Amophophallus Paeoniifolius: A Synthesis Paper
Amophophallus Paeoniifolius: A Synthesis Paper
Amophophallus Paeoniifolius: A Synthesis Paper
A Synthesis Paper
In Partial Fulfilment of
the Requirements in
Biology III
INTRODUCTION
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
(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
southwest India by S. R. Anil, E. A. Siril, and S.S. Beevy, the growth, morphology, and the
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.
Conservation status
foot yam conservation status is least concern. The crop has high productivity and wide
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
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
(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
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
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
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
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
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,
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
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
planting material that act as its vegetative reproductive structure. Then, the daughter corms, new
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
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
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
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
(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
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
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
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
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
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
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,
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
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
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
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.
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Sungkajanttranon, O. ก F ก F ก ก F กก Morphology and seedling growth of Amorphophallus