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Proceedings of the Indiana Academy of Science

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Proceedings of the Indiana Academy of Science Analysis of Some Natural and Artificial Interspecific Hybrids in Helianthus 1 R. C. Jacksons and A. T. Guard The importance of natural hybridization and subsequent introgres- sion as a source of raw material for evolution has been stressed by Anderson (1949) and other workers in this particular field. Heiser (1947, 1949, 1951a, b) has demonstrated that Helianthus annuus L, the common sunflower, hybridizes naturally with other annual species. Long (1955a, b) and Jackson (1955) have shown that morphologically similar and dissimilar perennial species of Helianthus may hybridize readily. This present study was initiated after observation of a large number of herbarium specimens indicated that Helianthus mollis Lam. was hybridizing naturally with several other perennial taxa. Descriptions of some of these hybrids have been reported in a previous paper (Jackson and Guard, in press). The purpose of this paper is to present an analysis of certain artificial and natural hybrids centered about Helianthus mollis. Methods and Materials Mass collections of the species and hybrid were made from various localities in Indiana. Collections of each species usually consisted of 25 individual plants from a single population. From 25 to 50 hybrid plants per population were collected, the number being dependent upon the size of the population and the variability present. The sources of the natural hybrids and species used in this study are given in table 1. Material for cytological study was collected from naturally occurring species and hybrids and from artificial hybrids and the parental species grown in the greenhouse. Immature heads were fixed in a mixture of two parts absolute ethyl alcohol and one part glacial acetic acid. After 48 hours, the microsporocytes were squashed directly in iron- acetocarmine. Usable slides were made permanent by the introduction of Venetian turpentine. Pollen grains were stained with 5 percent cotton blue in Aman's Lactophenol. Only those pollen grains with the cyto- plasm stained in a uniform deep blue after 48 hours were considered viable. Artificial crossing was accomplished by rubbing the heads of the plants together during anthesis of the disc flowers. Plants were bagged before and after crossing in order to insure against pollen contamination from other plants. 1 A portion of the field work for this study was made possible by a grant from the Indiana Academy of Science. 2 Present address: Dept. of Biology, University of New Mexico. 306 Plant Taxonomy 307 TABLE 1 Sources of the Species and Hybrids Used in This Study* Species or Hybrid Number Collector Locality H. mollis p72 C. B. Heiser N. St. Louis, Mo. H. mollis pl71 H. litis Rootsfork, Ark. H. mollis 716 R. C. Jackson Tippecanoe Co., Ind. H. grosseserratus p54 M. Ownbey Ft. Smelling, Minn. H. grosseserratus 717 R. C. Jackson Pulaski Co., Ind. H. giganteus p3250 C. B. Heiser Camp Douglas, Wis. H. giganteus 726 R. C. Jackson Starke Co., Ind. H. divaricatus 727 R. C. Jackson Starke Co., Ind. H. atrorubens p44 Baldwin Virginia H. mollis X 0.8 mi. N. of Chalmers, H. grosseserratus 700 R. C. Jackson White Co., Ind. H. mollis X 0.4 mi. N. Bass Station, H. giganteus 706 R. C. Jackson Starke Co., Ind. H. mollis X 1 mi. S. of San Pierre, H. giganteus 710 R. C. Jackson Starke Co., Ind. H. mollis X Jasper-Pulaski State H. divaricatus 714 R. C. Jackson Game Preserve, Pulaski Co., Ind. H. mollis X 1 mi. S. of San Pierre, H. divaricatus 715 R. C. Jackson Starke Co., Ind. 1 All plant numbers preceded by p represent collections kindly furnished to the writers by Dr. C. B. Heiser, Jr., Indiana University. Cytological Observations The species studied here all have the reported haploid chromosome number of seventeen. These counts were verified during this study. Meiotic stages beginning with diakinesis and containing through anaphase II were studied in the species and hybrids. Observations on pairing were made exclusively at diakinesis since this was the only stage where the chromosome pairs could be spread out sufficiently for detailed study. As a measure of pairing in the earlier stages, the mean minimum chiasma frequency at diakinesis was used. Closed bivalents were con- sidered as being held together by a single chiasma at either end while open bivalents were treated as being held together by a single chisma. Most of the closed bivalents were circular in shape and the open bivalents were held together at one end. No univalents were observed in any of the hybrids or species reported here. The chiasma frequency and pollen fertility of the species and hybrids are given in table 2. Standard error was calculated for chisma frequency. 308 Indiana Academy of Science TABLE 2 Mean Minimum Chiasma (Xa.) Frequency Per Cell and Pollen Fertility of the Species and Artificial Hybridsi No. of % Stain- Species and Hybrids cells Xa. frequency able studied range x s Pollen H. mollis (p72) |P 20-29 28.7 + 1.8 97.0 H. mollis (pl71) 50 29-26 22.5 + 1.5 98.9 H. grosses erratus (p54) 50 21-28 23.8+2.0 99.7 H. divaricatus (727) 50 20-26 22.6+1.2 100.0 H. giganteus (p3250) 50 20-30 22.0+1.8 99.4 H. giganteus (726) 50 20-28 23.5 + 1.7 99.7 H. atrorubens (p44) 50 29-32 31.0 + 1.0 92.4 H. grosseserratus (p54) X H. mollis (p72) 50 20-27 22.4+2.0 94.4 H. mollis X H. divaricatus (727) 50 19-24 20.8 + 1.5 49.4 H. giganteus (p3250) X. H. mollis (p72) 50 20-26 22.9 + 1.6 59.7 H. mollis (pl71) X H. atrorubens (p44) 72 17-22 18.9 + 1.5 34.0 K^ytological data for the hybrid between H. mollis and H. divaricatus are from a natural hybrid. The artificial hybrid has not yet been produced. Cytology of the parental species: Meiosis was normal in H. mollis, H. divaricatus L., and H. grossesarratus Martens. In material from two of five plants of H. giganteus L. from a natural population a single chromatin bridge was observed in 5 of 25 cells at first telophase, but in plant p3250 used in the artificial crosses meiosis was normal. In H. atrorubens L. (p44) atypical meiotic behavior was noted. At early diakinesis chains of four chromosomes were seen in some cells. Frag- ments were observed at late diakinesis, metaphase I and anaphase I. A single chromatin bridge sometimes accompanied by a fragment was noted in 5 of 54 cells studied at first anaphase. Cytology of the hybrids: Several natural hybrids between H. mollis and H. divaricatus were studied since the artificial hybrid has not yet been produced. In one plant a chain of four chromosomes was observed at diakinesis. No other aberrations were noted. In the artificial hybrid between H. mollis (p72) and H. grosseserratus (p54) all phases of meiosis were normal. The aberrations observed in H. giganteus (p3250) X H. mollis (p72) have been reported previously (Jackson, 1956). Briefly, fragments were found at early diakinesis and at first metaphase. A few single chromatin bridges were noted, and in one cell a double bridge was observed. Generally bridges were infrequent; apparently they break in early anaphase because the remnants of what appeared to be broken bridges were seen in a number of cells. The hybrid H. mollis (pl71) X H. atrorubens (p44) exhibited fragments at metaphase Plant Taxonomy 309 I, and a single chromatin bridge was observed at anaphases I and II. In several cells the bridge at first anaphase was accompanied by a fragment. Analysis of Populations Species and natural hybrids from which population analysis were made were collected in Indiana during the summer and fall of 1953 and 1954. Analysis of artificial hybrids and backcrosses were made from plants grown in the greenhouses at Purdue University. Measure- ments of all plants were taken after they were pressed and dried. The variation of morphological characters of the species and hybrids have been shown in histogram form using the Hybrid Index Method devised by Anderson (1936). It is a convenient way of graphically show- ing the differences within and between populations which are not easily grasped otherwise. The shortcomings of this method have been dealt with by Baker (1947), who criticized its use without adequate genetic data, by Heiser (1949) and by Anderson himself (1949). The method has been used extensively for studies of natural hybridization. In many of these studies artificial hybrids were not produced and the weighting of index values was arbitrary. In this study all but one of the Fi hybrids were produced, and a number of backcross progeny were grown and studied. By examination of the histograms in figure 8 it can be seen that Fi hybrid between H. mollis and H. oceidentalis reported elsewhere (Jackson and Guard, in press). Helianthus giganteus X H. mollis was an exception. This hybrid (fig. 4) resembled the pistillate parent more than the pollen parent. Thus, in examining histograms of natural hybrid populations, one should always refer to the position of the Fi hybrid scored by the same index. Variation to the right or left of the Fi may be taken as an indication that introgression has occurred in a particular direction. Parental species populations: Several populations of each species were sampled, but only one for each species has been used for the histograms presented in this section. Variation between populations in which hybridization had occurred was generally slight, and could most easily be attributed to edaphic or physiographic factors. Although most of the species discussed here are found in northern Indiana, they do not always grow in the same habitat.
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