Trichomes in Trifolieae
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Prof. Indian Acad. Sei, Vol. 85 B, No. 2, 1977, pp. 77-89. Trichomes in Trifolieae MOHINI GUPTA AND Y. S. MURTY Department of Botany, Institute of Advanced Studies, Meerut University, Meerut 250001 MS received 1 March 1976 ; in revised form 13 July 1976 ABSTRACT Trichomes of 39 species of Trifolieae belonging to six genera have been worked out. On the basis of structure they are classified into nine types and ontogeny of only four types could be studied. The structure and organographic distribution of each lype has been describect. All types of triehomes have been initiated from single initial cell. 1. INTRODUCTION Trichomes of Papilionaceaeespecially those of Trifolieae have been described by Solereder, 1 Metcalfe and Chalk, °" Fahn, 3 Toma, a Toma and Danis, 5 Toma and Buznea, 6 Shah and Kothari, 7 and Kothari and Shah? The use of trichomes in taxonomic delimitation has been stressed by Toma 4 in species of Trifolium, Shah and Kothari ~ and Kothari and Shah s in members of Viceae and Hedysareae. However, no detailed work is available on the distribution of trichomes on different organs and their ontogeny in Trifolieae. Therefore, triehomes of 39 species (tables 1, 2) belonging to all the six genera of Trifolieae have been worked out in detail. Customaiy micro- technique methods were employed. Epidermal peels were stained with Delafield's haematoxylin and mounted in glycerine. 2. OBSERVATIONS On the basis of their structure the nonglandular and glandular triehomes observed in members of Trifolieac arc classified as nine types. The ontogeny of four of the nine types has been studied. Tables 1 and 2 show the type along with their distribution and measurement of trichomcs in different species. 77 ]3 5--Feb. '77 78 MOHIN1 GUPTA AND Y. S. MURTY ug ,,-, b b ...... 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MtmTY x • ° xxxxxx xXx XX • ° ~,-.S~ x "~ ~ • . • ° • • . °° . ° ° °° • , • • • ° • • ..... • : : : : : : • r~ • ° X ..~ xx ".# x XX X X • . .... .-~.~ ° ~. • ~1,¢3 X 2.~ f"q ° ~ ~ ~ x~ XxXxxx ~z o"1~ x • ° XX • ° . • ° ° ° , o ° ° ° ° o~i1 ~X ,q.,h e,. 0 x A4~ >', X X X xx ~ o • • . • • • ° , ° ° ,£ ,b TRI~HOME$ IN TRIFOLiEAE 81 • , . • . , X X X,,4 X X x XX XX XX :x ..... ~ • • • • • ° ° • : • : : : "X ..... " " : • • . • . : • :x " " • • " " " " XX XX XX XxXx:-'IxXX o " ~ " "',,:, "x ..... ~ ~ o ,',, " • 0 . XX XxXX ;..',, X • • • , . • 0 ° X ' " 'Q " X x XXXx X e,I ~ X 4- ";' & o ,,m4 • 0 XX XX XX ..... • . , • ~ • • .. ~ :~. ,,4 82 MOHtNI-GUPTA AND Y. S. M~URTY i I ~ l i • ° ° ....... o g I ~o 0 m~:A~ ~6~ "*A6a l ~RICHOMES IN TRIFOLIEAL 83 NONGLANDULAR TRICHOMES These trichomes are uniseriate and are classified under four types Ate D. Type A.--This tl ichome is short and tlu ee celled with a thumb like apical cell (figure 1). Tjpe B.--This trichome is found only in Ononis and has 3-6 (occasion- ally 9;1 uniformly elongated cells showing~variations in the acuteness of the apical cell. It is acute in O. biflora (figure 2), obtuse in O. serrata (figui'c 4)and acunfinate in O. pabescens (tigure 3) aald O. natrix (figure 7). ONTO~3ENY.--The ontogeny of this trichonle has been studied in O. natrix. An ci)idermat cell functions as an initial which can be differentiated from the rest Of the cells 'by dense cytoplasm and a plominent nucleus. It divides tangentially to form a basal cell and an upper cell. The upper cell divides further (figures 5-7). Tj:pe C.--This trichome is sessile (figure 8) or has a she1 t stalk of one (figures 9-11) or two cells (figure 12). The apical cell has a long pointed apex (figures 8-12). When the stalk is single celled the basal cell protrudes out (figures 9-11, 13). The trichome may be straight, falcate or recurred. ONrOGEN¥.--The ontogeny is studied in Trifolium lappaceum. The hair initizfl can bc differentiated from the rest of the epidermal ¢ells by prominent nucleus. The initial enlarges into Papilla in which nuclem division is followed by scptum formation resulting in a smaller stalk cell and an elongated apical cell (figures 14-18). Type D.--The trichome is two celled with slightly protruding rectan- gular stalk cell and a very much elongated, thick walled, pointed apical cell with tuberculated surface (figmes 19-20). This trichome~may be straight recurred or falcate. GLANDULAR TRICHOMES ~ Tllcsc tricllolnes arc classified under five types E to I. Type E.--Each trichomc has single celled stalk and a globose multicellu far head. Each cell of the head shows a prominent nucleus and dense cyto- plasm. This type of hair is only recorded on the outer margin of the standard petal of Onanis biflora (figure 21). R4 MoHINI GUPTA A~ Y. S. MURTY i~'~"~i/~i I r . ~t~ *~ " N !£~i.i..iy I 005 mm ! ~ 13 .'ol1~'~ .. 17' 18 19 Fftmr~ 1-20 TRICHOMES IN TRIFOLIEAE 85 Type b:--It has uni or biseriate stalk of 3 cells in height and a club- shaped multicellular head (figures 22, 23). Type G.--It has multiseriate stalk of more than 3 rows of cells and one or more cells in height. The multicellular head is oval or club-shaped (figures 24, 25, 26). Type H.--It has a long uniseriate few celled stalk with a broad basal cell. The cells of the stalk are generally broader towards the base and narrower towards the apex. The head is multicellular and spherical a~ in Medicago scutellata (figures 27-28) (r elongated as in Ononis (figures 29, 34). ONTOGZNY.--It is studied in Ononis serrata only. The conspicuous tri- chome initial elongates and divides transversely to form a basal and an upper cell (figures 30-31). The upper cell divides by transverse divisions forming a row of cells (figure 32). The terminal cell of tiffs row enlarges, becomes globular with dense cytoplasm and a prominent nucleus, which divides longitudinally as well as transversely to form a multicellular head (figures 33, 34). Type L--The stalk may be one celled as in Trifolium subterraneum, (figure 36), T. hirtum, T. repens (figure 37), Trigonella stellata (figure 38) Medicago orbicularis (figure 35), M. arabica (figure 39) and Melilotus officinalis (figures 43, 48) or more than one celled as in Trigonella caerulea (figure 40), Medicago media (figure 49) and Melilotus sulcata (figure 50). Stalk may be very narrow or almost of the same size as compared to the base of the head (figures 39, 42, 44). Head is generally biseriate (figures 45-47)but occasionally multiseriate (figure 41). ONTOGENY.--Ontogeny has been worked out in Trigonella callicerasoites only. The prominent initial cell divides transversely into two. The upper cell divides further to form a uni, bi or multicelled stalk and an upper head initial cell which divides to form a multicellular head (figures 51-54). 3. DISCUSSION Nonglandular trichomes were reported earlier in some members of Trifolieae like in Trifolium alpestre, 1 Trifolium and Ononis, ~ species of Trifolium, 4 Melilotus alba and M. officinalis, 60nonis spinosa, O. pseudohircina, O. hircina, O. columnae 5 and in some species of Trigonella, Melilotus, Medi- cago and Trifolium. 7 In the present study nonglandular trichomes are observed B--6 86 MOHINI GUPTA AND Y. S. MURTY 35 36 O~mmA S2 44 S4 I 0~3~ mm ¢ I " ~tmm f I 9 SO' Figures 21-54 TRICHOMES IN TRIFOLIEAE 87 in 38 out of 39 species examined. Nonglandular hairs resembling in general features the types described here as B, C and D have been reported from members of Trifolieae by earlier workers without any mention about their distribution on different organs. But trichomes of ' A' type have not been reported earlier. It is interesting to note that the variations observed in the apex of the apical cell in different species of Ononis examined as acute in O. biflora, obtuse in O. serrata and acuminate in O. natrix and O. pubeseens can be usefu I for distinguishing the species from one another. Glandular trichomes which may be club-shaped with or without a distinct stalk have been reported in all the six genera of Trifolieae. ~ They are recorded in members of Melilotus, 6 Trigonella, Melilotus, Medicago and Trofolium ~ and Ononis.3, 5 In the present study out of 39 species belonging to all the six genera, 38 species showed glandular trichomes which are generally club- shaped with uniseriate stalk. However, biseriate stalk in Trifolium pratense and Ononis natrix and multiseriate stalk in Ononis natrix and Parochetus communis is also observed. Trichomes of Medicago scutellata and Ononis differ from all other members of Trifolieae examined in possessing a long uniseriate stalk. The shape of the head can be used for distinguishing the trichomes of Medicago scutellata from those of Ononis. In Ononis the head is elongated while in Medicago scutellata it is spherical.