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Biochemical and Morphological Analyses of the Hordeum Jubatum-Brachyantherum-Caespitosum Hybrid Complex

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Biochemical and Morphological Analyses of the Hordeum Jubatum-Brachyantherum-Caespitosum Hybrid Complex Brigham Young University BYU ScholarsArchive Theses and Dissertations 1969-06-01 Biochemical and morphological analyses of the hordeum jubatum-brachyantherum-caespitosum hybrid complex Gareth Roz Babbel Brigham Young University - Provo Follow this and additional works at: https://scholarsarchive.byu.edu/etd BYU ScholarsArchive Citation Babbel, Gareth Roz, "Biochemical and morphological analyses of the hordeum jubatum-brachyantherum- caespitosum hybrid complex" (1969). Theses and Dissertations. 8017. https://scholarsarchive.byu.edu/etd/8017 This Thesis is brought to you for free and open access by BYU ScholarsArchive. It has been accepted for inclusion in Theses and Dissertations by an authorized administrator of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. BIOCHEMICALAND MORPHOLOGICAL ANALYSES OF THE HORDEUM JUBATUM-BRACHYANTHERUM-OAESPITOSUMHYBRIDCOMPLEX A Thesis Presented to the Department of Botany Brigham Young University In Partial Fulfillment of the Requirements for the Degree Master of Science by Gareth R. Babbel June 1969 111 .A.OICNOWLEDGMENTS Grateful acknowledgment is made for the valuable sug- gestions, criticisms, and encouragement given by the chair- man of myadVisory committee, Dr. Howard o. Stutz, who sug- gested this problem. The valuable assistance of other members of my adv1sory committee, Dr. Fred G. White and Dr. Dayna Stocks, 1s appreciated. !he Department of Botany, Brigham Young University, prov1ded laboratory space, equipment, and supplies. Finally, my family, especially my Wife Ann, provided encouragement which aided immensely in the completion of this project. iv TABLEOF CONTENTS Page ACKNOWLEDGMENTS• • • • • • • • • • • • • • • • • 111 LIST OF TABLES••• • • • • • • • • • • • • • • • V LIST OF ILLUSTRATIONS• • • • • • • • • • • • • • Vi INTRODUCTION• • • • • • • •r-. • ,, • 1 \{ti cPt) t/.J,.llt5> • • • • • • • • MATERIALSAlID METHODS/ _f • • • • • • • • • • • • • 9 RESULTS • • • • • • • • • • • • • • • • • • • • • 14 DISOUSSIOM• • • • • • • • • • • • • • • • • • • • 37 SUMMARY • • • • • • • • • • • • • • • • • • • • • 52 :REFERENCES• • • • • • • • • • • • • • • • • • • • 54 V LIST OF TABLES Table Page 1. Summary of Seeds Set and Germination in Vari- ous Artificial Crosses in Hordeum. Small letters: j.·= !!• jubatum, br. = £!• brachy;an- therum, n.h. = natural hybrid ••••••• 15 Seed Set from Plants Growing in the Green- house Including Some Suspected Natural and Artificial Hybrids •••••••• • • 19 3 •. Summary of Chromatographic Data. The Diagnos- tic Substances Represent the Observed Spots 1n the Spot Patterns. Clearly Visible Spots = X,. Faint Spots = :x:. Compilation of the Hybrid Index 1 s Explained in the Text and in Table 4 •••••••••••••• 22 4. Summary of Diagnostic Characters and Assigned Values Used in Constructing Scatter Dia- grams and Hybrid Indices ••••••••• 27 5. Standard Deviations and Coefficients of Vari- ation in Lemm.a-awnLength and Anther Size for Individuals and for Populations of Hordeum in Idaho and Utah ••••••••• 34 Vi LIST OF ILLUSTRATIONS Figure Page 1. Typical Variation Found in ll• jubatp and H• brachyantherum in Utah Collection Sites. • 2 2. Normal (1.) and Aborted seeds (r.) from an Arti- ficial Cross between H. brachyantherum (Utah) and H• jubatum-(Utah). Ca. 6x ••• 16 3. Spikes of Artificial Hybrids and Parent Plants. Spike of Female Parent is to the Left of the Hybrid in each Case •••••••••• 18 4. Line Drawings of Representative Chromatograms as Seen with UV Light and Ammonia Vapors. A. Gb-170 H. brachyantherum (Utah), B. Gb-18 g. jubatum (Utah), C. Gb-18x170 Ar- tificial Hybrid, D. Gb-154 Natural Hybrid (Utah), E. Gb-46 li• Brachyantherum (Idaho), F. Gb-120 H• .JJ:tbatum7Idaho), G. Gb-17 · g. Jubatum TUtah) ••••••••••••• 23, 24 5. Comparison of the Range of Variation in Mor- phological Characters Scored. a= g. ju~- ~' Utah; b = H• jubatum, Idaho; c = arti- ficial and natural hybrids, Utah; d = H• brachyantherum, Utah; e = g. bracnyantherum, Idaho. Vertical Lines Encompass the Total Range of Variability, Horizontal Lines Denote the Mean • • • ••• • • • • • • • • • 26 6. Spikelets of H. brachyantherum (1.) and H. jubatum (r.). Note the Relative Differ- ences in Size of the Rachis Internode, Palea, and Lemma+ Awn of the Lateral Floret. x8. 27 7. Comparison of Anther Size. H. brachyantherum (1.), ~• jubatum (r.). x25 •••••••• 27 a. Pictorialized Scatter Diagram of a Mixed Population of H. jubatum, ~. brac~yan- therum., and Natural and Artificial tiybrids from Utah. (For Explanation of Figures see Table 4.) ••••••••• • • 31 Page Pictorialized Scatter Diagram of a ~axed Population of li• ubatum and li• brachy- antherum from Idaho. For Explanation of Figures see Table 4.) ••••••• ... 33 10. Normal li• jubatum (1.) and Morphological Variant (r.) all from Utah ••••• • • • 11. Comparison of Lemma-awn Length and F..ybrid Index in Hordeum Specimens from Utah. Symbols: O = li• jubatum, e = Artificial Hybrids, Q = Natural Hybrids, and O :::: H• brachyantherum • • • • • • • • • • • • 41 12. Comparison of Lemma-awn Length and Hybrid Index in Hordeum Specimens from Idaho. Symbols: O = !!• jubatum, • = ,g. brachyantherum. • • • • • • • • • • • • • 43 INTRODUCTION fhe Wild barley species, Hordeum jubatum L. and Rordeum prachyantherum nevsk:1, are common throughout western North A:inerioa. Although intermediate forms are usually found where these two species grow sympatric~lly, there are some locations where the 'bro remain sharply discrete, even though growing side b;y side. In this study an attempt was made to determine if natural by'bridization W1th subsequent introgression occurs between these species. Three populations were selected for study. Of these three populations, one near Heber, Utah and one near Provo, Utah are quite similar. l!• jubatum, !!• braohyanther1.1m and intermediates grow together at both sites~ Warm summers and cold Winters are characteristic of these two areas, al- 1 though Reber (6,ooo') is coo~er than Provo (4,300 }. The cli- mate of both of these two areas coupled with intensive agri- culture and grazing provides a great variety of available habi- tats for l!• jubatum and the intermediate forms (Fig. 1). !he third population which was selected for study is located in the Sawtooth Valley of Idaho. In this population !!• brachyantherum and l!• jubatum also grow side by side but no intermediate forms are apparent. The Sawtooth Valley of 1 Idaho 1s a high mountain valley (6,500 - 7,000 } with very cold Winters and cool summ·ers. Al though sheep and cattle have 2 Fig. 1. Typical variation found in li• jubatum and li• brachyantherum in Utah collection sites. grazed the valley since 1900 it is only sparsely inhabited by humans. A lack of intensive agriculture together with a very cold climate apparently provide only a narrow range of habitats for Hordeum species. H. jubatum and li• brachyantherum are both perennial wild grasses with reduced pedicillate lateral flowers (Hitch­ cock, 1950). As shown in Fig. 1, they can however, be easily distinguished morphologically. The awns and glumes of H. jubatum are very long (40 - 70 mm.), while those of li• brachy­ antherum are quite short (6 - 15 mm.). On the other hand, the palea (6.4 - 9.5 mm.) and culms (30 - 100 cm.) of li• brachy­ antherum are longer than those of H. jubatum (4.8 - 7.0 mm. and 30 - 60 cm.). Also, as reported by Mitchell and Wilton (1964), the internode on the rachis of H. brachyantherum is longer than that of li• jubatum. li• jubatum and li• brach,yantherum differ markedly in their ecological requirements. g. jubatum grows well on 3 saline soils in waste places and meadows at lower elevations, while!!• brachyantherutl_! grows primarily in moist meadows and 1 grassy slopes at higher elevations up to 9,000 • When the two species grow side by side, g. ~chyantherum is always found in the more mois•t portion of their commonhabitat. Although morphologically and ecologically distinct,~. juba~ and g. brachyantherum appear to be quite closely re- lated. They both have a chromosome number of 2N = 28 {Rajhathy and Morrison; Bowden, 1962), although Morrison (1959) has re- ported occasional specimens of!!• Jubatum which have a chromo- some number of 2N = 14. Intermediate forms found where these two species grow sympatrically have morphological characteristics intermediate between those of both species. Descriptions of such interme- diates have been given by Hitchcock (1950), Anderson (1959), Mitchell and Wilton (1964), and others. The length of the awns (15 - 35 mm.) is especially helpful in detecting inter- mediate forms in the field. Most authors consider intermediate forms to be natur- a1 hybrids between !!• juba.tum and !!• brachyantherum. Oova.s (1949) stated that it (the intermediate) "• •• seems to be an entity intermediate between!!• jubatum and!!• braohyan- theru.m." He postulates that this "entity" is probably the result of hybridization occurring when~. jubatum and!!• brachyantherum come in contact. In his Flora of Ala,ska, An- derson (1959) designates the intermediate. as a separate spe- cies,~- caespitosum, but also recognizes that it is a probable 4 hy'brid. Others who suggest that the intermediate 1s a natural hy'brid include Rajhathy and Morrison (1959), Bowden (1962), and Mitchell and Wilton (1964). The possibility of natural hybridization between!!• J!:!,- batum and li• prachyantherum was further suggested when Rajhathy and Morr1·son (1959) :found good cross-compatibility betwe~n the two species. They were able to produce F1 hybrids
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