REVIEW O W LITERATURlE

REVIEW OF LITERATURE

The survey of earlier work reveals that there is insignificant informaiio

Anrlrrtrium in general and in-vifro propagation in particular compared to the size 64

i.\,
genus. However, a brief review of the earlier work pertaining to all aspects of the c&

is delt under the different titles here.

Botany :

The name Anthurium has been derived from the Greek word "Anthos" meanin.

tail, referring to the morphology of the spadix.

The genus Anthurium is known to contain about 578 species of' which 50 ar

under cultivation and out of which 10-15 are known to the commercial trade (Bailey

1963); of those that are in cultivation, there are several hybrids, as the species ar

compatible and seem to cross readily. The leaf blade of Anthurium undreanttm i,

drooping and cordate; spathe cordate -ovate, thick in texture, 15-25 cm long, widely ope

- spreading; spadix is 7.5 cm to 10 cm long. yellowish with white band marking the zon
in which stigmas show protogyny (Criley, 1988). Kamemoto and Nakasone (1963

described the flower of Anthurium andreanum as hermaphrodite with 2 carpelled ovar.

and 4 anthers.

Chistensen (1971) studied the morphology of growth and flower formation ir

Anthurium scherzerianum Schott and Anthurium andreanurn Lind. He describes that the

plants have a juvenile phase during which a vegetative bud is produced in the leaf axis.

but in the subsequent generative phase, a flower bud is produced in the leaf axis. These
 buds become dormant after initiation. The flower de\lelopnient depends on breaking of

donnmcy. Airrlturium schcrzeriarllmr plants which remain in vegetative phase for longer

than normal branch rapidly and are knonn as bush plants con~merciillly.

Croat (1980) made a thorough study regarding the flowering bchn\,iour of

Neotropical Anrhurium species and reponed that more than 30 species flowered

frequently under green house condition.

Sheffer and Croat (1983) evaluated the chron~osomesnumber in few Airrlrrrrirrrri

species from North, Central and South America. The chronlosome counts ranged from

2n = 24 to 2n = 66 with 2n = 30 being the most common number. Aneuploidy, pol!'ploid!,

and presence of B chromosomes are the basic features of the genus. Rehena Ali (1979)

studied the cytology of different Anrhurium species and reported a natural triploid

Anrhurium crys~allinum.

Presence of B-Chromosomes in Anrhurium n~arocqueanumwas reported by Mari

and Karnemoto (1 983). Their analysis revealed the presence of 1-3 B-chromosomes in the

complement. However, as observed, presence of B-chromosomes had no effect on the

phenotype.

Higaki and his co-workers (Higaki er al., 1984) studied morphological and

anatomical aspects of Anthurium andreanum.

Ray (1987) described and classified the leaf types in Araceae including Anrhuriuni

on the basis of their morphology.

 Breeding :
Evaluation and improvement of At7rli1rriunr clones were taken up by Kmenloto

and Nakasone (1963). Of 1 13 clones evaluated. 13 were recommended for conrnrrrcial

cut flower production including "Haga White", the orange variety "Nitta". and red

varieties "Kaumana", "Ozaki", "Kansako" No. 1 and "Hirsoe".

Breeding for the improvement of At~thuriunlwas carried out by Kamen~otcrzi al.

(1968). Two high yielding seedling selections, a white and a pink, were named "l'niwai"

and "Marian Seefurth" respectively and introduced into the trade. Three bi-cnlourcd

clones, white green (UH 8), rose ope1 green (UH 16) and coral green (UH 3 9 , were

released later.

Three new cultivars of Anthurizrm and 2 seedling selections, "Anuenuc" and

"Chameleon" as suitable for cut flower production were introduced by Kamernoto and his

co-workers (1969).

Anthurium scherzerianum X Anthurium wendiinzerii a new hybrid was de\,eloped

by Kamernoto and Sheffer (1978). The 2 species were crossed successfully to produce a

hybrid with a greyish - orange spathe. Other characteristics such as the length and colour

of spadix and the length and the position of the leaf blade were intermediate between the

highly contrasting characteristics of the parental species. Fertility in hybrids was very

good, indicating close relationship of the two species.

The techniques of cross pollinating Anthurium scherzerianum and also the

presence of recessive characters (A 3 with anthocyanin, a 3 without anthocyanin. B =

whole spathe coloured, b spotted spathe) were discussed by Maurer (1979). When the
parents were AdBb. the descendants \\'ere 9 red (AB). 3 rcd spots an white (Ahh) and 4

white ( a d and aabb). The deficit in \vhite plant was provisionally attribured to their lack

of vigour.

Studies pertaining to major spathe colour in A ~ l ~ k ~ r r i uwas

m taken up h!.

Kamemoto el a/. (1988). They identified 2 major genes M, 0 for all the major colours for

Anthurlurn ; for example red. orange. pink. coral and white. Red and pink result \\.hen

both M and 0 are present; orange and coral result when only 0 is present: tlie double

recessive mmoo results in white while pink is heterozygous tor both M and 0. Crosses

between two pinks will produce offsprings in ratio of 9 red. 3 pink to orange-coral and 4

white.

Best cultivars for cut flowers have been described by Gajek and Schwarz (1980).

In medium sized 'Iga Gold' with a shining red spathe and a white spadix with a yellow tip

and the compact "Ellrina" with a vermilion light salmon spathe and a sulphur yellow

spadix are revealed.

Schmidt and Lavterbch (1985) have recognised 2 cultivar groups. (a) miniature

cultivars - generally under 20 cm tall with narrow leaves and short petioles, including
cultivars "Oud Orange", "Renata" and "Amazone" and (b) larger plants with broad leaves

and long petioles including cultivars "Lachs", "Flamenco", and K26.Different cultivars

which are suitable both as cut flowers and pot plants have been listed by Bhan 8: Desai

(1989) which are mostly hybrids of different Anrhuriurn species, invol\ing mainly

Anthurium andreanum and Anthurium scherzerianum.

Cultural aspects :

A detailed study regarding the gro\laingof A~irlirtrittr~rin Trinidad and Tohogo hy

Rapsey and C m (1969) included propagation and cultural practices, pests and disease

control etc. Christensen (1973) reported that root trimming of Anrhurilmt scher:crrnriurn

while picking reduced final plant size. but did not retard flowering. Lefring (1975)

concluded on the basis of his experinrent that plants grown under shade receiving at least

45 per cent of available light in the green house resulted in increased growth rae nnd

average flower production.

Higaki and Rasmussen (1979) reported that plants of "Ozaki Red" when sprayed

with PBA, BA or Ethephon at a concentration of 100, 500. 1000 or 1500 mgl" induced

adventitious bud formation. Maximum shoot formation was produced with BA at 1000

rngl" (3.6 shoots / plant ) followed by PBA at 1500 mgl-'(2.2 shoots I plants) and

Ethephon at 1500 mgl"(l .8 shoots /plant). Control plant exhibited no adventitious shoot

formation. Work on spacing was taken up by Higaki and his co-workers (1979): they

have recommended a spacing of 45 x 45 cm to accommodate 12000 plants per acre or a

closer planting of 30 x 30 cm, to provide 25,000 plants per acre. When closer spacing is

followed, for maximum production, heavy pruning should be done every fourth year, to

provide proper air-circulation. Rigid leaf pruning and spray schedule should be followed

to control diseases etc, They also suggested the application of fertilizers such as 5-10-10

(N.P.K.)
or 10-20-20 (N.P.K.) or 16-16-16 W.P.K.), at the rate of approximately 135 kg

of nitrogen per year per acre. Slow releasing or pelletised fertilizers are bener as they

cause less damage.

 The work of Vooyt (1979) rc~ealedthe effect of saline water on the de\~eloprnunt

of Atrthuriunr andrcunum. The cut flower yield of Atrthuriutn declined propressi\.ely as

salinity of the water increased. \\:ater containing sodium chloride was particularll,

detrimental. There was a marked difference in salt sensitivity between cultivars. \\'hen

desalinized water or rain water is to be used, extra calcium application is necessq .

In trials with cv. "Brazil Red" md "Antiqua". Strauer (1980) reported that culture

on rock wool (blocks & mattings) is satisfactory, if a suitable nutrient solution nith a

high K content is used.

Hetrnan ct al. (1981), studied the effect of substrate on physical propenies of

Anthurium ondrcanum assessing for plant height, number of leaves and leaf area co-

efficient. The best results were obtained on a substrate of peat + Sphagnum, peat + perlite

in equal proponions.

Studies pertaining to the effect of temperature on Anthurium andreanum hybrids

were taken up by Schenk and Bnrnden (1981). Five cultivars were exposed to air

temperature of 13 OC, 19 OC or 22 OC ~{itha constant peat substrate temperature 22 OC.

Cultivars "Homing Orange" and "Horning Rubin" were normally temperature neutral but

in "Homing Rubin" the total number of flowers per plant was reduced at 22 'c. For the 3

light Red "Vogel" cultivars yield and quality of flowers were best at 19 OC at nhich

temperature "Vogel" 202 produced 7 flowers per plant, the highest yield. At 10 OC

"Vogel" plants died, and at 13 OC they showed some leaf necrosis, Plant vigour increased

with raising temperature.

 Turski and his co-workers (1983) studied the effect of different substrates on

growth of Anthurium andrcanuni, The best substrates were those in which the basic

component was more of peat.

Use of powdered bark as a substrate for growing Arirl~rtri~tni

in pots was taken up

by Tesi and Faro (1985). They concluded that Anrhurr~tmcould be successfully gronn on

a bark based medium.

The work on seed sowing and gemination was done by Beele (1971). The best

germination resulted from picking the berr~esat the orange - red stage and fernicntlnp
them for four days in water at 22 "C to separate the seeds from the pulp. Storage was

sometimes necessary because berries from the same inflorescence ripened at different

times. But the maximum time for storage of fermented seed in water was five da!,s But

this reduced germination from 99% to 80% and 10 days in water ~t reduced it to 53%.

The best substrate for germination (95%) was peat. Seedling growth, especiali! root

development was better in white peat t perlite than in peat + perlite or coiferous l~ner+

perlite.

Fresh seeds of Anthurium scherzerionum hybrids germinated in light or darkness

at 10-35 OC. Germination occurred after 5 to 7 days after drying and storing at 20 OC for

24 h. Germination under favourable conditions were 70-75% (Bachthaler, 1977).

Bachthaler (1978) studied the germination of Anthurium scherzerianum hybnds at

different stages of beny ripening. Seeds extracted from (1) green / unripe (2) reddish 1

half ripe (3) red ripe and (4) reddish b r o w or over ripe berries were placed in petridishes

on damp sterile sand kept at 25 OC for 12 h of light. In the first three groups all the seeds
 showed gemination but in group 4 onl! 42 % gcrniinatcd. Group 2 & 3 were the first to

germinate and were the most suitable for commercial seed production.

Work on storage of seeds of .-!nthurium schcrzcrionum was also done h!,

Bachthaler (1979). Seeds stored within the berries at 3 "C suffered cold injury and fungal

infection. The best storage temperature was 10 OC. and after 6 weeks 60% of seeds

germinated. Seeds from berries treated with thiram Just before storage. 95% germinated

after 12 weeks at 10 'C and 60% after 16 weeks. Method of seed extraction of Anthurrtrrrr

schcrzerianun~hybrids was described by Maurer and Brandes ( I 979 a). The seeds treated

with 13% sodium carbonate at 20 OC for 25 hours or in 6% sodium carbonate + 1%

pectinase solution at 26-36 OC for 5 hours were the 2 methods resulting in seeds being

clean enough to sow singly.

Maurer and Brandes (1979 b) also worked on storage of seeds of Anthrrrium

scherzerianum hybrids. Seeds were stored as spadices, in berries or as cleaned seeds in

dry conditions. Seeds left on spadices or berries become too dry or rotted. In cleaned

seeds, germination decreased. Some seeds stored at 20 OC germinated but seeds stored at

5 OC became non-viable. Seed stored in water rotted.

Studies on storing of seeds of Anthurium scherzerianum hybrids was done by

Bachthaler (1980). Berries stored at 10 OC for upto 6 weeks germinated in 4 days where

as those stored at 5 OC and 2 OC took 5 and 7 to 8 days respectively. Germination was

100 % in seeds from berries stored at 10 OC for 3 weeks. Seeds from berries treated with

thiram germinated well (in 4 days) after 16 weeks storage. After 12 weeks storage at
 10 k seeds from untreated berries failed lo geminate, whcreas pemllnalion \\.as ahove

90% in seeds from treated berries.

Szendel and his CO-workers (1981) evaluated the conditions for gerniina~lonof

Anrhurium arrdreanunr and Arrrlrrtrirrnr scherzcriunum seeds. Seeds of Arrrlrrrr~~mr

andreanum harvested at maturity stages and those of Anrhrtrlunt ,sc/rer:crrrmrtnt at one

maturity stage (light orange) were fcrminated on 3 substrates at pH rangmg from 4 to 8 In

light or darkness at 18. 24 or 28 'c. In Arrrlrrrrium undreunum the best perminatlon was

obtained on peat substrate at pH 4 or 5 in l~phtat 28 'C using seeds han ested at an early

maturity stage (yellow - green to light orange). Anrhurium scher:errcmunt seeds

germinated best on peat having pH 4.

Flower development :

Studies relating to morphology and shelf life in flowers of Anthuriunr were

conducted by Watson and Shirakawa (1967) ; they described the stages in the

development of individual flowers on the spadix of cv "Ozaki Red". They also suggested

that picking when the stigmas are receptive should be avoided as water loss is greater at

this period. Dipping the spadix in paraffin wax was found to reduce water loss

considerably and enhance keeping quality of flowers.

The work related to physical characters of Anthuriurns to its vase life was done

by Akarnani and Goo (1972) and they concluded that vase life of cv "Kaumana" flo\vers

varied from 8-10 to 18-21 days in water at 78-87 OF -

and 54 74 % RH.Vase life was

inversely related to petiole length and diameter and flower weight.

 lwata and his co-workers (1979) studied tllc anthocyanins of Aml~rtrirt~~r

undrcanum cultivars. and identified them as pelargonidin - 3 - rhmnosyl glucoside and

cyanidin - 3 - rhamnosyl glucoside. Both pigments were present in red (spathe colour) as

seen in the cvs. "Ozaki". "Kaurnana", "Kozohara". "Kansako No.lt' and "Nalawwa".

And in the pink cv. "h~larianSeefurth", the Orange cv. "Nirta" and coral coloured cv,

"Tateishi Coral" contained only pelargonidin 3- rhan~nosyl glucoside. Later the! also

reported that a predominance of cyanidin - 3 - rhamnosyl glucosidc resulted in p ~ n kto

dark colours, where as a predominance of pelargonidin 3 - rhamnosyl - glucoside

resulted in coral to orange colour (Iwata el a/., 1985).

Harvesting and grading of Anthuriums, however. was done by Singh (1987).

Flowers of commercial value is harvested when approximately three - quarter of the

stigma along the spadix becomes receptive. Different grades are given based on total

spadix length and width of the spathe.

Grade Spathe measure

Miniature c8cm

Small 8 to 10cm

Medium 10 to 13 cm

Extra large > 15cm

Akamine and Goo (1981) worked on controlled atmosphere and storage for

Anthurium flowers and found that cold storage at 13 k successfully extended the storage

life of cut Anthurium flower for cultivar "Ozaki". But where refrigeration facilities were
 -
.mot available, storage in 2 10% 0: could he used udvantagcously at anrhient [enrpenture

of 24-25 'c.

Work by Kalknian (1983) revealed that vase life wns tlie longest \vitli flo\\.ers cut

when the spadix was almost coniplrtely ivhite. The average vase life for flowers cut at

this stage in winter and sumn~er\vu 11.5 to 25.2 days respectively. Cv "Avo-Cinthn" and

"Avo-Ingrid" had the largest a\,cnge vase life of 31.3 and 25.4 days respecti\.el!, in

winter and 29.1 and 28.9 days respectively in summer.

The use of waxes to extend post harvest vase life of Anthuriurns wns suggested

by Paul1 (1983) ; of the 8 products used to coat the flower. FMC-819 (Carnuba hased

wax) was most effective, increasing the vase life from 18 days in the untreated control to

36 days.

Paul1 and his co-workers (1985) described the physiological changes associated

with senescence of cut Anthurium flowers. Silver pulsing (for 40 minutes) of Anllt~~rlrtrn

andreanurn, cv. "Ozaki Red", flower stem was done to modify senescence process.

-
Florets on the spadix continued to open for 5 10 days after harvest in both treated and

untreated flowers. In both respiration rate was low until senescence began 8 days after

harvest, The rate of increase in respiration of silver treated flower was lower than that of

cantrols. In all cases ethylene production remained low throughout the post harvest life of

the flowers. Ten days from harven. spathe colour began to change from red to blue with

no significant changes in anthocynin ratios. Tissue pH rose from 5.2 to 5.6. The

concentration of tissue phenolics increased during senescence and intensified the colour

change by co-pigmentation. Tissue starch level declined by about 25 %. The ratio of

 sugars in the stem. spathe and spadix remained constant with a slight declinc in

concentration during post - harvest life. Silver pulsing of the stern reduced slem plugginf

and thus reduced the rate of change of all the senescence process observed.

Paull (1987) also studied the effect of storage duration and temperature on cut

Anrhurium flowers. The optimum storage temperature for A~rrlrrrriumat~drcanuntflowers

of the cultivars "Kaumma", "Nitta" and "Ozaki" was betweeti 14 and 17 "c. A silver

nitrate pulse (4 pM for 40 mins) given immediately afier har\,est increased post harvest

life of stored flowers. Maximum vase life was achieved with Ag + treated flowers

Micropropagation

In-Vitro seed germination :

Swaminathan (1986) reported that the Nitsch's medium was found to be the best

for seed germination (Anthurium andreanum cv 'Red'). NAA (0.1, 1.0 and 5.0 mgl") did

not have any effect but IBA (0.1, 1.0 and 5.0 mgl") delayed seed germination and

promoted the subsequent growth of seedlings. GA, also did not influence seed

germination but enhanced the subsequent growth of seedlings. The media containing

thiamine and banana pulp showed good seed germination and subsequent seedling

growth.

Zens and Zimmer (1988) obtained callus mediated multiple shoots from seeds of

An~huriumscherzerianum on Nitsch's media, while the reports of Randhawa (1990)

 showed the euly seed gemination w d development or first Ieal'stogc in Nitsch niediunr,

whereas the subsequent seedling growth was completely inhibited on Morel mediunl.

In-Vitro propagation through tissue :

Explants :

For in-1-itro propagation of Anthuriums, the segments of leaf, petiole, spathe,

spadix, pedicle. vepetati\.e buds, shoot tips and roots are used as the source of explants.

Leaf lamina segment have been widely used as esplants by many research

workers (Pierik. 1976: Navak and Nepustil, 1980 ; Geier, 1982 ; Eapen and Rao. 1985 :

Geier, 1986 a : Keller er al., 1986: Geier, 1987 ; Kuehnle and Sugii. 1991 ; Singh and

Sangrna, 1991 : Kuehnle. et. al. 1992 ; Nirmala and Sinph, 1993 and Matsumoto et ul.,

1996) in different species of Anthurium. Callus mediated plantlets were obtained from

leaf explants by Novak and Nepustil(1980) and Geier (1982).

The presence of the midrib in the leaf lamina segments has been reported to have

influence on callusing in Anthurium andreanum in both liquid and solid media (Pierik,

1976). The intensity and frequency of callus was found to be on the highest in leaf lamina

segment cultured with midrib viens in Anthurium petulum (Eapen and Rao. 1985).

However, Geier (1986a) found that the presence or absence of midrib had no effect on

:allusing in leaf explants of Anthurium scherzerianum. Position of the leaf from where

h e explant is excised and the surface (abaxial) touching the media was also imponant for

3etter callus production (Geier, 1986a). Leaf explants of Anthurium andreanum formed

:allus in 1.5-2.0 months on Murashige Skoog medium supplemented with 2 mgl" kinetin

:Keller et al., 1986).

 Regenerative callus was ohtained from leaf explants of ..ltirlrrrrirmr atidrrnrtrrrri

cultivars (Kuehnle and Sugii. 1991). Callus derived from leaf lamina segnients on

Modified Nitsch medium found to be poorly regenerative (Singh and Snngma 1991).

Kuehnle et al. (1992). obtained translucent embryonic callus at the basal ends of the leaf

blade explants. obtained fiom in-~~rrro

grown plants, with In a nionth of culture under dark

conditions in four c u l t i ~ x sof At?rirrtriumandreanum. viz.. UH 780. UH 965. UH 1060

and UH 1003. Nirntala and Singh (1993) also obtained regenerative callus on Nitsch

medium suppleniented \\ith BA and 2.4-D from leaf lamina segments. Matsumoto t r al.,

1996 derived somatic embryos from in-vitro cultured leaf lamina of Atiriitrrrum

andreanum cvs. 'Anuenue' and 'Toyama peach'. Mamet (1980) obtained the complete

plantlets from culture of meristem tissues in Anrhurium undreantrm.

Petiole segments of Anthurium scherzerianum showed low regeneration capacity

as compared to that of spadix segments (Geier, 1982). However, Eapen and Rao (1985)

obtained good regeneration in At7thurium patulum and Kuehnle and Sugii (1991) in

Anrhurium andreanum using the same explant.

Geier (1982) reported that spadix fragments of Anrhurium scherzerianurn

cultivated in-viho conditions. showed high capacity for regeneration than segments of

leaf, petiole and spathe under darkness on Modified Nitsch medium. Similar results were

obtained by Singh and Sangma (1991) and Nirmala and Singh (1993) in Anthirrium

andreanum.

Zens and Zirnmer (1986) reported that formation of callus and adventitious shoots

')y shoot tip explants were increased significantly by changing NHp-N : NO3-N ratio from
 1 :1 to 1 :5. Regenerative callus \\.a%
obtnined by Soczeck and Hmipel ( 1989) from s~nglc

node fragments of in-vi/r.o grown shoots of Anrhuriunr rmdrcor~rmrwhen cultured on half-

strength Murashige & Skoog medium containing various concentrations of dityerent

cytokinins. Zen cr al. (1993). reported the advantages of light and durliness In induction

and growth of callus as well iu the fwther growth of the plantlets derived tiom

adventitious buds. Similar results were also reported by Nirn~alaand Singh (1993) on

Vacin and Went liquid medium.

Nutrient medium :

The success of plant tissue culture as a means of plant propapalion is greatly

influenced by the nature of the culture medium used. Plant tissue culture media pro\.ide

major and minor nutrient elements and carbohydrates, Improved results were obtained by

providing trace amounts of organic compounds, notably vitamins. amino acids and plant

growth regulators (George and Sherrington. 1984).

Most of the reports on Anthuriums are based on Murashige & Skoog and Nitsch

media. Pierik et al. (1974), cultured embryos on Modified Murashige & Skoog med~um

containing half strength macroelements, microelements, sucrose (3%) and organic

constituents (except adenine, IAA and kinetin) and difco-agar (0.07%). Further. he

modified the macroelements of Murashige & Skoog medium for callus induction, callus

subculture, sprout regeneration and rooting of sprouts, but microelements and other

organic constituents were remained without change. He also found that lower

concentration of NH4N03was an essential factor in the induction of adventitious shoots

in callus tissue of almost all genotypes. Fersing and Lutz (1977) reported that medium
.applemented with yeast extracts stimulated shoot growth in ..lrr/huritm~.rc/icrccrmrrrm

but restricted in Anrh~irrrimundreartrrni whereas according to tlre repon of Eapen and R

I
(1985) half strength Murashige Skoog escept Fe EDTA. along with vitnn~insrvd 0,6

per cent agar was found to be most suitable for the production of callus mediated plant

regeneration from leaf segments ~ i t midrib

h and petiole segments in A ~ ~ ~ h ~ r r r r r t i i ~ i n r r ~ l ~ t r i i

For the culture of spadix segments of Anthurium schrrscrrurt~tm.Geier (1982)

employed Modified Nitsch medium (Nitsch, 1969) which contained low le\rls of'

ammonium nitrate (100 mgl"). Funher, he also made standard modifications to suit the

culturing of the leaf segments. He also indicated that 200 mg/l NH, NO3 was most suited

for callus and shoot formation, where as. 720 mgl" was effectwe for root formation and

changing NHd -N: NO, -N ratio from I:] to 1:5 increased the formation of callus and

adventitious shoot from shoot tip esplants in liquid medium (Zens and Zimmer. 1986).

Same workers (1988) also reported that Murashige & Skoog solid medium supplemented

with BA and NAA was found to be good for the production of callus mediated multiple

shoots from seeds of Anthurium scherzerianum. Root formation was not affected by

varying conceneations of ammonium nitrate in Murashige & Skoog medium (Lightbourn

and Prasad, 1990). Good and regenerative callus was obtained from leaf explanrs of

Anthurium andreanum on Modified Pierik medium containing 2, 4-Dand BA after 2 - 3

months. Whereas petiole explants callused better on Pierik, Modified Pierik and Finnie

and Van Staden media. While multiple plantlets derived from callus was on Kunisaki

medium (Kuehnle and Sugii, 1991).

 Modified Nitsch medium \\.as found to be good for the regenention of plantlcls

&m spadix segments (Singh and Sangma 1991). Leaf explants when cultured on niedia

~ t a i n i n gthe combination of 2 per cent sucrose and I per cent gelrite produced more

s on half - strength Murashige & Skoog medium \\.~th0.7 per cent

mmatic e m b ~ o than

k t o agar (Kuehnle c/ a/.. 1992).

Murashige & Skoog medium supplemented with 3 per cent glucose had the

greatest inductive effect on callus formation as compared to that of sucrose (Cen cr a/.,

1993). Leaf segments formed good callus on Nitsch medium containing BA and 1.4-D .
whereas vegetative buds callused in Vacin and Went liquid medium (Nirmala and Slngh.

1993). Reduced concentration of Murashige & Skoog ma,ior nutrient elements and

sucrose did not significantly influence the production of multiple shoots (Sreelatha cr a / . ,

1994).

Growth regulator :

Growth and morphogenesis in i n - v i m cultivars are regulated by the interaction

and balance between the growth regulators supplied with the medium and the growth

substances produced endogenously by cultured cells. Besides, many synthetic regulators

may in fact modify the level of endogeneous growth substances. some times in a fashion

which is heritable over many cell generations (George and Sherrington, 1984). The

breakthrough made in tissue culture is the discovery that root and shoot initiation is

basically regulated by interaction between two hormonal substances namely auxins and

zytokinins by Skoog and Miller (1957).

 ;16.Ilus induction and subculture :

Generally a high concentration of auxin and a low concentration of cytakinin in

& basal medium promotes cell proliferation and callus fonnation. Pierik il/ a1 (1974).

reported that in Anthurium andrearlllm different organs of adult plants were capable of

forming callus at 1-5 nlgll PBA added to Modified Murashige & Skoog medium. Callus

of leaf. spathe petiole and pedicel could be maintained on the basal medium supplied

with PBA I mgl" and NAA 0.1 mCl". Modified Murashige 8: Skoog liquid medium was

found to be good for callus growth Presence of NAA in the solid medium ~nducedroot

formation (Pierik. 1975).

Further Pierik (1976) reported that callus induction can be achieved with PBA

( I mgl"), 2,4-D (0.08 mgl") and can be subcultured with PBA (1 mgl'l) in the medium.

With BA (1 mgl") and 2,4D (0.1 mEl'l) in Nitsch medium can induce callus from spadlx

segments (Geier, 1982) and leaf segment (Geier, 1986 a) of Anlhurium scherzerianum.

Leaf, pedicel, spathe and petiole segments produced pink coloured callus in Anthltrwm

petulum (Eapen and Rao, 1985).

The leaf explants of Anthuri~tmandreanum formed callus in 1.5-2.0 months on

Murashige & Skoog medium supplemented with 2 mgl" kinetin (Keller et ab, 1986).

Variation in ploidy level was observed in the case of callus derived from shoot (Geier,

1988). Seeds of Anthurium scherzerianum produced caulogenic callus or callus with new

&oots and productivity depends on genotype and it decreased with increased NH4 : NO3

4jqati0 in the medium (Zens and Zimmer, 1988). Callus production increased as cytokinins

h c e n t r a t i o n was increased (0.25 mgl'l to 2.0 mgl") in Anthurium andreanum (Soczek

77
 m d Hempel. 1989). Best callusing was seen on nrcd~unisupplemented wlth 0.5 nigl"

2.4-D in the case of Anrhririum atrdrcatr~tmcv. 'Tulip' whereas In the case of cv . 'Tropical

Pink' 0.05-0.5 mgl" 2.4-D was found to be effective (L~ghtbournand Prasad. 1090)

Embryogeneic callus was obtained from basal ends of cut leaf' blade esplants wlthln one

month of culture in the dark on Murashige & Skoog nied~uni supplemented w~th

1-4 mgl-' 2.4-D and 0.33-1.0 mgl" kinetin (Kuehhnle cr u l . 1992) Medium

supplemented nith 3 per cent glucose was found to be goad for callus rnductlon as

compared to that of sucrose (Cen er al., 1993). Murshige & Skoog niedlum

supplemented ~ 7 t hNH, NO3, BA nnd 2.4-D induced callus from leaf midrib and spadix

under dark, but. vegetative buds needed light (Nirmala and Slngh. 1993). Murash~ge&

Skoog medium containing 1 mgl-' each of 2-iP and BA induced callus (Sreelatha el a/.,

1994). Regenerative callus was obtained on the media supplemented with 2-iP In

Anthurium andreanum cvs. 'Hazarija' and 'Ingrid' and .4nthurrum scherzerranum

cv. 'Belinda' (Yu and Paek, 1995).

Shoot bud differentiation :

Low auxin and high cytokinin levels in the medium is a general requirement for

shoot bud differentiation, though, Pierik et al. (1974), reponed that sprout formation

occurs in callus spontaneously on transferring the cultures to light ; Pierik (1976)

obtained good plant regeneration from callus on solid Murashige & Skoog medium as

~ m p a r e dto that of liquid medium supplemented with yeast extract, stimulated shoot

growth of Anthurium scherzerianum but restricted it in Anrhlcrium andreanum (Fersing

m d Lutz, 1977).
 Similarly, medium containing kinetin (3 mgl") was thund to he more effectlve in

&acing shoot differentintion as compared to that of BA and 2-iP (Lefring and Soede,

W9).Subculruring of callus (3-6 times) on homione free Modified Nitsch niedum

m l t e d in the production of true to type multiple slroots at 1000 lux light for I J ho& A

&y (Geier, 19s:).

Eapen and Raa (1985) reported that best response for shoot d~fferentlationwas

obtained with BA (1.0 n ~ ~ l and

" ) 2.4-D (0.1 mgl"). Kinetin and 2-iP stimulated shoot

differentiation to a lesser degree, while zeatin was ineffective in the case of Arirlirnrlmi

In Anthicrrum sclrcrzerianunr shoot regeneration was encourage from callus of leaf

cxplants when medium was supplemented with BA 0.5 mgl" and reduced level of

NH4N03 (200 mgl'l) (Geier, 1986 a).

Maximum shoot multiplication was observed on Murashige & Skoog medium

supplemented with NA.4 and BA from seeds of Anthuriuni scherzerianum (Zens and

Zimrner, 1988). Same result was obtained by Lightbourn and Prasad (1990) in

Anthurium andrsanum c ~ s'Tulip'

. and 'Tropical Pink' at 0.2-08 mgl" BA. Sreelatha er a[.

(1994), reported that kinetin (2 mgl") and BA 1 mgl" were effective in the production of

shoots. Treatment with kinetin did not produce callus, whereas BA and 2-iP induced

callus. Similar results were reported by Yu and Paek (1995) in Anthurium andreanurn

cvs. 'Hazrija' and 'Ingrid' and Anthurium scherzerianum cv 'Belinda'.

 Rooting :

Geier (1987) reported thc highest rooting of shootlets on harmone free

Nitsch medium at a light intensity of 200 lux for a duratio~iof I4 liours I day. Rooting

occurred after four \\eeks on cytokinin free medium in tlie case of .41trlrrn.iro?1

mdreonum (hfiet oi,l983). Medium containing 720 mgl" NH,NO, without harn~ones

accelerated root formation (Geier. 1986a). Root formation M8asno! influenced by varying

concentration of ammonium nitrate (Lightbourn and Prasad. 1990).

Cytology :

Determination of the chromosome numbers are the most common type of' cytological

studies made in the species of Anrhurrum. Among the 600 species kno\\n in the genus,

chromosome numbers are available for a little over 100 species.

Major contributions in the above field have come from Gaiser (1927). Pfitzer

(1957), Shanna and Bhattacharya (1966), Merchant (1968). Sheffer and Kamcmoto

(1976 a), Keneko and Kamernoto (1978), Ali (1979), Mari and Kamemoto (1983) and a

few others.

Cytological studies in the genus was first initiated by Gaiser (1927). He has

nponed the chromosome number in 43 taxa including several hybrids. Most of the counts

of the author have been confirmed by later workers.

 Mookerjea (19551 studying somatic chromosomes In a fe\v species has reponed

2n = 34, 35, 56 and 63. However. die author has not offered an\ plausible esplanat~onfor

the odd chroniosome counts of 35 and 63 in these plants.

Meiotic counts examined by Plitzcr (1957) revealed the chromosome nunibcr of

n = 15 in the species of Anthurruri~schcrrcriunum, Antlturrrtnr scllo\c ronrtm, Anlhrrrrrrm

domin~scence...lnthuriutit consobrir~um,Anthurlurn wurocqrrco~rrrn~,

Arirltlrrrurn radicans,

Anthurium rvrrchil, drrthurium arrdrcanum, Anrhurrunr per~/uphj,llrrm, Ar~tlrrrrr~tm

crystallinurn and Antklrrrum forgetti. However, n = 24 and 30 u'ere also found in

Anthurzum sca~rdensar~dAnlhuriun~digitatum, respectively. In addition. B chromosomes

were recorded in both PMCs and somatic cells of Anrhurium cr?~s/allinun~

and Anrhltrrum

forgetti.

Sharma and Bhattacharya (1961) investigated 3 species \'Iz.. Anrlrirrrum g1a:rovir.

Anthurium parulum and Anthurlum variable in which 2n = 28. 30 and 69 were recorded

respectively, Interestingly, all the three taxa revealed varying number of chromosome

fragments. The pollen abnormalities in these taxa were attributed to the presence of B

chromosomes.

Merchant (1968) studying the cytology of 12 species has recorded the sornarlc

chromosome numbers of 2n = 30 in 6 taxa viz., Anthurium ucutum, Anthurlum

aystallinum, Anthurium hurrisii, Anlhurium imperial, Anthurium nricrophyllunr and

vlnthurium subsignarum ; 2n = 40 in Anthurium grucile and Anthurium scolopendrrum

, ,nd2n = 60 in Anthurium undarum. The highest somatic chromosome number knoun so

% in the genus, 2n = ca. 124 in two species viz., Anthurium lucidum and an
undetermined species also come from this study. Hc has also rccordcd an odd somatic

number of 2n = 45 - 47 for the species Anrhuriltm sc~carilcns.Varying numher of

chromosome fngments. I in Antli~rrirmrnricrophyllrrni and diithuriuni arrhsi~nurunr.2 in

Anth~rriumcn~srallinrtnim d Antliroium imperial and 5 in .diirhurium Irtrrr.isii ha\,? also

been recorded. Anthurilmi undarrrnr also revealed the presence of I -B-chroniosonie in its

somatic cells.

Extensive study of somatic chromosome numbers of 63 species was taken up by

Sheffer and Lamemoto (1976 a). They observed that the most common number

encountered in these plants is 2n = 30 with only a few showing 2n = 60. Though.

karyotypes ate not described for any of the taxa examined. the authors mentioned the

presence of 'Loosely held satellite' in the complements of Anthuriurn ucrirrrlum.

Anlhurium scandens and Anthuriuni scolopendrium.

A cytological finding of importance has come from the work of Ali (1979) who

has reported a natural triploid in Anlhurium crysrallinum (2n = 45) from a collection

maintained at NBRI, Lucknow, India. On the basis of chromosome morpholop. the

materid was identified as an autotriploid. But, instead of the expected 15 trivalents at

meiosis only ten were found along with several bivalents and univalents. The situation

was explained by the author as due to inadequate chromosome length of some pairs.

Keneko and Kamemoto (1978) after studying two cultivars of Anikrrrium

zndreanum, the 'Kaumana' and 'Uniwai', varieties have recorded 2n = 30 and 2n = 30 +

2B respectively. The basic karyotype comprises 4 relatively large chromosomcs. 2

ratellited and 24 small chromosomes.
 A detailed account of the occurrence and tnnsnIls\lon of B-clironio~onics ~n

Anihrrrrum ~tarocqucar~rtrn
was gncn b) h4ar1 and Kmienioto (1983) They also stated

that tlie presence of B-chromosomes d ~ dnot effect the n~orpholog\ of the plan~s

However, the! had some effect on the b~valent format~on resultrig In certaln pollen

abnormallt~es

The work pertalnlng to basic chron~osomenumbers of 14 genera ot Ardceae ua5

done by Hotta and OLada (1987) They are of the oplnlon that ~ ~ t h each
l n genus the

bas~cnumber 1s stable (n = 15) and could be the bass of assessing ~nter-relatlonshlpsIn

An~hurrurn Srudles on ,Inrhurrunt andreanurn collected froni eastern Hlmala\ as h)

Sengupta and Chenrl re\ ealed that 2n = 30 as the chromosome number for the taxon The

somatlc compliment comprised of 4 fairly large, 22 medlum s~zedand 4 small sued

chromosomes

Chromosome numbers for a few specles are also lcnown from the work of

Campbell (1 905), Dela! (1 947), Haase-Bessel (1 928), Kurakuba (1 940), Malvesin (In

Fedoerov 1969). Nerl~ng(1969), Tsuchlya and Takada (1962), Sharma and Bhanacharya

(1966) etc

Phytochemistry :

Very ltttle chemotaxonom~calwork has been camed out tn the family Araceae In

Oeneral and genus Anrhrtrlum In part~cular However, a few reports arc known such as

-
Bate Smlth (1 968) who exarnlned 24 specles of Anaceae and ldentlfied the presence of

rercltm, kaempferol and procyanldln In most of them. The author stated that the farnlly
d d possess diverse phenolic compounds because of the prcscnce of sc\,cnl

bharacteriscd compounds. .

According to \Xrilliam cr (11 (1981), the most chmacterist~cs of flavano~d

mnstituents of the fatlily Araceae are the presence of fla\ one C-glj cos~des The! are

found in more than 50 % of the tam examined while flavo~iolswere found In about 25 %

of the plants. These authors have also found cafeic ester sulphates as n common

constituent of..lr~fhuriu~rr
iiookeri.