformed embryogenic tissue and in derived mature somatic embryos and regenerated plants. The integration of uidA and npt II genes into the Pinus radiata genome was demonstrated using PCR amplification of the inserts and Southern hybridisation analysis. The expression of both genes in transformed tissue was confirmed by Northern hybridisation analysis. More than 150 transgenic Pinus radiata plants were produced from 20 independent transformation experiments with four different embryogenic clones. Received: 9 May 1997 / Revision received: 18 September 1997 / Accepted: 18 October 1997     

Photosynthesis and wood structure in Pinus radiata D. Don during dehydration and immediately after rewatering

Abstract. Experiments were carried out on Pinus radiata (D. Don) trees grown as cuttings from clonal parent stock. Some of these trees were about 0.4 m high while others were about 5 m high; all were grown in containers. The stem diameters at the tops and at the bottoms of the large trees, rates of photosynthesis, and needle water potentials were measured both when the trees were well watered and as they dehydrated after water was withheld. The water potentials of well-watered plants was highest in the small trees and lowest at the top of the large trees. When water was withheld, photosynthesis was in most cases unaffected by a small reduction in water potential, but the rate of photosynthesis fell as water potentials declined further. The stems of the large trees expanded at a constant rate when the trees were well watered and for part of the dehydration period, while subsequent stem shrinkage and the fall in photosynthesis both occurred at approximately the same time.
Water potentials increased little in the 24 h after rewatering, and significant rates of photosynthesis were not measured until 2 or 3 d later while renewed stem expansion was not measured until 2 d after rewatering.
Water deficits reduced the lumen diameter of newly matured stem tracheids, but increased the thickness of their walls. After 1 month of water potentials of about −2.4 MPa, tracheid lumen diameter and wall thickness were both much reduced, and this reduction continued in tracheids maturing shortly after rewatering.     

The response of photosynthetic model parameters to temperature and nitrogen concentration in Pinus radiata D. Don

Responses of photosynthesis (A) to intercellular CO₂ concentration (c_i) in 2-year-old Pinus radiata D. Don seedlings were measured at a range of temperatures in order to parametrize a biophysical model of leaf photosynthesis. Increasing leaf temperature from 8 to 30°C caused a 4-fold increase in V_cmax, the maximum rate of carboxylation (10.7–43.3 μol m^?2 s^?1 and a 3-fold increase in J_max, the maximum electron transport rate (20.5–60.2 μmol m ^?2 s^?1). The temperature optimum for J_max was lower than that for V_cmax, causing a decline in the ratio J_max:V_cmax from 2.0 to 1.4 as leaf temperature increased from 8 to 30°C. To determine the response of photosynthesis to leaf nitrogen concentration, additional measurements were made on seedlings grown under four nitrogen treatments. Foliar N concentrations varied between 0.36 and 1.27 mol kg^?1, and there were linear relationships between N concentration and both V_cmax and J_max. Measurements made throughout the crown of a plantation forest tree, where foliar N concentrations varied from 0.83 mol kg^?1 near the base to 1.54 mol kg^?1 near the leader, yielded similar relationships. These results will be useful in scaling carbon assimilation models from leaves to canopies.     

Characteristics of single- and multi-copy microsatellites from Pinus radiata

 Dinucleotide microsatellites were isolated from Pinus radiata using both a standard genomic library and libraries enriched for microsatellites. Locus-specific primers were designed to amplify 43 unique microsatellites. Thirty two of these loci had interpretable PCR patterns, 11 of which were polymorphic in a screen of 19 P. radiata individuals; all 11 polymorphic loci contained at least 17 repeats in the sequenced plasmid. Six of the eleven primer pairs amplified multiple fragments per individual (3–8), suggesting that these loci were present in multiple copies in the genome. Genotyping a 48-tree P. radiata production population with seven of the most polymorphic microsatellites revealed an average of 17 bands per locus (the multi-copy microsatellites were treated as one locus). When tested on known pedigrees, both single and multi-copy microsatellites exhibited co-dominant inheritance and Mendelian segregation. Two loci had null alleles and one locus had a high frequency of non-parental alleles, suggesting a high mutation rate. Eight of these microsatellites, including five multi-copy loci, were placed on a partially constructed P. radiata genetic map. Four of the five multi-copy microsatellites had two or more sets of alleles that mapped to the same locus, and the fifth mapped to two unlinked loci. All seven tested primer pairs amplified PCR products from other species of hard pine, three amplified products from soft-pine species, and one amplified bands in other conifers. Received: 10 November 1997 / Accepted: 5 January 1998     

Competitive processes in a monoculture of Pinus radiata D. Don

Summary The changes between 9 and 40 years of age of the frequency distribution of tree diameters were studied in an experiment in a plantation of Pinus radiata D. Don in South Australia in which the spacing between trees at planting varied. Empirical functions were developed to relate tree diameter to growth rate in diameter, variance of growth rate in diameter and mortality rate. These functions were used in solving a forward diffusion equation to predict future diameter distributions. The contributions from each of the terms in the diffusion equation were examined and it was found that the drift (growth rate in diameter) term made the greatest contribution to the solution, whilst the diffusion (variance in growth rate) term had neglible effect. This implied that competition between individual trees for light was the dominant competitive process operating in this plantation, rather than below ground competition for soil resources. The ramifications of this finding to forest growth modelling are discussed.     

Response of transpiration and photosynthesis to a transient change in illuminated foliage area for a Pinus radiata D. Don tree

Sudden but transient changes in the fraction or illuminated foliage area in a well-watered 7-year-old Pinus radiata D. Don tree were imposed by completely covering either the upper 22% or the lower 78% of the foliage for periods of up to 36 h. Measurements of transpiration flux density (E), tree conductance (g_t), stomatal conductance (g_s) and net photosynthesis (A) were made to test the hypothesis that compensatory responses would occur in the remaining illuminated foliage when the cover was installed. When the lower foliage was covered there was an immediate decrease in g_t. However, when tree conductance was normalized with respect to the illuminated leaf area (g_t'), it increased between 50 and 75%, depending on the value of air saturation deficit (D). The effect was also apparent from concurrent measurements of increases in g_s and A up to 59 and 24%, respectively, for needles in the top third of (he crown. When the cover was removed these effects were reversed. The changes in the lower foliage when the upper foliage was covered were much smaller. Changes in bulk needle water potential were small. It is suggested that the observed responses occurred because of a perturbation to the hydraulic pathway in the xylem that could have triggered the action of a chemical signal to regulate stomatal conductance and photosynthesis.     

Nitrogen allocation and carbon isotope fractionation in relation to intercepted radiation and position in a young Pinus radiata D. Don tree

The three dimensional distribution of intercepted radiation, intercellular CO₂ concentration (C_i) and late summer needle nitrogen (N) concentration were determined at the tips of all 54 branches in a 6·2-m-tall Pinus radiata D. Don tree growing in a New Zealand plantation. Measurements included above- and below-canopy irradiance, leaf stable carbon isotopic composition (δ¹³C) and tree canopy architecture. The radiation absorption component of the model, MAESTRO, was tested on site and then used to determine the branch tip distribution of intercepted radiation. We hypothesized that in branch tip needles: (i) the allocation of nitrogen and other nutrients would be closely associated with the distribution of intercepted radiation, reflecting carbon gain optimization theory, and (ii) C_i would predominantly reflect changes in photosynthetic rate (A) rather than stomatal conductance (g_s), indicating that the increase in A for a given increase in N concentration was larger than the corresponding increase in g_s. Needle nitrogen concentration was poorly related to intercepted radiation, regardless of the period over which the latter was calculated. At a given height, there was a large azimuthal variation in intercepted radiation but N concentration was remarkably uniform around the tree canopy. There was, however, a linear and positive correspondence between N concentration and δ¹³C and needle height above ground (r² = 0·73 and 0·68, respectively). The very strong linear correspondence between N concentration and C_i (r² = 0·71) was interpreted, using gas exchange measurements, as supporting our second hypothesis. Recognizing the strong apical control in P. radiata and possible effects of leaf nitrogen storage in an evergreen species, we propose that the tree leader must have constituted a very strong carbon sink throughout the growing season, and that the proximity of branch tip needles to the leader affected their photosynthetic capacity and nutrient concentration, independent of intercepted radiation. This implies an integrated internal determination of resource allocation within the tree and challenges the current convention that resources are optimally distributed according to the profile of intercepted radiation.     

The influence of water deficits on needle conductance, assimilation rate and abscisic acid concentration of seedlings of Pinus radiata D. Don

Abstract Seedlings of Pinus radiata, 10–20 weeks old and hitherto fully watered, responded rapidly when water was withheld. Wilting occurred 9d later, at which time soil matric water potential at dawn (Ψ_m) was –1.06MPa and shoot water potential (Ψ) was –1.9 MPa. Small reductions in Ψ_m elicited large responses in assimilation rate (A) and leaf conductance to water vapour (g). Seedlings appear to be more sensitive to small water deficits than are older Plants of P. radiata. After rewatering, significant increases of A and g occurred within one day, but neither regained the values measured prior to the imposition of a single drying cycle. This residual effect of drought on A, after one or six drying cycles, was partially caused by a decrease in photosynthetic capacity. In plants wilted for the first time, the concentration of abscisic acid (ABA) in the bulk foliage increased 3.4 times as Ψ decreased to –1.77 MPa. In comparison, pretreatment with six drying cycles significantly reduced Ψ to –2.13 MPa (indicating some osmotic adjustment) and induced only a doubling of ABA concentration. However, these differences in Ψ and ABA concentration did not Persist after the plants of all pretreatments had been watered for 7 d, although g of drought-pretreatment Plants remained approximately half that of continuously-watered plants.     

The onset of photosynthetic acclimation to elevated CO2 partial pressure in field-grown Pinus radiata D. Don. after 4 years

The effects of CO₂ enrichment on photosynthesis and ribulose‐1,5‐bisphosphate carboxylase/oxygenase (rubisco) were studied in current year and 1‐year‐old needles of the same branch of field‐grown Pinus radiata D. Don trees. All measurements were made in the fourth year of growth in large, open‐top chambers continuously maintained at ambient (36 Pa) or elevated (65 Pa) CO₂ partial pressures. Photosynthetic rates of the 1‐year‐old needles made at the growth CO₂ partial pressure averaged 10·5 ± 0·5 μmol m^?2 s^?1 in the 36 Pa grown trees and 11·8 ± 0·4 μmol m^?2 s^?1 in the 65 Pa grown trees, and were not significantly different from each other. The photosynthetic capacity of 1‐year‐old needles was reduced by 25% from 23·0 ± 1·8 μmol m^?2 s^?1 in the 36 Pa CO₂ grown trees to 17·3 ± 0·7 μmol m^?2 s^?1 in the 65 Pa grown trees. Growth in elevated CO₂ also resulted in a 25% reduction in V_cmax (maximum carboxylation rate), a 23% reduction in J_max (RuBP regeneration capacity mediated by maximum electron transport rate) and a 30% reduction in Rubisco activity and content. Total non‐structural carbohydrates (TNC) as a fraction of total dry mass increased from 12·8 ± 0·4% in 1‐year‐old needles from the 36 Pa grown trees to 14·2 ± 0·7% in 1‐year‐old needles from the 65 Pa grown trees and leaf nitrogen content decreased from 1·30 ± 0·02 to 1·09 ± 0·10 g m^?2. The current‐year needles were not of sufficient size for gas exchange measurements, but none of the biochemical parameters measured (Rubisco, leaf chlorophyll, TNC and N), were effected by growth in elevated CO₂. These results demonstrate that photosynthetic acclimation, which was not found in the first 2 years of this experiment, can develop over time in field‐grown trees and may be regulated by source‐sink balance, sugar feedback mechanisms and nitrogen allocation.     

Growth and development of Pinus radiata D. Don: the effect of light quality

Abstract. Plants of Pinus radiata of two physiological ages, juvenile (seedlings and micropropagated plantlets) and adult (rooted cuttings from mature trees), were grown under lighting fromthree combinations of metal halide (MH) and tungsten halogen (TH) lamps for up to 10 months in controlled environment rooms. The three lamp combinations, MH alone, 50: 50 MH: TH and 25:75 MH: TH by wattage, produced red: farred ratios of 4.59, 1.51, and 1.15, respectively. Photosynthetic photon flux density was 700 μmol m⁻² s⁻¹. An increase in proportion of TH lamps markedly increased shoot elongation and internode length, decreased numbers of fascicles per unit stem length and increased the proportion of stem weight in both juvenile and adult material. In addition, in adult material, it increased the number of fascicle initials and expanded fascicles per growth flush, reduced the duration of the 'rest' phase between growth flushes, accelerated the rate of elongation growth during each flush, and increased apical dominance. Tracheid length, but not diameter or wall thickness, was significantly affected by light quality and found to be associated with longer internodes. Any treatment effects on needle weight or length, stem diameter or root weight were non-significant or very small. Different clones from either the juvenile micro-propagated material or the mature rooted cuttings each showed similar patterns of response, although they often differed in the degree of response to light quality. The main response could be related solely to the red: far-red ratio and the calculated phytochrome photoequilibrium. This is the first report of phytochrome-controlled photomorphogenesis in older specimens of a woody perennial. Recommendations for artificial light sources for growing P. radiata and some ecological implications of the results are presented.     

Pinus radiata D. Don in Chile. Fertilization trials with copper

Summary YoungPinus radiata D. Don were fertilized with copper oxychloride and copper sulfate on a random block basis with four replicates. Roots, needles, stems and shoot apices were analyzed separately for Cu, Fe, Mn and Zn contents at different times from fertilization. The results showed a rapid plant copper absorption, no effects being found on the Fe, Mn and Zn content. The dynamic internal distribution of copper among different plant parts is shown. The treatments applied did not cause any phenotypical alteration nor phytotoxicity in the plants, and no significant effect of the fertilizer chemicals on each other.     

Some observations on the spectral distribution characteristics of short-wave radiation within Pinus radiata D. Don canopies

Abstract. Measurements of the photosynthetic photon flux density (400–700 nm) and of the spectral distribution of photon flux density across the 370–800 nm waveband, were made under both clear and overcast sky conditions above and at various positions within two Pinus radiata canopies of different stocking but similar leaf area indices. The spectra obtained for the daylight conditions (i.e. above forest canopy) were generally similar to those published previously. The spectra for shadelight within the forest canopy showed no blue peak which was characteristic of previously reported measurements which were restricted to the diffuse radiation component. There was almost neutral absorption within the 400–700 nm waveband, and typical lower attenuation in the 700–800 nm waveband. The blue: red ratio was largely unchanged by either canopy type or sky conditions and varied between 0.57 and 0.81. The red: far-red ratio in shadelight was between 0.22 and 0.41 under clear sky and between 0.68 and 0.95 under overcast sky conditions. Values for daylight were between 1.16–1.22. Calculated phytochrome photoequilibrium values in shadelight were approximately 0.35 under clear sky and 0.46 under overcast sky conditions. In each case there appeared to be no differences between the two canopies with respect to these minimum values.     

Multiple-marker mapping of wood density loci in an outbred pedigree of radiata pine

The objective of this study was to determine the genetic location and effects of genomic regions controlling wood density at three stages, i.e., rings corresponding to ages 1–5 (WD1_5), rings corresponding to ages 6–10 (WD6_10), and outer wood density (WD14) in a full-sib pedigree (850.055×850.096) of Pinus radiata. The number of offspring measured at these three stages were 80, 93 and 93, respectively. Only a single linkage group of the parent 850.55 was considered for mapping quantitative trait loci (QTLs). A multiple-marker least-squares approach was employed for mapping QTLs for each of the three traits, using a single-QTL model. Logistic regression was used for multiple-trait QTL mapping. Critical values for test-statistic were calculated empirically by ’shuffling’ the data. A putative QTL with large effect on WD1_5 appears to be segregating at the 73 cM position (experimentwise P<0.01). The width of the 95% bootstrap confidence interval for this putative QTL was 40 cM (i.e. 56–96 cM). The effect of this QTL on the expression of wood density at later stages was diminished. From multiple-trait analysis, two marker locations (at 66 cM and 91 cM) were found to be significantly associated (experimentwise P <0.05) with the expression of wood density at different ages. These results are encouraging for the application of marker information to early selection in order to increase juvenile wood density, although the putative QTLs detected in this study need to be verified in an independent population. Received: 23 July 1999 / Accepted: 27 August 1999     

Photosynthetic acclimation to long-term exposure to elevated CO2 concentration in Pinus radiata D. Don. is related to age of needles

The effects of CO₂ enrichment on photosynthesis and ribulose-1,5-bisphosphate carboxylase/ oxygenase (Rubisco) in current year and 1-year-old needles on the same branch were studied on Pinus radiata D. Don. trees growing for 4 years in large, open-top chambers at ambient (36 Pa) and elevated (65 Pa) CO₂ partial pressures. At this age trees were 3·5–4 m tall. Measurements made late in the growing cycle (March) showed that photosynthetic rates at the growth CO₂ concentration [(pCO₂)_a] were lower in 1-year-old needles of trees grown at elevated CO₂ concentrations than in those of trees grown at ambient (pCO₂)_a. At elevated CO₂ concentrations V_cmax (maximum carboxylation rate) was reduced by 13% and J_max (RuBP regeneration capacity mediated by maximum electron transport rate) by 17%. This corresponded with photosynthetic rates at the growth (pCO₂)_a of 4·68 ± 0·41 μmol m^–2 s^–1 and 6·15 ± 0·46 μmol m^–2 s^–1 at 36 and 65 Pa, respectively (an enhancement of 31%). In current year needles photosynthetic rates at the growth (pCO₂)_a were 6·2 ± 0·72 μmol m^–2 s^–1 at 36 Pa and 10·15 ± 0·64 μmol m^–2 s^–1 at 65 Pa (an enhancement of 63%). The smaller enhancement of photosynthesis in 1-year-old needles at 65 Pa was accompanied by a reduction in Rubisco activity (39%) and content (40%) compared with that at 36 Pa. Starch and sugar concentrations in 1-year-old needles were not significantly different in the CO₂ treatments. There was no evidence in biochemical parameters for down-regulation at elevated (pCO₂)_a in fully fexpanded needles of the current year cohort. These data show that enhancement of photosynthesis continues to occur in needles after 4 years’ exposure to elevated CO₂ concentrations. Photosynthetic acclimation reduces the degree of this enhancement, but only in needles after 1 year of growth. Thus, responses to elevated CO₂ concentration change during the lifetime of needles, and acclimation may not be apparent in current year needles. This transitory effect is most probably attributable to the effects of developmental stage and proximity to actively growing shoots on sink strength for carbohydrates. The implications of such age-dependent responses are that older trees, in which the contribution of older needles to the photosynthetic biomass is greater than in younger trees, may become progressively more acclimated to elevated CO₂ concentration.     

Growth, dry weight partitioning and wood properties of Pinus radiata D. Don after 2 years of CO2 enrichment

Abstract Advanced selections (families 20010 and 20062) of P. radiata D. Don were exposed to either 340 or 660 μmol CO₂ mol ¹ for 2 years to establish if growth responses to high CO₂ would persist during the development of woody tissues. The experiment was carried out in glasshouses and some of the trees at each CO₂ concentration were subjected to phosphorus deficiency and to periodic drought. CO₂ enrichment increased whole-plant dry matter production irrespective of water availability, but only when phosphorus supply was adequate. The greatest increase occurred during the exponential period of growth and appeared to be tied to increased rates of photosynthesis, which caused accelerated production of leaf area. The increase in whole-plant dry matter production was similar for both families; however, family 20010 partitioned larger amounts of dry weight to the trunks than family 20062. which favoured the roots and branches. Wood density was generally increased by elevated CO₂ and for family 20010 this increase was due to thickening of the tracheid walls. Tracheid length was similar at both CO₂ levels but differed between families. These results suggest that, as the atmospheric CO₂ concentration rises, field-grown P. radiata should produce more dry weight at sites where phosphorus is not acutely deficient, even where drought limits growth; however, increases in wood production are likely only for genotypes which continue to partition at least the same proportion of dry weight to wood in the trunk.     

Segregation of allozymes in megagametophytes of viable seed from a natural population of jack pine,Pinus banksiana Lamb.

Summary Segregation ratios of allozymes in haploid female gametophytes obtained from viable seed were studied in a natural population of jack pine, Pinus banksiana. Stability of these ratios was assessed for three levels of the sexually reproductive crown as well as for four years of natural fertilization. Analyses of observed segregation ratios of four of five polymorphic isozyme loci showed good correspondence to the overall 11 ratios expected for simple Mendelian inheritance. Allozymes of glucose-6-phosphate dehydrogenase did not segregate in the expected 11 ratio. In addition, there were significant deviations from the expected segregation ratio for all the loci at some sampling positions on individual trees. Heterogeneity of segregation among trees, strata and years could be the result of pollen pool heterogeneity, segregation distortion and/or recessive lethal and semi-lethal gene combinations resulting in early embryo abortion. These types of segregation deviations in viable seed can affect the estimation of allele frequencies from bulked samples of a small number of individuals, the inference of heterozygosity/homozygosity of parental trees, and estimates of selfing rates.