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STEPHANOMERIA EXIGUA Ssp. CORONARIA THAT FLUCTUATES in POPULATION SIZE1

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STEPHANOMERIA EXIGUA Ssp. CORONARIA THAT FLUCTUATES in POPULATION SIZE1 GENETIC STABILITY IN A PERIPHERAL ISOLATE OF STEPHANOMERIA EXIGUA ssp. CORONARIA THAT FLUCTUATES IN POPULATION SIZE1 L. D. GOTTLIEB Department of Genetics, University of California, Davis 95616 Manuscript received September 5, 1973 ABSTRACT Allelic frequencies did not change at five polymorphic loci in seedlings grown from seeds collected in four consecutive years in a geographically peripheral population of the annual plant Stephanomeria ezigua ssp. coronaria (Compositae) , even though the population number fluctuated during this period by 50:i. Genetic stability is attributed to buffering effects provided by seed storage in the ground. Evidence described elsewhere suggests that this population was the recent progenitor of a new diploid species. The present re- sult indicates that fluctuations in number of individuals in the parental popula- tion were probably not involved in the origin of the new species. ISOLATED populations on the geographical periphery of widespread species often undergo drastic and unpredictable expansions and contractions in num- bers of individuals. Such fluctuations have been considered to set the stage for the establishment of novel genetic features and for speciation (CARSON1968; LEWIS1962, 1973). Evidence has been presented elsewhere (GOTTLIEB1973a) that a peripheral isolate of the annual plant Stephanomeria exigua ssp. coronaria (Compositae) in eastern Oregon was the recent progenitor of a new diploid species with which it is still sympatric. The derivative species, presently called “Malheurensis,” is self-pollinating and very similar morphologically to ssp. coronaria. Electrophoretic analysis demonstrated that its gene pool is a highly limited extraction of the alleles present in the progenitor population. The progen- itor population (as well as the derivative) experiences considerable yearly fluctu- ations in number of individuals in response to certain climatic factors, particu- larly the amount and timing of precipitation. Consequently, it was worthwhile to determine whether these fluctuations affect its allelic frequencies and could have been implicated in the origin of “Malheurensis”. The environmental conditions required for seed germination and seedling survival are often not satisfied at the study site in many years because of insuffi- cient and irregular rainfall. However, since seeds remain viable for long periods of time (following storage in the laboratory, 10-20% of field-collected seeds are viable after five years), it is likely that the soil at the study site contains a pool of seeds contributed over many generations, with the mating group of any one Supported by National Science Foundation Grant GB 29484X. Genetics 76: 551-556 March, 1974. 552 L. D. GOTTLIEB year comprised of individuals produced in different generations. The' data pre- sented in this paper document that the genetic input into this seed pool is stable from year to year regardless of the number of individuals in the mating population. MATERIALS AND METHODS Stephanomeria exigua ssp. coronaria is encountered in a diverse series of habitats and plant communities from eastern Oregon and western Idaho south to the islands off the California coast near Santa Barbara (GOTTLIEB1971). The subspecies is diploid, has annual generations, and possesses a sporophytic self-incompatibility system making it an obligate outcrosser. The popu- lation described in this report is the most northern one known in the subspecies; it occupies a site approximately 50 acres in extent, located 25 miles south of Burns, Harney County, Oregon. The seeds of the study population must be exposed to cold temperatures before they can germinate (GOTTLIEB1973a). Consequently, seeds presumably germinate in early spring, and the seedlings grow initially as rosettes. After several months, the rosettes send up a central stem which develops side branches that bear the flowering heads. Flowering begins by early July and seeds are produced until the plants die in early September. In additiosn to exposure to low temperatures, an adequate precipitation in the spring is re- quired for germination and seedling survival; but in eastern Oregon rainfall is highly unpre- dictable. Climatalogical data, available from the U.S. Weather Bureau Station at the nearby Malheur National Wildlife Refuge Headquarters, show that April, May and June, 1971, were wet with a total of 3.13 inches; the adult population in that year numbered approximately 25,000 individuals. The same months in 1972 were much drier, with a total of 1.06 inches, and the population numbered fewer than 500 individuals, a 50: 1 reduction. Average precipitation (35 years of data) for these three months is 2.64 inches. Census estimates were not made for earlier years except that following the dry spring (1.78 inches) of 1968, fewer than 100 individuals were observed. When precipitation is nearer to average values, my observations suggest that the mating group probably approximates 5000 individuals. Seeds collected from individuals growing in the field in 1969, 1970, 1971 and 1972 were available for study. The seeds were taken from plants sampled at ten-foot intervals along transects over the entire extent of the population. After germination in the laboratory, the seedlings were assayed by horizontal starch gel electrophoresis for three loci controlling glutamate oxaloacetate transaminases (GOT) and two controlling esterases (EST). Techniques are described in GOTTLIEB (1973a). These five loci were selected because they are highly polymorphic and their enzyme patterns can be unmistakenly determined. Formal genetic analysis o'f the electrophoretic patterns of the transminases has been reported (GOTTLIEB1973b), and the genetics of the esterases will be reported elsewhere. Twelve additional loci responsible for variation in electrophoretic mobility of enzymes, of which three are polymorphic by the five percent criterion, have been identified in this population (GOTTLIEB1973a and unpublished). The patterns governed by these three loci are not consistently readable and they are not included in this study. The data presented here for 1971 differs in some respects from that reported previously (GOTTLIEB1973a) which also was based on seeds of 1971. EST-I and EST-2 are now designated EST-2 and EST-3, respectively, since a third locus controlling more anodal esterases has been detected, and allele 43 is now called allele 44. The small differences in allelic frequencies between the two reports reflect the smaller sample size previously studied, differences in observed frequencies of rare alleles, and the improved resolu- tion obtained in the more recent gels. RESULTS AND DISCUSSION The data inTable 1 demonstrate that even though the population of S. ezigua ssp. coronaria numbered approximately 5000 individuals in 1969 and 1970, flushed to 25,000 in 1971, and crashed to fewer than 500 in 1972, allelic frequen- cies at all loci remained remarkably stable. The chi-square test of independence GENETIC STABILITY IN STEPHANOMERIA 553 TABLE 3 Allelic frequencies obserued for five loci in seedlings grown from seed dlected in each of four years in a peripheral populution of Stephanomeria exigua ssp. coronaria Year Locus and allele 1969 (12,16) 1970 (14,25) 1971 (43,137) 1972 (52,125) GOT-1-5 1 0.31 0.24 0.18 0.20 -54 0.69 0.76 0.82 0.80 GOT-2-31 - 0.02 0.01 0.02 -38 1.00 0.98 0.99 0.98 GOT-3- 8 0.03 - 0.01 0.01 -1 1 0.13 0.19 0.16 0.19 -1 9 0.84 0.81 0.83 0.80 EST-- - - - <O.Ol -51 0.10 0.02 0.03 0.06 -54 0.57 0.60 0.41 0.41 -56 0.30 0.26 0.47 0.47 -58 0.03 0.05 0.W 0.02 -61 - 0.02 - - -null* - 0.02 0.W 0.03 EST-3-38 - - - 0.01 -46) 0.43 0.33 0.33 0.37 -42 - 0.11 0.01 0.05 -44 0.47 0.47 0.47 0.51 -46 0.10 0.09 0.19 0.06 The numbers in the parentheses are the average number of plant families sampled and the total number of individuals in these families. The alleles are identified according to the migration in mm of their enzyme products. * Frequency of homozygous individuals only. calculated on the basis of observed numbers of alleles showed no significant changes over years: for GOT-I, x;= 3.88, P < 0.25; for GOT-2, x:= 1.93, P < 0.50; for GOT-3, xi= 5.01, P < 0.50. For the more highly polymorphic esterase loci, the calculations were made using the two most frequent alleles and pooling the additional alleles in low frequency into a single class: for EST-2, xi = 10.97, P < 0.05; and for EST-3, x: = 7.59, P < 0.25. The small fluctuations which occurred may reflect sampling errors attributable to the much smaller number of seeds available for the 1969 and 1970 samples, or possible differences in seed viability (not seedling survival since following germination seedling survival was uniformly high for all four age classes). The important observation is that regardless of the size of the mating group in the different years the allelic frequencies of their progenies fluctuated with a very narrow amplitude and that even rare alleles were not lost. Comparison of observed and expected numbers of heterozygous individuals suggests that the population is mating at random (Table 2). Only EST-2 showed a significant heterozygote deficiency but, since genetic analysis has identified 5 54 L. D. GOTTLIEB TABLE 2 Proportion and number of observed and expected individuals heterozygous for five loci in seedlings grown from seed collected in four consecutive years in a population of Stephanomeria exigua ssp. coronaria Year Locus 1969 1970 1971 1972 GOT-I ob) 0.25(4) 0.41(11) 0.26 (35) 0.23 (29) * ex) 0.43 (6.9) 0.36(9.7) 0.30( 41 .I ) 0.32(40.6) GOT-2 ob) 0.00 0.04(1) 0.02(3) 0.05 (6) ex) 0.00 om(1.1) 0.012(2.7) 0.04 (5.0) GOT-3 ob) 0.31(5) 0.23 (6) 0.25 (37) 0.27 (34) ex) 0.28(4.5) 0.31(8.1) 0.29 (42.3) 0.32(40.3) EST-2 ob) 0.40(6) 0.4Q(9) 0.32(42)*** 0.30(38)*** ex) 0.57(9.1) 0.57( 11.9) 0.61 (81.1) 0.61 (77.5) EST-3 ob) 0.53(8) 0.48(11) 0.48(63)* 0.50(59) ex) 0.58(8.7) 0.65(15.0) 0.63 (82.5) 0.60(70.2) * Significant, P < 0 025, *** P < 0.001.
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