Amophophallus paeoniifolius

A Synthesis Paper

In Partial Fulfilment of

the Requirements in

Biology III
INTRODUCTION

Elephant foot yam, Amorphophallus paeoniifoliu, locally known as “Pungapong” in the

Philippines, is an herbaceous and perennial C3 crop, originating in south eastern Asian countries

such as the Philippines, Indonesia, Malaysia, Bangladesh, India, and China as a native crop

(Chandra, 1984; Sugiyama and Santosa, 2008). As it is a rich source of carbohydrate, protein,

minerals like calcium, iron, phosphorous, vitamin A, B, C, flavonoids, and fiber, it is used by

many countries as a local staple food and as medicines (Kay 1987; Shilpi et al. 2005). Due to its

high production yield and as a major ingredient for various Indian cuisines, Elephant foot yam is

commercially cultivated. In northern and eastern states, wild and local grown cultivars are

generally used for making vegetable pickles and medicine preparations for various ailments. This

crop also offers export potential since it is not commercially cultivated in other countries (Misra

and Shivalingaswamy, 1999; Misra et al., 2001). Along with the crop’s countless benefits,

Elephant foot yam is known for its intriguing “penis shaped” flower. With the aid of following

researches: Growth and Productivity of Elephant Foot Yam (Amorphophallus paeoniifolius

(Dennst. Nicolson): an Overview (2009) by V. Ravi, C.S. Ravindran and G. Suja from the

Division of Crop Production, Central Tuber Crops Research Institute and Morphological

variability in 17 wild elephant foot yam (Amorphophallus paeoniifolius) collections from

southwest India by S. R. Anil, E. A. Siril, and S.S. Beevy, the growth, morphology, and the

nutritional and medicinal uses of A. paeonoiifolius were assessed.

(about elephant foot yam-facts)

Origin

The genus Amorphophallus originates from and is mainly distributed in the Old World,

especially in the tropics from Africa to the Pacific Islands, but also extending to temperate areas

in China and Japan. The genus is not well known, the total number of species is possibly more

than 170. A. paeoniifolius occurs wild and cultivated from Madagascar eastwards via India and

South-East Asia to Polynesia, including also southern China and northern Australia. Because it

easily escapes from cultivation and naturalizes, its exact origin is unknown.

Distribution

Common in most or all, provinces of Luzon and in Mindoro, in thickets and secondary

forests, along roads, trails, etc., at low and medium altitudes in settled areas.

- Occurs in India through Malaya to Polynesia.

Conservation status

As categorized by International Union for Conservation of Nature (IUCN), the Elephant’s

foot yam conservation status is least concern. The crop has high productivity and wide

agroecological adaptation and exhibits suitability for the agroforestry system.

Growth
 Elephant foot yam plants grow well in medium to light soils (coarse-textured sandy soils)

with adequate amounts of organic matter because they prefer well-aerated soils. The crop can

tolerate temporary flooding, but anaerobic waterlogging causes corm rot.

Corm dormancy

Amorphophallus corms exhibit dormancy for about 3 - 4 months after harvest. As a result

of this, planting and harvesting are done at a particular time of the year. Amorphophallus is

propagated by corms as such or by cut corm pieces having a part of apical meristem. Sprouting

percentage was more (98%) with top cut portion of corm than the cut corms from lower half of

the mother corm (Dhua et al., 1988; Nedunzhian and Mohankumar 1997; Mondal and Sen,

2004).

The bottom portion of the corm is not used generally as planting material due to lower

efficiency of sprouting (Dhua et al., 1988; Nedunzhian and Mohankumar, 1997; Mohankumar

and Ravi, 2001). Therefore a greater portion of (about 25%) of the harvested produce is again

lost as source of planting materials.

Ethrel or ethephon was reported to induce early sprouting in Amorphophallus corm

(Dhua et al., 1988; Bala Nambisan and Indira, 1992). Treating cut pieces of corms from lower

half with chemicals significantly improved sprouting, subsequent growth and yield. Among the

different chemicals used, thiourea, potassium nitrate and CCC were effective in promoting

sprouting.

Thiourea (200 ppm) and KNO3 (1000 ppm) and kinetin (5 ppm) increased corm

sprouting by 24.3 – 92.0, 17.8 and 13.4% respectively as compared to control (Table 4, Dhua et

al., 1988; Kumar et al., 1998). However, mean corm weight was greater in plants from corms
treated with thiourea (100 ppm), potassium nitrate (KNO3) (500 ppm) and CCC (0.02 ml l-1)

yielding 722, 821 and 806 g per plant respectively (Dhua et al., 1988).

However, corm yield per ha did not increase significantly in plants from corms treated

with chemicals, as compared to plants from untreated corms. Exposing the whole corms to

smoke for 6 h per day for 6 weeks increased sprouting by 58.3% as compared to untreated corms

presumably due to ethrel in smoke.

Similarly exposing the corms to 45oC for 6 h per day for 3 weeks increased sprouting by

83.3% as compared to untreated corms (Mohankumar and Ravi, 2001). Temperature (32°C) and

thiourea had greater influence on breaking dormancy (Archana Mukherjee et al., 2009). This

may be due to increase in availability of sugars due to increase in respiration at higher

temperature.

Compared to smoke and heat treatments, soaking corms in different chemicals [KNO3,

thiourea, ammonium sulphate (NH4SO4)] for a short period (20 - 30 minutes) had no significant

effect on inducing early sprouting (Mohankumar and Ravi, 2001). However, treating the apical

portion of corm (after removing the apical bud) with thiourea and subsequently wetting the

apical portion for a period of 10 days induced early sprouting with more number of sprouts

(Archana Mukherjee et al., 2009).

Darkness had adverse effect on sprouting (Kumar et al., 1998). In A. konjac, abscisic

acid (ABA) and ferulic acid were extracted from the dormant corms and exogenous application

of ABA (10 mg l-1) and ferulic acid (400 mg l-1) inhibited sprouting and growth of the terminal

buds of non-dormant corms suggesting that ABA and ferulic acid are inhibitors of sprouting of
dormant corms (Sun et al., 1996). Corms are acrid before dormancy, but the acridity decreases

after dormancy (Santosa et al., 2003).

Flower

The plant blooms annually around the beginning of the raining season. The flower bud

emerged from the corm as a purple shoot, and later blooms as a purple inflorescence.

The pistillate (female) and staminate (male) flowers are on the same plant and are crowded in

cylindrical masses as an inflorescence. The top part is responsible for secreting mucus that gives

off putrid, pungent smell that is used to attract pollinating insects, the middle part of the

inflorescence contains staminate, and the base of the inflorescence contains pistillate. The

stigmas of the female flowers will be receptive on the first day of the bloom, when the pungent

smell will draw pollinating insects inside, and the inflorescence will close, trapping them for a

night to allow the pollen deposited on the insect to be transferred to the stigmas. Later in the

second day, the female flower will no longer be receptive of pollens, the male flowers will start
to bloom, and the inflorescence will open again. This allows the pollen to be deposited on the

emerging insects to be pollinated on different flowers, while preventing the pollens from the

same inflorescence to fertilize itself, preventing inbreeding.[citation needed]

After 24-36 hours after the first bloom of the inflorescence, the inflorescence's female

flowers will start developing into berries bright red fruiting bodies, and other parts of the

inflorescence will start wilting away. The berries are red when ripe and are not quite round,

being subglobose or ovoid.[7]

Shoot characteristics

The new growing part of the A. paeoniifoliu arises from the corm, where its sprouting

period depends on the dormancy status of the planting material. Upon completion of the plant’s

dormancy period, the new shoot sprout will emerge subsequently after it is planted. Leaf

emergence is delayed when the apical buds of seed corms are damaged. Leaves arose earlier

when whole corms were planted than when cut corms were planted irrespective of corm size

(Sen et al., 1996).

After the plantation of the whole corms, bud portions or upper half sections of the plant,

buds are expected to sprout two to three weeks prior planting. However, when vertical 1/2, 1/4

and 1/8 corm sections and lower half corm sections were planted, the buds are expected to start

sprouting sprout 4-7 weeks prior planting (Sugiyama & Santosa, 2008).

As soon as the sprout is initiated, continual development of the new shoot may be

completed within 30 days (Plate 1. A to F).

planting material that act as its vegetative reproductive structure. Then, the daughter corms, new

corms, enlarge by using assimilates synthesized by the leaves.

The leaves of the plant arise from the base of the stem, grow vertically (erect) and

remain solitary, having sizes ranging from medium to very large. It is compounded, where it is

made up of two or more parts. Also, it is pinnate, resembling a feather-like appearance where the

similar parts are arranged on opposite sides of an axis.

Commonly, the genus Amorphophallus have corms that have one bud situated in its

uppermost portion, the apex. The bud is found inside the cavity in the head part of seed corms.

Three or four small cataphylls, a reduced or scarcely developed leaf, found in the head part of

corms cover the apical bud in which the first leaf growth has already differentiated at planting.

The cataphylls elongate concomitantly with leaf development. Possibly, they protect a leaf from

damage by soil impedance during development. Furthermore, subepidermal cells of cataphylls

may contain needle-like crystals of calcium oxalate which presumably offer protection to a

young leaf from damage by pests. Cataphyll size depends on corm size and plant age; they

wither after the leaves become mature.

Leaves are composed of a petiole (pseudostem) and three rachises with many leaflets.

The number of leaves developing during the growing season is dependent on corm age. During a

growing season up to 12 leaves may be produced successively. As such, more than 2 leaves may

coexist at the same time. The number of leaves is also determined by the size of planting

materials. Plants originating from small corms (10 g) produce three to eight leaves, while large

corms (500 g) usually produce one or two leaves during a growing season. Under weedy
conditions leaves are submerged (Fig. 3) under weeds and the number of leaves, total leaf area,

leaf thickness and fresh masses of corms decreases markedly (Santosa et al., 2006c).

When pre-flowering and post flowering corms with similar fresh masses were planted

both types of corms sprouted at about the same time; however, leaf sizes (length of petioles and

rachis) were larger in preflowering corms than in postflowering corms (Sugiyama and Santosa,

2008).

Up to 150-250 leaflets per leaf may be produced per leaf and it may vary among

accessions. The leaf area of any 1 of the 3 lobes of A. campanulatus leaves showed a highly

significant correlation (r = 0.93 to 0.97) with total leaf area (Patel and Mehta, 1987).

The number of stomata in lower epidermis increased from 10.22 per unit area at 50 DAP

to 17.78 per unit area at 150 DAP (Gopi et al., 2008). The stoma has 2 adjacent cells surrounded

by 4 subsidiary cells (Plate 2. a and b). The leaf area index increased with time and reached a

maximum (6.1) at 120 DAP at a planting density of 1,40,000 plants ha-1 (Das et al., 1997).

On the other hand, the LAI reached 4.4 and 5.4 at a planting density of 1,00,000 and

1,20,000 plants ha-1 respectively. Petioles (pseudostem) looks like the stems of normal plants

and are cylindrical in morphology. In general, large petioles indicate that the corm is also large.

The mean shoot length varied between 47.3 - 122.5 cm depending uponthe variety, plant spacing

or size of planting material used (Mukhopadhyay and Sen, 1986; Ravindran andKabeerathumma,

1991; Sen and Das, 1991; Goswami and Sen, 1992; James George and Nair 1993; Geetha, 2001;

Suja et al., 2005, 2006). Increase in N application from 50 to 150 kg ha-1 increased shoot length

by 11%, (Mukhopadhyay and Sen, 1986) or did not increase shoot length and girth (Geetha,
2001). Increase in K application from 50 to 150 kg ha-1 did not have any significant effect on

shoot growth (Mukhopadhyay and Sen, 1986; Geetha 2001).

Regardless of plant spacing, increase in size of planting material increased plant

(pseudostem) height and plant height was maximum (84.6 cm) when 1 kg cut corm piece was

used as planting material. Closer plant spacing (60 x 45 cm) increased plant height (53.8 cm)

than wider plant spacing (90 x 90 cm) (James George and Nair, 1993). Plants produced from

whole seed corms were taller than those produced from cut pieces of corm of the same size. This

may be due to early sprouting and better root ramification (Sen and Das, 1991).

The canopy spread varied between 72.2 and 143.8 cm (Ravindran and Kabeerathumma,

1991; Sen and Das, 1991; Goswami and Sen, 1992; James George and Nair, 1993). Regardless

of plant spacing, increase in size of planting material increased canopy spread and canopy spread

was maximum (132.7 cm) when 1 kg cut corm piece was used as planting material at wider plant

spacing (90 x 90 cm) (Sen and Das, 1991). The canopy spread was more in plants raised by

planting whole seed corms than that in plants produced from cut pieces of corms of the same

size. This was presumably due to early sprouting and better root ramification (Sen and Das,

1991).

Biomass production of shoot (leaf and pseudostem/ petiole) increased up to 120 days

after planting (DAP) and 150 DAP respectively and declined thereafter whereas the corm dry

weight and total dry matter production (TDMP) showed a steady increase up to the maturity. The

corm dry – matter production (CDMP) per ha increased with increase in planting material size or

plant density and highest CDMP (25.6 t ha-1 and 19.4t ha-1 respectively) was observed at 6
MAP by using 250 g cut corm pieces as planting material or with high plant density (14 plants

m-2) (Das et al., 1997).

The crop growth rate (CGR) increased gradually up to 120-150 DAP and sharply

declined at maturity as the crop growth ceased. However, the relative growth rate (RGR)

continued to decrease with crop age and was the highest at early growth stage (Das et al., 1997).

The leaf area increased with increase in planting material size or plant density and highest leaf-

area index (5.4) was observed between 4 and 5 months after planting by using 250 g cut corm

pieces as planting material or with high plant density (14 plants m-2) (Das et al., 1997).

Similarly, CGR increased with increase in planting material size or plant density and

highest CGR (25.3-32.2 g m-2 day-1) was observed at 5 months by using 250 g cut corm piece

as planting material. The CGR was 22. 4 g m-2 day-1 at a plant density of 14 plants m-2 (Das et

al., 1997). Treating corm pieces from bottom portion of corm with growth regulators viz.,

thiourea, KNO3 and GA3 effectively influenced the growth characters and GA3 gave maximum

corm yield (Das et al., 1997).

Application of triazole compounds (systemic fungicides) triadimefon (TDM),

paclobutrazole (PBZ) and propiconazole (PCZ) through soil drenching increased total root length

(by 8.85 - 75.92%), dry weight of whole plant (by 71.44 – 84.91%), intercellular CO2

concentration (by 25.12 – 27.91%), leaf thickness, number of spongy and palisade cells, number

of chloroplasts per cell, net photosynthetic rate (PN) (by 15.7 – 28.92%) and water use efficiency

(WUE) (by 56.81 – 87.9%) as compared to untreated control plants. In contrast, total leaf area,

transpiration rate (TR) and stomatal conductance decreased (Gopi et al., 2005, 2008, 2009).
Root characteristics

Roots grow out from the surface of newly developing daughter corms at the base of the

pseudostem through the remnants of the cataphylls concomitantly with leaf emergence. These

roots extend horizontally and are densely distributed at a shallow depth of top 15-30 cm soil

depth. The roots are cylindrical and 2 to 5 mm thick.

Roots grow more than 1 m in length under adequate soil moisture conditions or under

adequate rain and are known as “rain roots”. Under dry soil conditions, the root length decreases

to less than 30 cm length. The transverse section (T.S.) of root shows about 25 layersof thin

walled parenchymatous cortex cells surrounding a central stelar portion with 8 protoxylem points

(Plate 3 a, b and c).

Effect of water deficit stress

Little research work has been done on the response of Amorphophallus to water deficit

stress. Soil moisture status does not influence sprouting but further development of new shoot

depends on adequate soil moisture.

1000-1500 mm of rainfall per year is optimum for the crop (Jansen et al., 1996).

Many plants enter dormancy earlier than usual when the rainy season is shorter than 4

months and supplementary irrigation is necessary for high productivity when the rainy season is

shorter than 4 months. Plants produced a larger number of leaves under frequent watering (1-, 3-

and 5 day intervals) than under 7- and 15-day intervals; the third leaves were produced in

treatments up to 7-day intervals, but neither the second nor the third leaves were produced in 15-

day intervals. Furthermore, frequent watering produced large leaves and extended their life span

compared with less frequent watering (Santosa et al., 2004b).

A decrease in the dry mass of seed corms was more evident with frequent watering,

suggesting that reserved carbohydrates in seed corms are not easily metabolized under a limited

water supply. The ratios of dry mass of daughter corms to that of seed corms are 6.1, 1.1, 0.6, 0.4

and 0.2 at 1-, 3-, 5-, 7-, and 15-day intervals, respectively. The high ratio of dry mass of daughter

corms to that of seed corms under frequent watering treatments could be ascribed to the fact that

the soil water availability affects not only the utilization of dry matter in seed corms but also the

production and translocation of photoassimilates into daughter corms (Sugiyama and Santosa,

2008). The roots dried earlier than usual when the soil water content decreased to less than 40%

of field capacity (Santosa et al., 2004b). The crop tolerates water deficit stress conditions for

about 30-60 days but prolonged stress may affect corm yield (Santosa et al., 2004b).

In green-house conditions, plant growth was not affected when plants were watered at 1,

3 or 5 day intervals. Nevertheless, infrequent watering (watering at 7 or 15 days intervals)

reduced corm yield and forced the corms to enter into dormancy. Soil moisture conservation
methods like mulching induced higher percentage of early sprouting, greater canopy spread,

plant height, greater mean corm weight and corm yield (Mohankumar et al., 1973). The corm

yield of elephant foot yam was greater under surface irrigation (40.0 t ha-1) and microirrigation

@ 80% CPE (37.0 t ha-1) than under rainfed conditions (25.2 t ha-1) (M. Nedunchezhiyan,

personal communication).

Seed dormancy

Successful seed production has been reported in Amorphophallus (Arakeri, 1950). Seed

dormancy of 5- 6 months has been reported in this crop (Arakeri, 1956). Exposing seeds to

running water for 6 days resulted in highest sprouting (55.5%) as compared to control (2.7%).

However, exposing seeds to water for more than 6 days resulted in lower percentage of sprouting

(Rajendran and Hrishi, 1976).

Response to shade

Elephant foot yam tolerates shade conditions. Therefore, it can be intercropped between

young trees. Corm yield decreased by 66% when the light intensity is reduced to 25% of full

sunlight (Pushpakumari and Sasidhar, 1992).

In contrary, Santosa et al., (2006) reported that the fresh biomass of corms increased with

a decrease in light intensity; 75% shading produced the largest corms and 0% shading produced

the smallest corms.

Under full sunlight, necrosis and curling at either the edge or the tips of leaflets occurred

causing 25% loss of the crop. No damage was observed in the 25, 50 and 75% shading.
 However, the shading treatments significantly decreased the leaf number. The short life

span of leaves might enhance the production of new leaves resulting in a larger number of leaves

under full sunlight. Shading treatments significantly affect the length of petioles and rachis.

Plants developed the shortest petioles under full sunlight but the longest under 75% shading.
REFERENCES

Das, S. S., Sen, M., Dey, Y. N., De, S., & Ghosh, A. K. (2009). Effects of petroleum ether

extract of Amorphophallus paeoniifolius tuber on central nervous system in mice. Indian journal

of pharmaceutical sciences, 71(6), 651

De, S., Dey, Y. N., & Ghosh, A. K. (2010). Phytochemical investigation and chromatographic

evaluation of the different extracts of tuber of Amorphaphallus paeoniifolius (Araceae). Int J

Pharm Biol Res, 1, 150-157.

Dey, Y. N., De, S. H. A. N. K. H. A. J. I. T., & Ghosh, A. K. (2010). Evaluation of analgesic

activity of methanolic extract of Amorphophallus paeoniifolius tuber by tail flick and acetic acid-

induced writhing response method. Int J Pharm Biosci, 1, 662-8.

Harish, A., Rashmi, M., Krishna Murthy, T. P., Blessy, B. M., & Ananda, S. (2014).

Mathematical modeling of thin layer microwave drying kinetics of elephant foot yam

(Amorphophallus paeoniifolius). International Food Research Journal, 21(3).

Lim, T. K. (2015). Amorphophallus paeoniifolius. In Edible Medicinal and Non Medicinal

Plants (pp. 443-453). Springer Netherlands.

paeoniifolius. International journal of advanced scientific research and technology, 2(2), 99-

111.

Punekar, S. A., & Kumaran, K. P. N. (2010). Pollen morphology and pollination ecology of

Amorphophallus species from North Western Ghats and Konkan region of India. Flora-

Morphology, Distribution, Functional Ecology of Plants, 205(5), 326-336.

Ravi, V., Ravindran, C. S., & Suja, G. (2009). Growth and productivity of elephant foot yam

(Amorphophallus paeoniifolius (Dennst.) Nicolson): an overview. J. Root Crops, 35(2), 131-

142.

Santosa, E., Lian, C. L., Sugiyama, N., Misra, R. S., Boonkorkaew, P., & Thanomchit, K.

(2017). Population structure of elephant foot yams (Amorphophallus paeoniifolius (Dennst.)

Nicolson) in Asia. PloS one, 12(6), e0180000.

Singh, A., Srivastava, K. C., Banerjee, A., & Wadhwa, N. (2013). Phytochemical analysis of

peel of Amorphophallus paeoniifolius. Int J Pharma Bio Sci, 4, 810-815.

Singh, Anuradha, and Neeraj Wadhwa. "A review on multiple potential of aroid:

Amorphophallus paeoniifolius." Int J Pharm Sci Rev Res 24.1 (2014): 55-60.
Sungkajanttranon, O. ก F ก F ก ก F กก Morphology and seedling growth of Amorphophallus

paeoniifolius (Dennst.) Nicols. propagated by seed.