CHEMOTAXONOMY OF GERANIACEAE^
T. SANT PRASAD REDDY AND L. L. NARAYANA Department of Botany, Kakatiya University, Warangal
a b s t r a c t
The Chemotaxonomy of 10 ^ecies di»tributed in 3 genera of Geraniaceae has been investi gated. They exhibit similarities as well aa differences in their chemical characters. Leuco-antho- cyanins are present in the seeds ofall the spodes studied, except Erodium maiacoidet. The presence of triterpenoids in ErotHum matacaidti. Geranium dunctum and G. earolimanum and steroids in G. molU has been inferred. These are however absent in the re.naining taxa. The relatio.Dhips of Geraniaceae with Oxalidaceae, Balsaminaceae, Tropaeolaceae and Linmanthaceae have been discussed in the light of chemical data gathered from present study and the available data from other disciplines. It is pointed out that while the similarides in chemical characters indicate a probable relationship between them, the differences justify their treatment as independent families.
INTRODUCTION ha^ been placed under the order Gora- niales. There is however, difference of Geraniaceae comprises 5 genera and opiniort about the circumscription of the 750 species (Willis, 1966). The members order, the number of families included of the family are cosmopolitan in dis under it being fieven according to Ben- tribution, mostly extending from Tem tliam and Hooker C1862-1883), twenty perate to sub-tropical regions of the World according to Engler and Pranti (1931), (L. H. Bailey, 1969). Most of the species twenty one according to Bessey (1915), are cultivated for their ornamental value. nineteen according to Takhtajan (1967), However, some of them are medicinally ten %pcording to Benson (1957), thirteen important (Chopra, Nayar and Chopra, according to Thom e (1968) and five 1956). according to Hutchinson (1959, 1973) Bentham and Hooker (1862-1883) and Cronquist (1968), treated Geraniaceae as a tribe, Geraniae While the family has received con under the family Geraniaceae, while Bes- siderable attention from the point of sey (1915), Engler and Pranti (1931), floral anatomy, embryology, anatomy an4gM Benson (1915), Hutchinson (1959), Thakh- palynology, it has not received much tajan (1967), Cronquist (1968) and Thome attention from the point of Chemotaxo (1968) accorded it the status of an in nomy. Gibbs (1974) reviewed the work dependent family, Gcraniaceae. In all on the Chemotaxonomy of Geraniaceae. the systems of classijficatbn the family Recently Bate-Smith (1973) worked on
1. AccQited for publication on November 4, 19B5. F&st author (X. S. P. S..) is grateful to Univetiity Grants Commission, New Delhi for the award of teacher feilowihip under Faculty Itnprovemsnt Programme. We e x p tm our deq|) seme of gratitude to the Director, Univetyity Botanic Gatdea, Denmark for ptovid- ing tbe «eedi of the difiismat i^eoiek the flavonoids of Geranium and discussed Hydroxyquinones, Tannin test and Alka the taxonomic implications of the distri loid test. bution of the substances. The present The Cigarette test was negative in all study deals with the Chemotaxonomy of the taxa except Geranium dissectum, Ero- Germittm colitmbimm L., G. dissecUm L., dium malacoides and Pelargonium inodorum G. molle L., G. pusillum Burm fil., G. pyre- which showed Oxalis reaction indicating naicum Burm fil., G. caroliniamm L., G. the acidic nature of the cell-sap. Oxalis rotuHaifolivm L., Erodiim cicutarium (L.) reaction was first reported by Dykyj- L’ He’ rit., E . malacoides (L.) WiUd., Sajfertova (1958) in the species of OxaUs. Pelargonium inodorum Willd. Stems, roots and leaves were tested separately to detect the presence or ab MATERIALS AND METHODS sence of leuco-anthocyanins. The diffe All the materials in the present study rent species differ in the site of localiza were collected from the plants raised tion of leuco-anthocyanins (present study). from seeds obtained from Denmark. They arc present in the stem and roots of Using fresh materials, consisting of Geranium carolianum, G. dissectum, G.pusillum, roots, stems, leaves and flowers saponin G. rotundifolium, Erodium malacoides and test ‘A’, Syringin test ‘A’, Leuco-antho- Pelargonium inodorum and absent in Erodium cyanin test ‘A’, Juglone test ‘A’, Aurone cicutarium. Leuco-anthocyanins are present test ‘A’, Ehrlich test, HCN test, HCl/ in traces only in the leaves of Geranium Methanol test, Maule test, Cigarette test, rotundifolium and absent in all the remain Hot-Water test and the test for hydroxy- ing taxa (present study). However they are quinones were carried out. present in the seeds of all the species Using 80% methanolic extracts of under study with the exception of Erodium shade dried plants collected during malacoides. flowering and fruiting the tests for Sapo- According to Bate-Smith & Ribe- nins. Tannins, Phenols, Carbohydrates reau-Gayon (1959) the presence or absence (Molisch test), Flavonoids (Shinoda test). of leuco-anthocyanins in the testa reflects Alkaloids, Indoles (Ehrlich test), Lignans, more consis entiy the character and rela Leuco-anthocyanins, Triterpenoids (Nol- tionships of the family as a whole than ler’s testj. Steroids (Salkowski reaction) does their presence in the leaves. The and Labat test were carried out. presence of leuco-antliocyanins in leaves i.s influenced by herbaceous habit in the RESULTS AND DISCUSSION plant, while the seed coat is not so in The data gathered from the present fluenced or only to a lesser extent. When study are presented in the Tables I, II leuco-anthocyanins are present in the and III. leaves of a plant, they are almost always From the tables, it will be obvious present in the seeds also, but they may be that the different taxa of Geraniaceae present in the seeds, although absent from under the present study resemble one the leaves. Thus herbaceous members of another in the uniformly positive reaction Geraniaceae differ so far as the leaves are for Flavonoid test, Phenol test and Mo-,, concerned from the woody membcs of lisch test and uniformly negative reaction the Geraniales in having no leuco-antho for Hot-Water test, HCN test, Syringin cyanins, but the seeds are consistently test‘A , Juglone test, Maule test, Aurone positive foi leucc -anthocyanins. test ‘A , Sapamn test ‘A’, E,hrlich test, Bate-Snuth and Lemer (1954) poin- T A B L E I
Tsm vnrR nssR m aterials
1 Auniae tert *A'. 2 Cigwrctte tot, 3 Ehrlich tot, 4 HCl/Methanol tot, 5 HCN tot, 6 Hot-Water tot, 7 Hydroxyquinone test, 8 Juglone teat; 9 Leucoanthocyanin test'A ’, 10 Maule test, 11 Saponin test, 12 Syringin tot.
Name oftbe plant 12345678 9 10 11 12
ErtMum cieutarium (L) , O.R — St— — O.R — — Lf~ — — •— L’ H e 'rit. R t— St— Rt—
EroJivn malocoitUs . O.R. — St+ — O.R — — Lf— — ? — (L) WiUd, R ti- St + R t+
Gtntmm caiolimamm . — — St— — — — — IX— — — — L. Rt+ St+(traa») R t+ (trao o )
Gmamtm caluman L,
OenniunKiifsictim L. . O.R — St-f- — — — — LI'— R t+ S t+ Rt +■
G trm iim molU L.
Geramwn pusillum . — — St+ — — — — Lf— — — — Burm. fil. R + S t+ R t+
Geramum pyttnaieum Burm. fil. , — — — — — — — — — — —
Gtraniam rotimdifolutm . — — S t+ — — ^ — — Lf-t- — — — L. Rti- St+ R t+
Pilargpnitm inaiorum . O.R — St-f — — — — U— — ? — wad. R tl- S t+ Rt-i-
O.R>«OsalU reaction; Lf«»Le*f; St=“ Stem ; Rt=»Root ted out that there is a tendency for the leuc»-anthocyanms in one or the other forination of ileuco-anthocyanins in tiie part of the plant body of the taxa studied more primitive taxa and absent from the (present study), their origin from a stock more advanced members of a phyletic which had ieucoanthocyanins,, can be series. Thus on this basis, Herbaceae of inferred. Hutchinson (1946, 1948) as a whole may Absence of cyanogenesis and alka- be regarded as phylogenetically advanced loids seem to characterise the family (prc- ovwr Lignosae. In view of the presence of sent study). TABLE II T e w s w it h '8 0 % m e t h a m o u o bx t * aots
1 Alkaknd tcM, 2 FUvonoid test, 3 Indoles test, 4 Labat test, 5 Leoco-anthoeyanin test, 6 liebermann' Burchard test, 7 Ugnan test, 8 Molisch test, 9 Noller’s test, 10 Phend test, 11 Salkowaki reaction, 12 Saponin test, 13 Tannins test.
Name of the plant 1 2 3 4 5 6 7 8 9 10 11 12 13
Endkim cicutarium {L) — + __ _ _ — + + _ -1- _ _ __ V H e’rit.
E ndiim nuUacoides (L) — + - - + + + + + + _ _ _ WiUd. Red
Gtranium carolinianum L. — + ~ + + + — + + + — — — blue green
Gtranium columbinwn L. — r — — — — t- — + _ _ _
Gemaum dismtuin L. — + _ — -t- + ~ + + + _ _ _ Gtraniunt motU L. — _ _ _ + — + _ f —
Gtrmiwn ptisUlam — + — — + — 4- — — — Bunn. &1.
Gtrmrnm frfwuacum — + — — ~ — — + — + — — Uurm. til.
Geranium rolundifolium L.— + __ + — + — — — Pelargonium imdenm —• X — — + — — + — + — — — WiUd.
TABLE III
Test with T e« with Test with Type of Name of the plant Mg/HGl conc. H SO aq. N ,0 H 4 compound
Eradium cicutarium (L.) L’ He’ rit. YeUow Yellow Brownish yellow Isoflavones £. malacoides (L) Willd. Rcddiih yeilow Reddish yellow YeUow Flavonol Gtranium carolimanam L. Yellow Olive groeo Yellow IsoBavones G. eolumbimim L. Pale yellow YeUow Pale yellow Isofiavooes G. dissectum L. Yellowish Yellowish YeUow Isofiavones green green
G. moiWL. YeUow Ydlow YeUow Isofiavones
G. pusillum Burm. fil. YeUow Reddish yellow Yellowish brown1 Flavone* YeUow YeUowiahred YeUow Flavonot Reddish Reddish ydlow Ydlow Flavonol YeUow YeUow Ydkwr Ifoflavones Negative reaction for Liebermann- positive results. Burchard test, Nollcr’s test and Salkowski Gibbs (1974) found a close correla reaction in Geramum columbinum, G, pusil- tion between positive HCl/Methanol test Um, G. pyrenaieuM, G. roimdifolium, Ero~ and positive reaction for leuco-anthocya- divm cimtarium and Pelargonium inodorum nins. In most of the taxa of present study, indicate that both tritercpenoids and ste there is a close correlation between positive roids are absent in these taxa. HCl/Methanol test and positive reaction By the positive reaction for Lieber- for leuco-anthocyanins, except in Geranium mannBurchard test (Red colour), Noller’s carolinianvm where IeucoGeranium molle responded positively where their doubtful presence is inferred. lor Liebermann-Burchard test and Salko In the following paragraphs, a dis wski reaction and negatively for Noller’s cussion on the relationships of Geraniaceae test indicative of the presence of steroids with families like Oxalidaceae, Balsami- and absence of triterpenoids. Thus the naceae, Limnanthaceae and Tropaeola- different members in the present study ceae, generally considered to be related differ in the presence or absence of steroids to Geraniaceac is presented taking into and/or triterpenoids. consideration ihe chemical data as well as Though tannins were absent uni data from other disciplines of Botany. formly in the Ten taxa investigated, Batc- Geriuiiaccae closely resemble the Oxa Smith and Metcalfe (1957) reported the lidaceae in many of its chemical features. occurrence of ellagic acid in Geranium The resemblances are in the absence of pyrm aiam . raphides, aluminium accumulation, muci Flavonoids arc present in all the taxa, lage and negative reaction for Hot-Water though they differ in the type. On the test. Oxalis reaction, leuco-anthocyanins basis of colour reactions with Mg/HGl, (preset study; Gibbs, 1974) and tannins concentrated sulphuric acid and aqueous are known in both taxa (sec Gibbs, 1974). sodium hydroxide, in the presence of iso- Cyanogenesis unrecorded in Geraniaceae flavones in Geranium columbinum, G. dissec- is known in three taxa of Oxalidaceae tum, G. molle, G. carolinianum, Erodium (Bal Reddy, 1979). In the distribution cicutarium and Pelargonium inodorum, Fla- pattern for the phenolic acids also Gera vonols in G. pyrenaicwn, G. rotundifolium niaceae exhibit similarities as well as diffe and E . malacoides and flavones in G. pusil- rences with Oxalidaceae. Both taxa are lum is inferred. known to contain gentisic acid, ferulic The stem and root of Greanium dis~ acid, p-coumaric acid, while myricetin, sectum, G. pmillum, G. rotundifolium, Erodium ellagic acid, quercetin, kaempferol, shiki- malacoides and Pelargonium inodorum were mic acid and caffeic acid are known only fotuid to be uniformly positive for HCl/ in Geraniaceae and are absent in Oxali Methanc>r test, while G. molle, G. pyreriai- daceae, cum and E . cifutarium responded negatively. Geraniaceae and Oxalidaceae resem In G, carolimanum, tte stem showed a ble one another in essential ^bryological negative reaction while the root gave (Thathachar, 1942; Narayana, 1962^ Go- vindappa and Boriah, 1956; Johansen, bundles in the petiole, absence of a ring 1950j, floral anatomical (N'arayana and of mechanical tissue in the pericycle Arora , 1963a; Narayana, 1966), anato and presence of raphide sacs (Metcalfe mical (Metcalfe and Chalk, 1950) and and Chalk, 1950). cytol(^ical characters (Warburg, 1938; Though the ba.sic plan of the flower Darliogton and Wylie, 1955; Fedorov, in Geraniaceae and Balsaminaceae is the 1974). However, Geramiaceae are cury- same, the structure and anatomy of palynous whereas Oxalidaceae are stco- the flower in Balsaminaccac present ad palynous (Erdtman, 1971). vanced traits over Geraniaceac (Narayana, Chemotaxonomically, the Balsaniina- 1966). Thus the Balsaminaceae, though ceae are comparatively little known when resemble Geraniaceac in certain floral compared to Geraniaceae. The two taxa traits, appear di.stantly related because resemble one another in the absence of of the acquisition of characters unknown aurones, indoles, alkaloids, hydroxyqui- in Geraniaceae. . nones, aluminium accumulation and pre Pollen morphology does not indicate sence of leuco-anthocyanins. Negative re any close relationship between Gerania sults were recorded in both taxa for ceae and Balsaminaceae (Erdtman, 1971). Ehrlich test, Hot-Water test. Cigarette The haploid chromosome numbers test, Juglone test ‘A’. Presence (doubtful) n=8, 9, 10 (Darlington & Wylie, 1955) as well as absence of saponins has becji and diploid numbers 2n=>18 and 20 reported in Geraniaceae and Balsamina- (Fedorov, 1974) are met with both in ceae. Though the tannins have not been Geraniaceae and Balsaminaceae, indica detected in the taxa under present study, tive of a probable relationship. they were reported in both Geraniaceae The data on the chemistry of Tro- (Bate-Smith, 1973) and Balsaminaceae by paeolaceac is scanty for a comparison with Gibbs (1974). Cyanogenesis, unknown Cieraniaccae. The available data shows in Geraniaceae was reported in one taxon that there arc similarities as well as diffe under Balsaminaceae. Presence of gentisic rences between them. In the absence of acid, p-OH benzoic acid, has been on raphides, mucilage, aluminium accumula record in Geraniaceae while there is no tion, myricetin, delphinldin, cyanidin, si- information in Balsaminaceae for com napic acid, cyanogenic glycosides and the parison, Raphides, present in Balsam negative reactions for aurones, Cigarette inaceae, are absent in Geraniaceae. test, Hot-Water test, Ehrlich test and the Geraniaceae resemble Balsaminaceae presence of quercetin, kaempferol, gentisic in certain cmbryological features. How acid, p-OH-benzoic acid and caffeic acid ever in characters such as 2-celled micro- are observed in both taxa (present study; gametophyte, dorsal raphe, well-differen Gibbs, 1974). Sedoheptulose and leuco- tiated endothelium, tendency towards anthocyanins present in Geraniaceae are unitegmy, Allium and Polygonum type not recorded in Tropaeolaceae (Gibbs, of embryo sac ontogeny, cellular endos 1974). Tannins occur in certain taxa of perm with micropylar and chalazal both the families (Gibbs, 1974). haustoria, the Balsaminaceae are at In essential embryological characters, variance with those of Geraniaceae. Geraniaceae resemble Tropaeolaceae (Na Anatoinicaliy the Balsaminaceae stand rayana, 1967). However Tropaeolaceae apart from Geraniaceae in such featiures stand apart from Geraniaceae in embryo- ^ having an arc instead of a circle of logical features such as teninucellatc tmUes, aggr«sBive suspcnsor or haustoria, in Geraniales or Sapindales and favoured Sdmmd type of embryogcny and absence the creation of a separate order Limnan- (rf paritetal cell. thalcs. Anatomical data on record (Mfctcalfc Limnanthaceae resemble Geraniaceae and C^aik, 1950) do not give any definite in anatomical characters like widely spac clue about the relationship of Tropaeo- ed vascular bundles in the stem and ab laceae with Geraniaceae. They however sence of mechanical tissue in the pericycle justify its treatment as an independent (Metcalfe and Chalk, 1950). family. Floral anatomical data on Limnan The available data on the floral thaceae are not a\ailable for comparison anatomy of Tropaeolaceae (Saunders, witli the data of other families. 1937) and Palynology (Erdtman, 1971) Palynological characters of Limnan do not permit us to draw any conclusions thaceae do not permit us to draw any regarding the relationship with Gerania conclusions (Erdtman, 1971; (Ironquist, ceae. 1968). Gytologically the basic chromosome Gytologically Limnanthaceae shows number n»6, 7 reported in Tropaeolaceac resemblances witli Geraniaceac in its basic (Darlington & Wylie, 1955) is unknown in chromosome number n = 10 (Darlington Gieraniaceae. However diploid chromo and Wylie, 1955). some numbers 2n==24, 28, 30 and 42 From the forerunning discussion, it reported in Tropaeolaceae arc also known is obvious that Geraniaceae exhibit simi in Geraniaceae, indicative of a probable larities as well as differences not only in relationship of both the taxa. chemical characters but also in otlu-r Viewing from different aspects it may characters with Oxalidaceae, Balsamina- be concluded that a relationship between ccae, Tropaeolaceae and Limnandiaceae. Geraniaceae and Tropaeolaceae cannot While tlie similarities indicate a probable be ruled out. close relationship between them, the diffe Geraniaceae and Limnanthaccae re rences jstify their recognition as inde semble each other in the negative reacdon pendent families. for Hot-Water test, HCN test, Juglone test ‘A’, Ehrlich test, Saponins and in the REFERENCES absOicc of aurones and raphides. The presence of tannins, ellagic add, querce- BAitBY, L. H. 1969. ManmU o f Cultivated Plants. 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