The Inheritance of Tight and Loose Paleae in Avena Nuda Crosses
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VOLUME VII AUGUST, 1918 ){0. 4 THE INHEI~ITANCE OF TIGHT AND LOOSE PALEAE IN AVENA NUDA CROSSES ~. BY t. ST CLAIR CAPORN. (With six text-figures.) EXPEICIMENTS were conducted~ three varieties of ordinary oats with tight paleae, viz. :- Thousand Dollar. Ligowo. Nubiseher Sehwarzer (Nubian Black). As the N~ generation of the cross with Nubiseher Sehwarzer has not yet been harvested, being part of this year!s'erop, it may be necessaly to issue the results obtained fi'om it in a supplementary account, though reference to its earlier stages will be made in this paper. A short description of each of the parents.will indicate the nature of the characters involved. Tho~sand Dollar. This is a typical tight grained oat of the spreading, or open, panicle kind. The grains are of good size, fairly long, and firmly ensheathed 'in the paleae. They do not, except when it is exceptionally prolonged and violent, become naked on threshing. The inner paleae are thin but stiff, the outer thick and curled round the edges of the inner. So welt developed is the intervaseutar sclerotic tissue that the fine longi- tudinal ribs of the outer paleae appear only as faint, markings, the whole surface, save at the extreme tip, being perfectly smooth. The spikelets are usually two-grained, though a panicle will often contain a few which are 3-grained. Very rarely a plant will be found which has two or three 4-grained spikelets. In this case the upper- most grain is generally very small and all the peduncles are short and 1 The work herein described was taken up at the 1,~2 stage. The data concerning the parents and the Fl generation, however, were rather vague and scanty, and had ~o be relnves~igated by me concurrently with the anaIysis of the F a crop. Joum. of Gem wi 16 230 Inheritance of Tight and Loose Paleae in Oats straight. It is important to note this condens,~tion of tile occasional large spikelet in tlle tigllC oat as compared witll tlle extreme laxity of bile Avena nuda type. /~ff0720. In all i~s essentials tllis variety resembles Thousand Dollar. It also has white, tight, awned outer paleae and spreading panicles. The grain, llowever, is somewllat plumper and tllere seems a greater tendency towards 3-grained spikelets, though no C-grained have been noticed, possibly because no large field crop llas been available for inspection as in the case of Thousand Dollar. Fig. I. Characteristic spikelet from a "pure loose" containing 8 flower~. Nubischer Schwarzer is a shorter strawed oat than the two just described. It also differs in being a black barbarian. Save for the size and shape of the grain all other points are similar. A vena ~suda. Two distinct features charac~erise the spreading panicles of this species :~ (a) The long, dangling spikelets in which there may be as manyas nine flowers (cf. Fig. 1), and (b) The readiness with which tile grain drops out fi'om between the paleae when mature. A. ST CLAm CApOR~ 231 The parental stock used showed impurity if' the "nuda" type be taken as one in which the paleae are all absolutely membranous, tha~ is to say, with no trace whatever of intervascular sclerosis. For some plants had at least one obviously tight grain, often more, the pro- portion of tights even rising to 4,0~ in extreme cases, while othem would have a grain or two with paleae moderately stiffened in a broadened midrib region. One important character, however, is common to all these forms: they never tlfrow q~'sprinff with ALL the grains tight. In this respect, tlierefore, even if not in others also possibly concerned in the partial Lightening of the paleae, they nvay be regarded as constituting a pure line. In colour the parent plants were mixed greys and whites. Greys are very easily overlooked in Aven~, nuda because colour is developed only on the small inner paleae. THE ~1 GENERATION. Spilcelets. The panicles ot ~0he F~ plants contained a varying proportion of many-flowered spikelets of the "nuda" type. In the basal regions the two-flowered spikelet predominated. 2aleae. Every conceivable gradation fi'om the "pure tight" of the tight grained parents to the wholly membranous palea occurred. On rubbing out, it was noticed that sometimes there was an excess of tight grains, and sometimes not. Six separate panicles, which all threw the same range and similar proportions of F.. types, and must therefore have been F~ heterozygotes of the same factorial composition, were constituted as follows TABLE I. Arena nuda ?. • Thousand Dollar" O. Panicle 1. Excessof loose grains. Tightsabout of total ,, 2......... ~} ,, ,, 3. ,, tight ,, ,, ,, Ligowo ? • Avena q~uda ~. Panicle 4. Excessof tight grains. Tights about ~ of total ,, 5......... ,, ,, 6.. ,, loose ,, ,, ~- ,, It was ~t first thought that the reason for this fluctuation might lie in the fact that single panicles were examined instead of whole phmts ; but 16--2 232 Inheritance q/" Tight and Loose Paleae in Ocets a more detailed analysis of the entire plants in this generation of the Nubiseher cross proved that, although there are cases in which the exact nature can only be ascertained when the basal tillers are also taken into consideration, the main panicle is generally indicative of the whole plant. The necessity for inspecting the other panicles arises chiefly when the excess of either tight or loose grains is very great. In such cases all, or most, of the paleae of the preponderating type may be concentrated into the largest panicle, and the opposite extremes have to be searched for on the subsidiary panicles. In the detailed analysis just referred to the panicles on each //'~ plant were taken separately, in the manner shown in Table II, and the paleae borne on the different branches classified node by node, com- mencing at the base. A "pure tight" palea has no paperiness, a "pure loose" is entirely membranous, while an "intermediate" is a very variable mixture of these two extremes (el. Fig. 2}. Fig. 2. Types of paleae found on F1 plants. A--pure tight; a grain is shown in both dorsal and ventral view, the latter showing the margins of the outer pale clasping the inner one. B =intermediate forms. O=a pure loose palea. The selerotised parts are repre- sented black. The percentage of "pure tights," it will be observed, ranges fl'om 20'4% to 68'7%'; that of the "pure looses" from 7'8% to 4,6'8~ The rise of the one appears to be correlated by tlle fifll of the other, not only when different plants are compared, but also in different zones in the same plant. In plant A, for example, the first nodes of the four panicles bear 7'7 pure tight and 7 pure loose paleae out of a total of 115, i.e. the relative proportions are as 67 ~ : 6'1 ~ In the ease of the second nodes they are as 4@8 O/o : 20'8 ~ For the third nodes they are as ... 26'8 ~ : 39 r For the fourth they are as ...... 5 O/o : ~5 ~ And for the last they are as ...... 7'4., ~ : 44'4 o/o. A. ST CAPO~N 233 TABLE II. .Nq&ischer Sehwa'rzer x Ave~a n',.cla. Analysis of paleae on 5 F 1 plants. [P.t.=puretlght. P.l.=pnreloose. Int.=d:membranous.] Distribution of paleae at Ist node 2rid node 3rd node ,ith node 5th node ]Percentage Plant Panicle P.t. Int. P.I.P.t. Int. P.I. P.t, Int. 1~.1. l).t. Int. P.1. P.~. Int. 1).1. t).t. I'.I. A 1 27 8 2 22 15 15 2 4 5 0 1 3 0 0 0 2 16 18 4 9 10 9 0 3 7 0 2 5 0 2 4 3 23 5 1 15 3 0 ~ 5 3 2 1 3 1 1 5 7 4: 11 0 0 9 12 1 4 4 2 0 4 0 1 6 1 Totals "77 31 7 55 40 25 11 14 16 1 10 9 2 13 12 45'2~ 21'4~ 1 4 17 12 5 9 20 2 5 7 0 5 8 0 3 8 2 27 4 2 13 11 0 5 10 11 1 1 t2 2 7 19 3 4 7 6 5 17 10 5 10 11 5 5 6 0 5 18 4 4 2 ! 0 8 10 1 3 7 0 2 8 0 2 14 Totals 39 30 21 23 45 40 13 28 36 6 13 34 2 17 59 20'4~ 46.8~[o t 7 10 7 2 13 12 1 $ 10 0 5 7 O 1 10 2 26 9 4 13 9 7 7 8 5 0 6 7 0 2 4 3 11 14 9 7 15 6 3 8 5 0 3 2 0 3 5 4 6 3 0 i0 6 2 9 4 1 5 3 0 2 3 7 5 10 6 2 3 8 5 10 1 5 2 3 6 0 4 6 6 13 2 2 5 12 I0 1 5 3 6 5 15 0 0 8 7 7 2 0 II II 5 8 4 4 3 "3 8 3 2 7 8 24: 1 0 !1 7 2 8 6 3 4 1 2 4 1 2 Totals 10t 47 24 62 81 49 47 44 36 20 29 47 9 16 49 3(]'4% 30'9~ D 1 30 0 0 18 0 0 5 2 2 1 ! 0 1 1 0 2 i0 8 2 12 12 2 3 4 3 0 2 6 0 2 6 3 12 0 0 6 0 0 I0 1 1 1 1 2 0 2 4 Totals 52 8 2 36 12 2 18 7 6 2 4 8 1 5 t0 63% 16.2% E 1 11 8 0 16 5 0 10 3 0 2 0 0 2 2 2 2 3 0 0 ,I i 0 4 1 0 1 1 0 0 2 0 3 19 1 5 5 1 1 1 1 0 1 1 1 tip broken off Totals 33 9 5 25 7 1 15 5 0 4 2 1 2 4 2 68,7~ 7.8~ 234 Inheritance of Tight and Loose Paleae in Oats Fig.