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58 THE BRITISH SMUT FUNGI Ustilago hordei (Pers.) Lagerh. Covered Smut of Barley and Oats [Beticularia segetum BuUiard, 1791, P-p.] Uredo segetum subsp. hordei Persoon, Synopsis, p. 224, 1801. Uredo carbo de Candolle, 1815 [nov. nom. for U. segetum] p.p. Ustilago segetum (Pers.) Ditmar, 1817 [as ' U. segetum Link'] p.p. Ustilago carbo (DC.) Tulasne, 1847, p.p. Ustilago hordei (Pers.) Lagerheim, Mitt, badischen bat. Ver., p. 70 (March) 1889.' Ustilago avenae var. levis KeUerman & Swingle, 1890. Ustilago kolleri WiUe, 1893. Ustilago levis (Kellerm. & Swing.) Magnus, 1894. Sori in the spikelets replacing the ovaries and more or less of the tissues within the glumes. Spore mass firm, brown- or purpUsh-black, partly (or rarely completely) hidden by the glumes. Spores spherical to subspherical, pale yellow or greenish-brown, lighter in colour on one side than the other, smooth, 7-11 (av. 8-5-9-0) fj, diam. On barley (Hordeum) and oats (Avena) causing Covered Smut. July-Sept. Widespread. Common. Spore germination has been studied by Fischer von Waldheim (1869), Kellerman & Swingle (1890), Brefeld (1895), Herzberg (1895), McAlpine (1910), Paravicini (1917), and others (see Liro, 1924). Stakman (1913), using a-race from barley, found that some spores germinated on water in 6^ hours and nearly all in 24 hours. The growth of promycelia and sporidia follows the same plan as U. avenae (see p. 61). Infection of the host occurs during germination from spores lying on the surface of the grain or between the pales and the caryopsis. See p. 43. Racial specialization. Jacqzewski (1925) reported on the natural occurrence of U. hordei on rye in Siberia, and in the United States it occurs on Agropyron cristatum and Elymus glaucus jepsoni. Fischer (1939 a), using paired monosporidial cultures of covered smut from these hosts, produced infection on two varieties of barley and on the following grasses: Agropyron caninum, E. canadensis, E. glaucus jepsoni, E. sibiricus, Hordeum nodosum, and Sitanion jubatum. This race was physiologically distinct from covered smut of oats. Faris (1924 b) gives a table showing the infection of four differential varieties of barley by five physiologic races. Rodenhiser (1928), dealing mainly with cultural races, found that two differed also in pathogenicity. Aamodt & Johnston (1935) reported on two physiologic races at Alberta. Semeniuk (1940) in the same State detected four physiologic races in 1935-7 with an unexplained change in pathogenicity in 1938. Allison (1937) found that 27 out of 28 collections of covered smut could be differentiated on six varieties of barley, the type of infection varying with the race. Tapke (1937 b) distinguished eight races (including those of Faris) on five varieties of spring barley. Odessa was susceptible to all races. Later, 13 races were differentiated on eight varieties (Tapke, 1945, Table 1). New races were obtained by screening certain collections. Apart from pathogenicity, races differed in the size of colour of the chlamydospores, relative smoothness of the walls, degree of compactness of the heads, and mode
THE BRITISH SMUT FUKGI 59 of exsertion. The most promising variety for breeding for resistance was Pannier C. 1. 1330. A change in pathogenicity attributed to hybridization resulted from the inoculation of Odessa with a mixture of two races of smut (Tapke, 1944). UstUago vaillantii Tul. Ustilago vaillantii Tulasne, Ann. Sci. not., Bot., Ser. 3, vii, p. 90, 1847. Sori in the anthers and, less frequently, the ovaries. Spore mass powdery, brownish-black. Spores globose, irregularly globose, or somewhat elongated, pale greenish-yeUow, smooth or slightly granular, 6-12 X 6-9 /x. On Chionodoxal uciliae, Muscari botryoides, M. cyaneo-violaceum, Scilla bifolia, S. verna. April. Widespread. Spore germination. Figures of germination made by Schroeter (1877), Brefeld (1883), Schellenberg (1911), Massee (1914), and Davie & Wilson (1914) suggest a similarity between this species and U. longissima (see p. 56). According to Schroeter (1877) a long eUiptical cell (16-18x3-5^ yu) arises on a short stem (3-5 X 2 /i) from which it is soon released; subsequently it becomes septate and cuts off sporidia directly or on short sterigmata. Additional sporidia (sometimes 12 X 3, usually 4-6 X 2 /n) may develop from the promycelium (Fig. 2 e). Spores retain their viability for at least three months after being dried (Massee, 1914). Paravicini (1917) confirmed this method of germination in material from Scilla bifolia, observed fusions between sporidia, and figured nuclei. Infection of the host. The fungus is systemic and passes from the parent to newly formed bulbs. It wUl also infect young Scilla seedlings (Massee, 1914). Racial specialization. Ciferri (1938) distinguishes the form on S. bifolia as U. scillae Cif. Ustilago grandis Fr. Reed Smut Ustilago grandis Fries, Systema, iii, p. 518, 1832. Erysibe typhoides Wallroth, 1833, fide de Toni in Sacc. Syll., 1888. Ustilago typhoides (Wallr.) Berkeley & Broome, 1850 [Notices of British Fungi, No. 480]. Sori in the culms as raised, brown, longitudinal streaks, sometimes completely surrounding the culm and extending from one node to the next, at first covered by the epidermis (Plate I, Fig. 4). Spore mass powdery, brownish-black, weathering away to leave the culm bare. Spores globose or somewhat elongated, pale brown, smooth (or, under an oil immersion objective, granular) 10-12 X 7-10 ;x. On Phragmites communis. July-Oct. Cambridgeshire [Herb. Kew]; Norfolk [Herb. I.M.I. 17263] Spore germination. Kiihn (1877) described germination, noting a tendency for the promycelia to separate from the spore before producing sporidia. Brefeld (1883), who found that spores would germinate in autumn and remain viable
Page 1 and 2:
THE BRITISH SMUT FUNGI ^ (USTILAGIN
Page 3:
FOREWORD THIS volume was started wh
Page 6 and 7:
6 PBEFACE We also acknowledge the b
Page 9 and 10: INTRODUCTION THE smut fungi, which
Page 11 and 12: INTRODUCTION H of bushels of wheat
Page 14 and 15: FIG. 1. Vstilago avenae on Arrhenat
Page 16: ^^ THE BBITISH SMUT FUNGI Young hea
Page 19 and 20: BIOLOGY 15 less than the normal, in
Page 21 and 22: BIOLOGy 17 and united with it, fina
Page 23 and 24: BIOLOGY 19 eermination in a collect
Page 25 and 26: BIOLOGY 21 genus, and the manner of
Page 27 and 28: BIOLOGY 23 species of Entyloma, and
Page 29 and 30: 3036^ BIOLOGY 25 in others. Dickins
Page 31 and 32: CYTOLOGY THE discovery of nuclei in
Page 33 and 34: GENETICS INCOMPATIBILITY KNIEP'S di
Page 35 and 36: GENETICS • 31 direction of growth
Page 37 and 38: GENETICS 35 when a few sporidia fro
Page 39 and 40: GENETICS 37 lines retained their di
Page 41 and 42: TECHNIQUE COLLECTION AND EXAMINATIO
Page 43 and 44: TECHNIQUE 41 be undesirable. Weak a
Page 45 and 46: TECHNIQUE 43 in one season (see p.
Page 47 and 48: TECHNIQUE 45 barley with loose smut
Page 49 and 50: TECHNIQUE 47 (1939), Holton & Heald
Page 51 and 52: TECHNIQUE 49 Allen's, modification
Page 53 and 54: CLASSIFICATION THE morphological ch
Page 55 and 56: THE BRITISH SMUT FUNGI MOST of the
Page 57 and 58: FIG. 2. Spore geimination in Ustila
Page 59: THE BRITISH SMUT FUJJGI 57 from one
Page 63 and 64: THE BRITISH SMUT FUNGI 61 noticeabl
Page 65 and 66: THE BRITISH SMUT FUNGI 63 Evidence
Page 67 and 68: THE BRITISH SMUT FUNGI 65 winter in
Page 69 and 70: THE BRITISH SMUT FUNGI 67 Ustilago
Page 71 and 72: THE BRITISH SMUT FUlJGI 69 material
Page 73 and 74: THE BRITISH SMUT FUNGI 71 results o
Page 75 and 76: THE BRITISH SMUT FUNGI 73 TTstilago
Page 77 and 78: THE BRITISH SMUT FUNGI 75 Species i
Page 79 and 80: " TJSE BRITISH SMUT FUNGI 77 Ustila
Page 81 and 82: THE BKITISH SMUT FUNGI 79 On Carex
Page 83 and 84: THE BRITISH SMUT FUNGI 81 Exsiccati
Page 85 and 86: THE BRITISH SMUT FUNGI 83 Tilletia
Page 87 and 88: THE BRITISH SMUT FUNGI 85 bunt ball
Page 89 and 90: THE BEITISH SMUT FUNGI 87 globose t
Page 91 and 92: THE BRITISH SMUT FUNGI 89 FIG. 13.
Page 93 and 94: FIG. 15. Spore geimination in Taboi
Page 95 and 96: FIG. 17. Spore germination in Urocy
Page 97 and 98: THE BRITISH SMUT FUNGI 95 separate
Page 99 and 100: THE BRITISH SMUT FUNGI 97 8ori in t
Page 101 and 102: THE BRITISH SMUT FUNGI 99 Urocystis
Page 103 and 104: 30390 THE BRITISH SMUT FUNGI 101 pr
Page 105 and 106: FIG. 20. Spore geimination in Entyl
Page 107 and 108: THE BRITISH SMUT FUNGI 105 Sori in
Page 109 and 110: On Ranunculus ficariae and B. scler
Page 111 and 112:
THE BRITISH SMUT FUNGI 109 Infectio
Page 113 and 114:
THE BKITISH SMUT FUNGI 111 [Sporidi
Page 115 and 116:
THE BRITISH SMUT FUNGI 113 Ustilago
Page 117 and 118:
REFEEENCES 115 BBETT, M. A. (1940).
Page 119 and 120:
REFERENCES 117 ^ DniON WESTON, W. A
Page 121 and 122:
REFERENCES 119 GRIFFITHS, MAEIOK A.
Page 123 and 124:
REFERENCES 121 HtjTTiG, W. (1933).
Page 125 and 126:
REFEBENCES 123 LiMBOtruN, E. J. (19
Page 127 and 128:
REFERENCES 125 NOBLE, E. J. (1934).
Page 129 and 130:
REFERENCES 127 RoDENHiSEE, H. A., &
Page 131 and 132:
BEFERENCES 129 STAKMAK, E. C, XCEEN
Page 133 and 134:
EEFEEENCES 131 WAXTBE, J. M. (1934)
Page 135 and 136:
INDEX to generic and specific names
Page 137 and 138:
Muscari Ustilago vaillantii, 59 Myo
Page 139 and 140:
Ustilago ficuum Reich., 113 —flps
Page 141 and 142:
2i}:y0 ANDHRA PRADESH AGRICULTURAL
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