EFFECTS OF TREE AGE ON SECONDARY XYLEM AND PHLOEM ANATOMY IN STEMS OF GREAT BASIN BRISTLECONE PINE (PINUS LONGAEVA)

Xylem and phloem tissue samples were collected from various-aged Great Basin bristlecone pine (Pinus longaeva D. K. Bailey) stems in southern Utah and southeastern California to determine whether the vascular cambia of older trees produce fewer xylem rays, shorter-lived xylem and phloem ray cells, fewer phloem sieve cells, and a thinner phloem. Increment cores were examined to determine whether ‘aged’ cambia produced narrower tracheids that might reduce water translocation. Sapwood thickness was measured and sapwood growth layers were counted on these cores. Regression and Classification and Regression Tree (CART) analyses of sample data found no age-related changes in cambial products. Phloem and xylem production appeared normal at all ages, with no evidence of cambial malfunction.     

THE FATE OF THE DWARF SHOOT APEX IN BRISTLECONE PINE (PINUS LONGAEVA)

The fate of the pine dwarf shoot (DS) apex after needle initiation has been controversial. Dwarf shoot primordia of Pinus longaeva were examined to determine the developmental basis for DS with and without interfoliar buds. Interfoliar buds are microscopic buds derived from the original terminal apex of the DS. In October, all the DS primordia are similar in size and appearance. However, as the needles elongate in the following June the apices of more proximal DS decrease in size, such that by July there is a clear diminishing size gradient of apical domes in going from the most distal to the most proximal positions. The distal DSs start to form bud scales in July and have fully formed interfoliar buds by mid-August. In contrast, those DS apices lacking protective bud scales at needle maturity become suberized and can never proliferate into long shoots. The distal placement of interfoliar buds may be due to a group effect, where each developing DS inhibits the more proximal DSs in the long shoot terminal bud.     

SECONDARY GROWTH IN NEEDLE LEAVES OF PINUS LONGAEVA (BRISTLECONE PINE) AND OTHER CONIFERS: QUANTITATIVE DATA

Needle leaves of Pinus longaeva can be accurately dated and remain alive on branches for 30 or more yrs, making this species ideal to study secondary growth in leaves. Field observations and regression analysis of needle age versus mean needle length both indicate primary (elongation) growth of needles is completed in the first year. Statistical analysis of cell counts for one- to 33-yr-old needles indicate production along the entire length of needles of 1.0-1.7 cell layers per year of secondary phloem, but no secondary xylem. Microscopic measurements and cell counts reveal that with advancing needle age phloem increases radially and transfusion tissue buckles, but the number of endodermal cells and xylem width do not change. Living phloem remains constant in amount (ca. 9 layers) with advancing needle age, indicating yearly replacement of old by new phloem. For comparison to P. longaeva, needle leaves were also studied for ten other conifer taxa with maximum needle longevities ranging from 3 to 19 yrs. In needles of each taxon no secondary xylem is produced, but secondary phloem production occurs throughout the post-elongation lifespan of the needles regardless of maximum needle longevity.     

PATTERNS OF VARIATION IN LEVELS OF HOMOPLASY

Patterns of variation in levels of homoplasy were explored through statistical analyses of standardized consistency indexes. Data were obtained from 60 recent cladistic analyses of a wide variety of organisms based on several different kinds of characters. Consistency index is highly correlated with the number of taxa included in an analysis, with homoplasy increasing as the number of taxa increases. This observation is compatible with a simple model of character evolution in which 1) the probability of character-state change increases with the total number of branches in a tree and 2) the number of possible states of a character is limited. Consistency index does not show a significant relationship to the number of characters utilized in an analysis or to the taxonomic rank of the terminal taxa. When the relationship between consistency index and number of taxa is taken into account, there is no significant difference between plant and animal data sets in the amount of homoplasy. Likewise, the level of homoplasy in morphological and molecular data sets does not appear to differ significantly, although there are still too few molecular studies to be confident of this result. Future comparisons of consistency indexes, including studies along the lines established here, must take into account the influence of the number of taxa on homoplasy.     

MICROGEOGRAPHIC PATTERNS OF GENETIC AND MORPHOLOGICAL VARIATION IN SAVANNAH SPARROWS (PASSERCULUS SANDWICHENSIS)

Surveys of genetic population structure are often limited to large geographic scales because geographically close populations are indistinguishable. Genetic uniformity across adjacent demes can be interpreted as evidence for cohesion (panmixia) or recent divergence. However, poor genetic resolution at microgeographic scales can also arise from the use of overly conservative (slowly evolving) markers. This study examines the ability of hypervariable, minisatellite loci to discriminate among geographically close populations of Savannah sparrows (Passerculus sandwichensis) and to track morphological differentiation at a microgeographic scale (interregional distance < 55 km). Savannah sparrows breeding at five island and two mainland sites in the Bay of Fundy, New Brunswick, Canada, show concordant patterns of variation in external morphology (seven characters) and multilocus DNA fingerprinting profiles (S_xy): island sparrows are phenotypically larger and genetically more similar to each other than they are to mainland sparrows. This pattern of variation is consistent with both adaptive (natural selection) and nonadaptive (genetic drift) mechanisms of population divergence. Based on minisatellite diversity, the effective size of both island and mainland populations is 37, an estimate substantially lower than census population sizes. These data are discordant with observations of sparrow vagility and abundance and suggest a closer examination of microgeographic patterns in avian systems.     

GENETIC PATTERNS ASSOCIATED WITH GROWTH VARIABILITY IN PONDEROSA PINE

Variability in annual growth increment in Pinus ponderosa was measured and heterozygosity level estimated in order to examine the relationship between genetic heterozygosity and phenotypic homeostasis in natural populations of a long-lived perennial plant. Pollen was collected from 190 ponderosa pine trees in the upper montane region of the Front Range in Colorado and four isozyme systems were resolved electrophoretically from these samples. Each tree was cored, aged, and each annual ring width measured. Statistics estimating within-tree variation in growth increment were then related to heterozygosity levels. The results indicated a significant positive association between level of heterozygosity and growth variability. Two speculations are offered to account for this relationship.     

PATTERNS OF GENETIC VARIATION IN SALIX SECTION LONGIFOLIAE (SALICACEAE)

Enzyme electrophoresis was used to examine genetic relationships within a taxonomically complex group of diploid willows (Salix section Longifoliae). Forty-eight populations representing seven morphogeographic taxa were analyzed for 15 putative enzyme loci. Levels of genetic variation (A, P, H_e, and H_T) differed greatly among populations, but were generally lower than those reported for other species with similar life history traits. In populations and species from southern latitudes, where populations tend to be smaller and more isolated, genetic variation decreased and population differentiation increased. Genetic identity values suggest that the section is divisible into four major elements: 1) Salix interior, 2) S. taxifolia, 3) the S. exigua group, and 4) S. melanopsis. The first three occupy eastern, southern, and western portions of the continent, respectively. Salix melanopsis is sympatric with the S. exigua group in western North America, but is ecologically distinct. The S. exigua group is differentiated into four major geographic entities: S. hindsiana from west of the Sierras in California and southwest Oregon, S. sessilifolia from west of the Cascades in central Oregon to British Columbia, and a northern and southern race of S. exigua in the Intermountain West. The relationships suggested by allozyme data do not agree with traditional taxonomic concepts. Evidence of hybridization was obtained at five points where taxa come into contact, and Salix fluviatilis may be of hybrid origin. Taxonomic confusion appears to be largely the result of frequent hybridization events and overemphasis of a few highly variable morphological traits. The section appears to be a syngameon undergoing geographic speciation, with one taxon gaining reproductive isolation by a shift in habitat preference.     

MORPHOLOGICAL STUDIES OF THE ROOT OF RED PINE,PINUS RESINOSA I. GROWTH CHARACTERISTICS AND PATTERNS OF BRANCHING

Observations were made of the seasonal root growth behavior under natural conditions and under controlled conditions in plant observation boxes. Under natural conditions root growth conformed to the commonly reported pattern of a surge of growth in the spring, a mid-summer low, and a renewed burst in the fall. Growth of individual roots was cyclic. Growth patterns ordinarily varied according to root diameter and branching and in the plantations were modified by soil moisture conditions. Observations of roots during periods of constant elongation showed that the distance from the root apex to the first lateral root primordium varied directly with growth rate. Laterals did not arise in strict acropetal succession, and lateral root abortion was common, particularly in large-diameter, fast-growing roots. Observations of root initiation in relation to seasonal growth increments and to dormancy structures showed an increase in numbers of laterals on both the proximal and distal portions of a seasonal increment.     

凉水国家自然保护区天然红松林遗传变异的RAPD分析

采用随机扩增多态DNA(random amplified polymorphic DNA RAPD)技术,研究了凉水国家自然保护区天然红松林的遗传变异水平及其分布规律。16个10bp长度的随机引物在8个实验样地内72个个体中共检测了96个位点,其中55个是多态位点。在种水平上,红松的多态位点比率为P=58.51%;Nei's遗传变异指数为H=0.2868;Shannon's指数为I=0.3654,所有的变异中,有92.47%的变异存在于样地内, 7.53%的变异存在于样地间。根据各遗传多样性在不同样地的分布,初步探讨了红松遗传变异水平与其种群发生及生境的相互关系。     

FINE-SCALE GENETIC STRUCTURE OF WHITEBARK PINE (PINUS ALBICAULIS): ASSOCIATIONS WITH WATERSHED AND GROWTH FORM

The fine-scale genetic structure of a subalpine conifer, whitebark pine (Pinus albicaulis Engelm.), was studied at nested geographic levels from watershed to adjacent stems in the eastern Sierra Nevada Range of California. A combination of several characteristics contributed to unpredicted genetic structure in this species. This includes being one of only 20 pine species worldwide with wingless, bird-dispersed seeds; having the reputed capacity to reproduce vegetatively; and forming distinct growth morphologies at different elevations in this part of its natural range. Genetic differentiation, as measured with 21 allozyme loci, among the three studied watersheds is virtually negligible (F_ST = 0.004). This is a surprising result because the upper-elevation sites vary somewhat in slope aspect; thus, aspect was confounded with watershed effect. Differentiation between the upper-elevation prostrate krummholz thickets and lower-elevation upright tree clump growth forms is modest (F_ST = 0.051). Much stronger differentiation was measured among the individual thickets and clumps within their sample sites (F_ST = 0.334). Within krummholz thickets, multiple individuals are present and genetic relationships often resemble half- to full-sibling family structure (mean r = 0.320). Canonical trend surface analysis in two intensively sampled thickets indicates greatest genotypic variation in the direction of the prevailing wind. At lower elevations, most (72%) of the tree clumps contained more than one genotype; the remaining clumps are probably multistemmed trees. Within tree clumps, family relationships are closer than those for krummholz thickets—commonly full-sibling to selfed structure (mean r = 0.597). Genetic structure is apparently profoundly influenced by the seed-caching behavior of Clark's nutcracker (Nucifraga columbiana Wilson). Western pine species typically show little among-population differentiation and high levels of within-population genetic variation. In whitebark pine in the eastern Sierra Nevada of California, genetic variation is highly structured, especially within the natural groupings—krummholz thickets and upright tree clumps.     

云南松居群核型变异及其分化研究

在分析研究6个云南松居群核型变异的基础上,应用常规统计方法和巢式分析方法,在常规染 色体水平上探讨云南松居群间、个体间和细胞间的变异式样及其分化。结果如下:(1)云南松居群核型 变异不显著,6个居群的核型公式均为2n=24m(6～10SAT),核型类型均为1A。(2)云南松居群仅相 对长度系数、臂比和次缢痕数目及其分布有小的变化。6个居群染色体相对长度系数(I．R．L)分别为: 滇中居群P1=16M₂+6M₁+2S;滇东南居群P2=14M₂+8M₁+2S;滇西居群P3=12M₂+10M₁+2S; 地盘松居群P4=14M₂+8M₁+2S;细叶云南松居群P5=14M₂+8M₁+2S;云南松与思茅松渗入杂交 居群P6=10M₂+12M₁+2S。(3)巢式方差等级分析表明,云南松染色体结构变异有10％左右来源于居群间,有90％左右来源于居群内个体间或细胞间。     

GENETIC POPULATION STRUCTURE OF DROSOPHILA TRIPUNCTATA: PATTERNS OF VARIATION AND COVARIATION OF TRAITS AFFECTING RESOURCE USE

Drosophila tripunctata is an ecological generalist, using both fruits and mushrooms as breeding sites. Isofemale strains of this species were established from seven populations over a wide part of its range and assayed for electrophoretic variability, oviposition-site preference, and larval performance on several types of substrates. Significant variation among strains within populations was found for oviposition-site preference, larval development time on tomatoes versus mushrooms, and tolerance (as measured by development time) of the mushroom toxin α-amanitin. Even populations at the periphery of the range, which electrophoretic data suggest have been through bottlenecks, harbored levels of variation for oviposition-site preference approximately equal to that found in central populations. All correlations between preference and various measures of larval performance were close to zero. Thus, there is no evidence for sympatric divergence of host races or for coadapted complexes of genes related to host specificity. Strains with higher-than-average amanitin tolerance tended to develop more rapidly on tomatoes than on nontoxic mushrooms, whereas the less-tolerant strains had slower development on tomatoes. This suggests that there may be genetically based correlations and trade-offs in larval performance on different breeding sites. No geographic differentiation among populations was found for either oviposition-site preference or any measure of larval performance. There is also very little electrophoretic variation among populations. Thus, the species as a whole, rather than local populations, appears to be the unit of evolution with respect to resource use in D. tripunctata.