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Polymorphism in pollen of Salvia leucantha(Lamiaceae)Asha Gupta a & Chhaya Sharma aa Birbal Sahni Institute of Paleobotany , 53 University Road, Lucknow, 226007, IndiaPublished online: 01 Sep 2009.

To cite this article: Asha Gupta & Chhaya Sharma (1990) Polymorphism in pollen of Salvia leucantha(Lamiaceae), Grana, 29:4, 277-284, DOI: 10.1080/00173139009428938

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Grana 29: 277-284. 1990

Folymorphism in pollen of Salvia leucantha (Lamiaceae)

ASHA GUPTA and CHHAYA SHARhfA

Gupta, A. 6r Sharma, Chhaya 1990. Polymorphism in pollen of Salvia Iettcatriha (Lamia- ccae). - Grana 2?? 277-281, 1990. Odensc 25 Octobcr 1990. ISSN 0017-3131.

Lhl and SEhl study of the pollen from single specimen of Snhia lertcaniha Cav.. gathered from Kumaon mountains in Western Ilimalaya have shown significant variations in aperture number and type. shape and size of pollen grains, hitherto not reported. Normal pollen is 6-colpate but 4, 5, 7, 8 , 9, 10, 1 I-colpate to spirapcrturate also occur to make it a hetcroge- nous assemblage. In shape, pollen varies from oblate, suboblate, oblate-sphcroidal, prolate- sphcroidal to subprolate, ivhereas, in size it ranges from 15 Iim to 40 Iim. Furthermore. besides the marked variations in monads, the polymorphism is also expressed in tcrms of dyads and triads. Exine ornamentation however, does not vary, it being reticulate-retipilate sexine pattern in all the different types under Lhl. and shotking double sculpturing under SEhl.

Aslin Giipta atrd Clrlrajn Sharttra. Birbal Sohtri Ittsiiirtte of I’alaeobotatiy, 53 Utiiversity Road, Ltrckrro~c~ - 226 007, India. (Matiitscript received 6 Sepietnher 1989; revised versiotr accepted 8 May 1990.)

Lamiaceae (Labiatae), a cosmopolitan family com- prising nearly 200 genera and about 3500 species (Willis 1973) is well represented in the Indian sub- continent.

The genus Salvio is \veil distributed all over the world and has over 700 species, many of them culti- vated as ornamentals for their bright coloured in- floresence, flower and foliage, both in the tropical as ivell as temperate climate. Itowever, a few species occur in alpine or subalpine regions of the Himalaya and other montane regions. In the Indian subconti- nent, the genus is represented by over 36 species occurring across a wide range of climatic types from extremely xeric conditions in the plains of Indian desert to the Iieights of alpine and subalpine zones in the Himalaya (Dhar 6: Kacliroo 1983).

Sdr.ifi Ieiictiritlia is native to hlexico but has nat- uralised recently in some subtropical and tempcrate belts of Himalaya. I t has however, been reported from hlussoorie - at 2100 m (Raizada 6: Saxena 1978), Naini Tal - about 2000 m (Gupta 1968) and Eastern Himalaya between 1500-1700 m (Hara 1966).

Laniiaceae is basically stenopalynous, having ei- ther 3 or 6-colpatc pollen (Erdtman et al. 1961). As carly as in 1956, palynological studies in Salvia was carried out by Ikuse (1956; 7 spp.) and after him a

number of workers have investigated its several spc- cies, for example Erdtman et al. 1963 (1 sp.), Embo- den 1964 (lospp.), Nair 1965 (4 spp.), licndcrson ct al. 1968 (56 spp.), Varghcsc 6: Verma 1965 (4 spp.). Vij 6: Kashyap 1975 (4 spp.), Markgraf 6: D’Antoni 1978 (1 sp.), Bir 6: Saggoo 1981 (5 spp.), Forlani 1981 (1 sp.), Saggoo & Bir 1983 (5 spp.), Gupta 6: Sharma 1986 (3 spp.). Except for a few cultivars in which polyploidy or an increase in chromosome number had led to the proportionate increase in the number of colpi (Maurizio 1956), 6-colpate pollen has been reported in all the species. However, ab- normal pollen has also been reported earlier by a number of workers in different plant taxa namely, Nair 6: Sharnia (1968) in Sisyriibriiitii iriu, Nilsson (1967, 1970) in Gcntianaceac, Erdtman (1969) in Iturippo sylvestris, Chaturvedi (1974) in Nyrtyliaea slellafa, Van dc Beck (1974) in Riibiis, Clarke (1976) in Hypericirm, Dupont (1977) in Jetiseribolryci, Fer- guson (1980) in Ceraforiia, Guinet 6: Graus-Cavag- netto (1986) in Ncpriirtio, etc.

Recently Kumar 8: Subramaniam (1987) suggest that there is little variation in chromosome number in the Lamiaccac, most spccics having 2 n = 22. 1 io\vevcr, spccics in cultivation may exhibit varia- tion in thc chromosome numbcr evcn within the same taxon.

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hlATERIALS A N D hlETi IODS I'ollcniferous material \vas collected during October 1958, from one of the plants inhabiting the forest near Kainchi, a roadside sm;ill settlement on tvay from I3him Ti11 to Naini Tal. The voucher specimen has been deposited with the llcrharium. at Birbal Sahni Institute of Palaeobotany, l.ucknow, under the Registration No. 11420.

The pollen material ivas acetolysed (Erdtman 1943, 1360) and slides w r c prepared in glycerine jelly. hlorph- ological observations \vere made with ;I Leitz Diplan micro- scope. using 10 x eyepiece and 100 x objective (oil-im- nicrsion) whereas, the photomicrographs with Lcitz auto- m;itic photo-equipment.

For scanning electron microscopy (SEhl), the acctolyscd pollen grains w r c mounted on aluminium stubs and coated \vith gold-palladium. The photomicrographs ucrc taken with l'hillips Holland 505 microscope.

Pollen descriptions and size measurements arc based on r;indom sclcction of 4&50 acctolyscd pollen grains and the terminology follotved here is after Erdtnian (1952) and Erdtman ct al. (1061) tvith some modifications.

RESU IXS

Pollen grains in Sah-io lciicorrtlra are predominantly monads (Fig. 1 A-W; Fig. 2 A-E; Fig. 3 A-F) but dyads (Fig. 2 F-hi; Fig. 3 G , 11) and triads (Fig. 2 N) also occur in very low frequencies (Table I ) . hlonads arc gencrolly 6-colpate (%YO), followed by S-colpatc (IS%), and much less frequent I-colpatc (So/), 5-

colpatc (5%). 1 I-colpate (2%) and spirapcrturate (5%) which amount almost 92% monads. The fre- quency of dyads and triads is nearly 6% and 2% respectively. Resides the above statcd different apcrtural types, tlie pollen grains have shown a great range of variation in their s i x and shape. It has further been observed that generally the size of the pollen increases with tlie increase in colpi number as cvidcnt by the fact that as small ;IS 15 p i arc the 4-colpatc one whcrcas 40 Itm pollen largest i n size arc spirapcrturate. However, dyads and triads are normally larger i n size compared to distinctly smaller monads. Different pollen shapes, occurring within this species vary from ohlatc, suboblate, oblate-

colpatc (2%), 7-colpatc ( I Y O ) , 9-colpate (2%), 10-

spheroidal, prolate-spheroidal, to subprolate. Exinc thickness ranges from 1 Itm to 2 Itm with a reticulate- retipilate pattern which is either homobrochate or hetcrobroc1i;itc. Colpi are either equi or unequi dis- tance placed. Colpi margins are surrounded with equal or smaller meshes.

The various pollen types encountered during the present investigations - nine forms under monads as \sell ;IS dyads and triads are described below .

L hl- 0 BS E RVATl ONS

hforrods J - Z O I J O C O I ~ ~ C typc (Fig. 1 A D ) . - Oblatc-sphcroi- dal (15 X 15.5 p i ) . Colpi long, thin, running almost pole to pole with acutc apices, equidistantly placed. Exine 1-1.5 pni thick. Sexinc & as thick as ncxinc, scmitcctatc, reticulate-retipilate, honio-heterobro- chate, muri with single row of bacula.

5-zoriocolpate type (Fig. 1 E, F). - Oblate-spheroidal (18 X 19 pn). Colpi long. thin, running almost pole to pole with acute apices. -C equidistantly placed. Exinc 1.5 pm thick. Scxinc iis thick as nexine or slightly thicker, scmitectate, reticulate-retipilate, homo-heterobrocliate, muri with single r o w of bac- ula.

6-zonocolpote rype (Fig. 1 G-L). - Suboblate (19 x 23 pm), oblatc-spheroidal (19 X 20 Itm), prolate- spheroidal (19 x 17 I'm). Colpi long, thin, running almost pole to pole with acute apices, equi-to un- equidistantly placed, colpi margins slightly wavy at times. Exine 1.5-2 Itm thick. Sexine slightly thickcr than nexinc, seniitectate. reticulate-retipilate, hctcr- obrochatc, muri with single row of bacula.

7-zoriocolpate type (Fig. 1 hl) . - Oblate (17 X 21 p ) . Colpi long, thin, running almost pole to pole with acute apices. equidistantly placed. Esine 1.5 Iim thick. Scxine k as thick as nexine, seniitectatc, reticul;ite-retipilate, hctcrot)rochatc, niuri with sin- gle row of bacula.

Fig. I. A-W Light micrographs of pollen grains of Sdvin Iertcmi!/iu X 1ooO. (A-D) 4-colpate pollen in polar vicw. (E, F) 5-colp:itc pollen in pol:ir view. (G-I) 6-colpate pollen in polar view. (J-I-) S;ime in equatorial view showing subprolate, oblate-spheroidal and suboblate shapes respectively. (hl) 7-colpate pollen in polar view. (N) 8-colpatc pollen i n equatorial vicw. ( 0 - R ) S a n ~ in polar vicrv. (S) 9-colpatc pollen in polar view. (T) 10-colpate pollen in polar view. (U . V) Same in polar view shwing 2 tri:ingular :imh. (W) 1 I-colpate pollen in polar view.

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8-zoriocolprrre type (Fig. 1 N-R). -Oblate (22 x 32.5 jirn, suboblate 26 x 31 pn), oblate-spheroidal (28 x 29 jim). Colpi long, thin, running froni pole to pole with acute apices, unequi to equidistantly placed. hfesocolpia always larger at the equator. Exine 2 jirn

thick. Sexine k as thick as nexine or slightly thicker at times, sernitcctate, reticulate-retipilate, hetcro- hrochate, rnuri with single row of bacula.

A. Gzrpra atid C. Slzarttia

- FREQUENCY

OCCASIONAL

RARE

DOMINANT

RARE

CO-DOMINANT

RARE

OCCASIONAL

RARE

OCCASIONAL

OCCASIONAL

RARE

RARE

9-torzocolpnre rype (Fig. 1 S). -Oblate (20 x 30 - 26 x 35 jirn), oblate-spheroidal (22 x 24 jirn). Colpi long, thin, running almost pole to pole with acute apices, unequidistantly placed. hlesocolpia larger at equator. Exine 2 pn thick. Sexine k as thick as nexine, semitectate, finely retipilate-reticulate, het- erobrochate, muri with single row of bacula. _ _ _ - - --- 10-zonocolpare type (Fig. 1 T-V). - Oblate (23 X 36 Iirn), oblate-spheroidal (30 x 33 Iirn). Colpi long, th in , running from pole to pole with acute apices, unequidistantly placed. hlesocolpia larger at the cquator. Exine 2 Iirn thick. Sexine f as thick as nexine, sernitectate, finely retipilate-reticulate, het- erobrochate, muri with single row of bacula.

~ -

If-zoriocolpare type (Fig. 1 W). -Suboblate (28 x 33 p). Colpi long, thin, running from pole to pole with acute apices, equidistantly placed. Exine 2 jirn

thick. Sexine slightly thicker than nexine, sernitec- tate, finely retipilate-reticulate, heterobrochate, rnuri with single row of bacula.

Spirnpertrrmle rype (Fig. 2 A-E). - Suboblate (32 X 40 Iirn), oblate-spheroidal (33 x 36 Iirn). Colpi long, thin, number not discernible. Exine 2 jim thick. Sex- ine thicker than nexine, semitectate, reticulate-reti- pilate, heterobrochate, rnuri with single row of bae- ula.

Djntls

\Virh f cqiralsize ofttionads (Fig. 2 G-hl). - hlonads suboblate (13 x 16 pin), 4 to 6-colpate. Colpi long and thin. Exine 1.5 jirn thick. Sexine as thick as nexine, sernitcctate, reticulate-retipilate, heterobro- chate, rnuri with single row of bacula.

TABLE 1.POLLEN TYPES IN SALVIA LEUCANTHA

:ATEGORY POLLEN TYPE l

4- ZONOCOLPATE

7- ZONOCOLPATE

8- ZONOCOLPAT E

I 9- ZONOCOLPATE

I 10 - ZONOCOL PAT E

11- ZONOCOLPATE

S PI R A P E R T U RAT E

WITH 2 EQUALMONADS

WITH UNEQUAL MONADS

With irrieqiral size of tilorinds (Fig. 2 F). - hlonads oblate-spheroidal (18 x 19 Iirn & 31 x 32 p), generally 4 to 6-colpate, rarely spiraperturate. Colpi long and thin. Exine 1.5 Iirn thick. Sexine generally as thick as nexine, sernitectate, reticulate-retipilate, heterobrochate, rnuri with single row of bacula.

Trinds (Fig. 2 N) Triads 23 X 14 jirn. hlonads of I equal size (13 X 14 pn, 14 x 15 jini and 14 x 15 Iirn), 4 to 6-colpate. Colpi long and thin. Exine 1.5 Iim thick. Sexine as thick as nexine, sernitectate, reticulate-retipilate, heterobrochate, rnuri with single row of bacula.

SEM obsrenwrioris (Fig. 3 A-H) In SEM, pollen grains reveal some interesting fea- tures, not possible when seen through Lhl. For ex- ample, pollen have actually double ornamentation.

Fig. 2. A-N Light micrographs of pollen grains of Solvin Icirccinrhri X 1 0 . (A-E) Spirapcrturatc pollen sho\ving apcrtural oricntation and scxinc pattcrn. (F) Dyad with uncqual monads. (G-hl) Dyads with equal mon:ids showing scxinc pattcrn, cxinc thickness and apcrturnl variation. (N) Triad.

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Lumina of primary reticulations arc further studded with comparatively much smaller secondary reticu- lations. Similarly, number of mcshcs in secondary reticulations apparcntly correspond to thc size of the primary lumina i.c. smaller the primary lumina, lcsser the mcshcs, \vhcreas in case of largcr lumina, comparatively more sccond;iry reticulations arc seen. Furthermore, extra smallcr meshes surround- ing colpi margins arc also distinct in SEhl.

A. Cirpta arid C. Slinrrria

DISCUSSION AND CONCLUSION

The present study sets out to exaniine the pollen of Snliin hicaritha through Lhl and SEhl, revealing interesting range of morphological variations in sin- gle pollen assemblage. Ever since Ikusc (1956) car- ricd out pollcn studies in Snlr*ia, rcporting 6-colpate pollen grains, the same typc has been invariably reported by subsequent palynologists in other in- vestigatcd spccics of this vast genus, esccpting for a few cultivated tam. It is howcvcr, for the first time that i n one of the taxon of Lamiaccac namely Sahio Icircarithn, about ;I dozen abnormal o r atypical typc of grains arc being rcported, adding to its signif- icance that pollen material invcstigatcd by the au- thors had its sourcc from one of the plants growing in thc wilderness of temperate Himalaya whcrc this exotic spccics has naturalized very recently though not yct common in the arca.

Pollerr apertures In the family Lamiaccac, 6-colpate and 3-colpatc pollen are normal types. In Saltin leircoritlici, 6-col- pate pollcn is the most common, the normal typc. Surprisingly no 3-colpate pollcn has been encoun- tered ticre when it is coniparcd to thc over all range of aperture variations which account for 5 ,7 ,8 ,9 , 10 6r 1 1-colpate types though in much varying frequcn- cics. Infact, Erdtman (1952) had pointed out that in Lamiaceac, 3-colpate pollen is shed at 2-nucleate stage whereas, 6-colpate ones at 3-nuclcate stage. A critical cytological approach is probably the answer to hiivc sonic insight before one is able to correctly interpret such a phcnomcnon of extrcmc flexibility scen in the pollen characteristic of Sah-ia leircarrthn.

Nevertheless, hlaurizio (1956) dcmonstratcd in Snlvia praterisis that in its 2n race ;ill pollen grains arc 6-colpate whcrcas in 4n race most frcqucntly 8-colpate (or 9-10) and 6 ,7 , 11 and 1Zcolpatc grains were seldom. Similarly, in Sahia spleritleris its 2n race has 6-colpi, 3n mostly 6-colpi or rarely 7-colpi but in case of 4n race the number of colpi rangcs from 6-10.

t’ollcri shapes Diffcrcnt pollen shapes have been encountered in Salvia leircaritha i.c. oblate, suboblate, oblatc-sphe- roidal, prolate-spheroidal and subprolate. Accord- ing to Erdtman (1961), induction of polyploidy leads to a tendcncy in pollen grains to round off i.e. the value of WE x 100 tends to move towards 100 (sphc- roidal). Such a trend however, has not bccn ob- scrvcd during the present study and contrary to this as many as 4 diffcrcnt shapes in S. leirrciritlin have bccn observed within 6-colpate pollen alone.

Exilic orrinrrieritatiori Exine character remains rather static in this species and show reticulntelretipilate pattern i n all the dif- ferent pollcn types under Lhl and doublc ornamen- tation under SEhl . t’ollcri size It has been observed that nornially the incrcasc in the number of colpi has resulted in thc proportionate increase in pollen sizc which varies from as small as 15 Iim (4-colpatc typc) to as large as 40 pm (spirapcr- turate). Saggoo & Bir (1953) in thcir cytopalynolog- icnl study obscrvcd that in Lamiaceac thc variation in clironiosonie nuniber does not effect the sizc of the pollen grains. Thcy studicd 9 members of the family and observed small, mcdium as \vcll as largc size pollen grains in taxa having lowest chromosomc number (2n = 16). Earlicr, Erdtman (1961) too, infcrrcd that thcrc is no correlation bettvccn chro- niosomc number and pollen grains sizc among thc members of Laniiaceac. Thus, it could he infcrrcd that incrcasc in the size of pollen grains viz-a-viz the increasc in the numbcr of colpi is probably in no \wy related to the chromosomal bchaviour.

Fig. 3. A-H Scanning clcctron micrographs. (A) 6-colpate pollcn in polar v i w sho\ving double reticulate scxinc pattern. (B) Central portion of thc sarnc rnagnificd. (C) 8-colpatc pollen in polar vicw. (E) Spirapcrturatc pollen. (D. F) Portions of same rnngnificd. ( G , 11) Dyads sliotving doublc rcticulatc scxinc pattern (s:irnc scale bar).

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Polo[)nioryhism in yolleti of Salvia leirctitirha (Larniriceae) 283

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281 A. G i r p nrid C. Shariiia

ACKNOWLEDGEhlENTS The Authors are grateful to Dr. B. S. Vcnkatachala, Direc- tor, BSIP, Lucknow, for the encouragement and facilities. Thanks are also due to our colleague Dr. hl. S. Chauhan, for his help in the field work.

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