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The International Journal Of Engineering And Science (IJES)
||Volume||2 ||Issue|| 7 ||Pages|| 01-05||2013||
ISSN(e): 2319 – 1813 ISSN(p): 2319 – 1805
www.theijes.com The IJES Page 1
Physiological, Biochemical and Cellular Changes Associated With
the Ripening Of Bitter Less Bitter Gourd
(Momordica Dioica Roxb. Ex Willd.) Fruits
1,
Payal T. Shah and 2,
T. V. Ramana Rao
1,
Department Of Biosciences Sardar Patel University VALLABH VIDYANAGAR- 388 120 Gujarat, INDIA
2,
Asst. Professor Shree P. M. Patel Collage, ANAND, Gujarat: INDIA
-------------------------------------------------------ABSTRACT ---------------------------------------------------
Fruits of Momordica dioica were selected at seven sequential developmental stages, starting from very young
stage to post ripened stage and were analyzed. From the study physiological and biochemical changes it was
concluded that gradual decrease was found in chlorophyll-a (5.25 fold), chlorophyll-b (13.0 fold), total
chlorophyll (8.23 fold), starch (6.5 fold) and free amino acids (14.4 fold), while in case of total proteins
decrease was observed from pre-mature stage to post ripened stage 67.2 percent. In case of RNA, decrease was
found 55.1 percent from mature stage to post ripened stage. However, gradual increase level was found only in
carotenoids (2.5 fold) and in total sugars (209 percent), non-reducing sugars (317 percent) and phenol (2.9
fold) increase was found from very young stage to pre-ripened stage. In contrast the amount of anthocyanins,
reducing sugars and DNA was unstable.Hydrolytic enzymes viz., Amylase, Invertase and Peroxidase; the
activity of Amylase decreased from mature stage to post ripened stage (86.73 percent). While activities of
Invertase and Peroxidase was noticed in 94.3 percent decreased and 44.64 percent increased from very young
stage to mature stage respectively.In contrast, the activities of cell wall degrading enzymes (such as cellulase,
polygalactouronase (PG) and pectin methyl esterase (PME)) 6.38 fold increase activity was noticed in cellulase
activity from very young stage to ripened stage while during ripening PG activity gradually increased with 2.37
fold, whereas the activity of PME was noticed inconsistent. These biochemical changes also reflect on cellular
changes such as increasing cell number, enlargement of cell, decreasing cell content, separation of middle
lamella etc., were observed during the mature stage to post ripened stage. The low production of ethylene and
an increased rate of respiration indicates that the bitter less bitter gourd fruit falls under the category of
climacteric fruits.
----------------------------------------------------------------------------------------------------------------------------------------
Date of Submission: 10 June 2013, Date of Publication: 10.July 2013
----------------------------------------------------------------------------------------------------------------------------------------
I. INTRODUCTION
Momordica dioica (Bitter less bitter gourd, kakrol, Teasle gourd) is a perennial, dioecious climber
with thickened roots, which belongs to cucurbitaceae family occurring in Southern Asia. Bitter less bitter gourd
is relatively small and oval in shape. This fruit is in demand for internal as well as external markets due to its
medicinal properties and is also rich in calcium, phosphorus, iron and carotenoids. Its immature fruits are
berries; softly echinate which are used as a vegetable (More and Nayar, (1998). Luo, et al., (1998) found three
triterpenes and two steroidal compounds were isolated from the dry root of Momordica dioica. Their structures
were elucidated by spectral analyses (MS, IR, 1HNMR, 13CNMR and DEPT) and chemical methods.Ali and
Srivastava (1998) found two new chemical constituents for the first time from the fruit of Momordica dioica
along with the known sterol compound and an unknown pentacylic triterpene isolated from the seeds. A more
or similar study on phyotochemical constituent was previously studied by Shantha, and Radhakrishnaiah (1993)
in the Momordica sps. From the above report it was derived that this plant is not only helpful for the medicinal
purpose but is also commercially important. There are very few reports cited on physiological studies such as
increase in respiration, ethylene production, carotenoids synthesis, chlorophyll degradation and hormonal
changes was previously reported (Kays and Hayes (1978); Zheng, (1986) Tan et al., (1999), Rodriguez et al.,
(1976), Huong & Raymundo (1999), Basu et al., (1994), in M. charantia but not in M. dioica as a result this
fruit was selected for present study.For giving idea about fruit ripening this study was alienated in five divisions
(i) Changes in pigments (ii) Changes in primary metabolites (iii) Changes in hydrolytic and cell wall degrading
enzymes (iv) Changes in cellular structure & (iv) Changes in respiration rate and ethylene production.
Physiological, Biochemical And Cellular Changes…
www.theijes.com The IJES Page 2
II. MATERIALS AND METHODS
The fruits of bitter gourd bitter less were collected from local fields at seven sequential stages, starting
from very young stage to post-ripening stage. The length and breadth were measured for the collected fruit
samples before they were subjected for their physiological and biochemical analyses and their remarkable data
were tabulated (Table 1, Pl. 1.1). Biochemical analysis of pigments (Chlorophyll, Carotenoids & Anthocyanins)
and primary metabolites (Sugars, Starch, Total Proteins, Total Phenols & Free Amino Acids) were measured
from the collected fruit samples by using different methods cited by Thimmaiah (1999). The estimation of
nucleic acids (DNA & RNA) was performed by the methods of Devi (2002). Amylase, Invertase, Peroxidase,
Polygalactrouronase and Pectin Methyl Esterase were extracted and assayed according to the procedure
described by Selveraj and Pal (1984) & Thimmaiah (1999). Data presented at each stage were the mean of three
replicates (± Standard error, S.E.). Data of the experiment were subjected to statistical analysis using Duncan’s
multiple range test (DMRT) (Bliss, 1967).The histological and ultra structure studies were carried out from the
last four stages (including mature stage to post ripening stage) of bitter gourd fruits. The collected fruits were
fixed in Formalin Acetic Acid (FAA) (Berlyn & Miksche, 1976). Customary methods described by Johansen (6)
were followed for the dehydration and embedding of these FAA fixed materials. The sections of 8 - 10 µm thick
were cut with the AO Spencer's rotary microtome from the paraffin embedded materials and then they were
stained with safranin - fast green combination. The stained sections of fruit were observed under Carl-Zeiss
MP3 400 Microscope and photomicrographs were taken with same microscope with an Image Analyzer Facility.
For the TEM studies, the preparation of fruit materials has been done as per the methods cited by Dawes (1971)
and the ultra thin sections were observed under the Philips Tecnai (High Tension 200 KV) TEM. The amount of
ethylene synthesis and the rate of respiration were measured from mature, pre- ripening, ripening and post
ripening stages with the Perkinelmer Autosystem XL Gas Chromatography as per the procedures cited by
Thimmaiah (1999).
III. RESULTS AND DISCUSSION
3.1.Changes in Pigments:-
The quantitative analysis of the Chlorophylls revels that the amount of Chlorophyll a (0.042 mg/gm),
Chlorophyll b (0.065mg/gm) and Total chlorophyll (0.107 mg/gm) were very high in very young stage, but as
the fruit proceeds towards the maturation, ripening and post ripening stages the amount of chl. a (0.008 mg/gm),
chl. b (0.005 mg/gm) and total chl. (0.013 mg/gm) were found declined 5.25 fold, 13.0 fold and 8.23 folds
respectively. Carotenoids (0.067 mg/gm) were found gradually increasing from very young stage to almost 2.5
folds at post ripening stage (0. 170mg /gm). Anthocyanins dose not show consistency in their quantity during
the maturation, ripening and post ripening stages of fruit. The maximum amount of anthocyanins (6.365 mg/gm)
found in the young stage of fruit.Merzlyak et al., 2003 found that during the analysis of pigments, amount of
chlorophyll were decline, carotenoids increases due to the pigments changes at terminal stages of leaf and fruit
development in many plant species. At these stages plant tissues retain certain amounts of carotenoids or
carotenoids synthesis was stimulated on the background of chlorophyll degradation. The increasing level of
caroetnoids and decreasing level of chlorophyll also recently noticed by Wang and Zhong (2005) during
ripening of tomato fruit. The decreasing level of chlorophyll a, chlorophyll b and Total chlorophyll and
increased level of carotenoids were also previously reported in the cucurbits during ripening and storage period
by Tan et al., (1999), Rodriguez et al., (1976) and Huong & Raymundo(1999). Anthocyanins were found
unstable during ripening which was observed by Underhill and Critchley (1993) in Litchi fruit due to
polyphenol oxidase degradation.
3.2.Changes in primary metabolites:-
The quantitative analyses of sugars (i.e. total sugars, reducing sugars and non reducing sugars) indicate
that the amount of total sugars was gradually increased from very young stage (0.986 mg/gm) to pre-ripened
stage (3.047 mg/gm) but there after their amount declines which displayed 209 percent increase. However, the
quantum of reducing sugars does not show consistency. In contrast non-reducing sugars was gradually increased
from very young stage (0.686 mg/gm) to pre-ripened stage (2.864 mg/gm) which displayed 317 percent
increase, but thereafter declining nature was seen in remaining stages.A more or similar observation was also
noticed in the apple fruit during there ripening process (Sabir et al., 2004). From the data it was concluded that
increase in sugar might be due to hydrolysis of polysaccharides or due to less acidity and ascorbic acid contents.
The contents of reducing sugar and non-reducing in spongy tissue may be also due to lower activities of
invertase and amylase (Luiz et al., 2001).The amount of starch was observed 1.380 mg/gm at very young stage
but thereafter a steady decline was found in the post ripened stage 0.211 mg/gm which displayed 6.5 fold
decrease. According to Luiz et al., 2001 during development stage of mango, starch was hydrolyzed. However,
as the fruit becomes over-ripe, only traces of starch were detected and amylase activity was substantially
reduced.
Physiological, Biochemical And Cellular Changes…
www.theijes.com The IJES Page 3
The sugars in the pulp spongy tissue were accounted by the presence of higher content of starch, which
remained unhydrolysed due the low activity of amylase.The quantitative analyses of the total proteins, free
amino acids and total phenols in M. dioica reveal that initially the amount of total protein does not show
consistency during their growth development and ripening process. The amount of total protein was decrease
from pre-mature stage (7.644 mg/gm) to post ripened stage (2.500 mg/gm), which showed 67.2 percent
decrease. The amount of total phenol was gradually increasing from very young stage (2.074 mg/gm) to pre-
ripened stage (6.123 mg/gm) which showed 2.95 fold increase thereafter it declines in later stages. In, contrast
of proteins and phenols the amount of free amino acids shows 14.4 fold gradual decrease.Hulme et al., (1968)
and Dilley (1972) reported that in the early stages of fruit protein synthesis is stimulated and reaches a peak
value of climacteric thereafter it declines slowly and gradually during ripening process. An increase in protein
synthesis during early climacteric period has been measured in many fruits observed by Sharma (2000). The
sharp decrease in free amino acids towards maturity and onset of ripening has been attributed to their
incorporation into protein required for the synthesis of various ripening enzymes (Frunkel, Klein and Dilley,
1965). This declining trend of amino acids has been noticed by a number of workers during the approach of fruit
maturity (Sharma, (2000)) According to (Dilley, 1970 and Sharma, 2000) the phenolics content of most fruits
declines from high levels during early growth to low levels when the fruit is considered to be physiologically
mature and therefore, susceptible to the induction of ripening. A more or similar observation was found by
Kumar and Goswami (1985) that the presence of higher concentration of phenolics during early stage of
development provides protection mechanism to the phytohormones like auxins, gibberellin and cytokines, which
play an important role in cell division and cell enlargement. The quantitative analysis of nucleic acids (i.e. DNA
& RNA) has been carried out in the seven sequential developmental stages of fruit. The amount of DNA
remains inconsistent during ripening while, RNA does not shows consistency in earlier stages. RNA was found
decrease from mature stage (0.261 mg/gm) to post ripened stage (0.117 mg/gm), which displayed 55.1 percent
decrease.
During the climacteric period the RNA was increased and there after its decline up to ripening stage,
while in the case of DNA it is fluctuated during the ripening noticed by (Sharma 2000).Amylase activity was
found to be gradually declining from very mature stage (0.377 activity) to the post ripened stage (0.050
activity), which displayed 86.73 percent decrease. In contrast, Invertase shows increase in first stage (0.037
activity) and declines upto mature stage (0.019 activity), which displayed 94.3 percent decrease and remains
unstable for remaining stages. In case of Peroxidase was gradually increased from very young stage (0.274
activity) to mature stage (0.495 activity) that displayed 44.64 percent increase. These hydrolytic enzymes were
observed very high in the early stage of development and thereafter they declined towards the ripening due to
starch disappearance, hydrolysis of sugar, climacteric periods or related to growth development process
respectively. The activities of hydrolytic enzymes increased from early mature stage to peel colour turning stage
and declined in later ripening stage of pineapple fruit during ripening was observed by Selveraj (1993). A more
or similar observation also noticed by previously Hulme (1970) and Seymour et al., (1993).
The activities of cell wall degrading enzymes viz., Cellulase, Polygalactrouronase (PG) and Pectin
Methyl Esterase (PME) during the ripening of fruit were presented in (Table 1.5A, Fig. 1.5B). From analyzed of
cell wall degrading enzymes indicate that there was an increment noticed in the activities of cellulase from very
young stage (0.013 activity) to ripened stage (0.083 activity) which displayed 6.38 fold increase. PG enzymes
were gradually increase 2.37 fold during ripening. While the activity of PME was gradually decline from mature
stage (1.009 activity) to post ripened stage (0.800 activity) which displayed 20.7 percent decrease.It has been
widely recognized that cell wall changes are related to fruit softening and that these modification are due to the
action of cell wall enzymes. The accumulation of these enzymes during ripening and their often-concerted
activity will result, eventually, in degradation to many cell wall polymers leading to cell separation and
associated softening. During ripening increasing activities of cell wall softening enzymes were reported by
Selveraj & Raju (2000) in Kagzi Lime fruit.
3.3.Changes in cellular structure:-
Histological Studies:-
The development anatomy of fruit has provided information about the prepositional cell division and
cell enlargement in various parts of fruit. For better understanding of histological changes the fruits were
selected from the mature stage to post ripened stage. It was found that mesocarps cells were divided into three
categories (i) Outer mesocarps (ii) Middle mesocarps (iii) Inner mesocarps cell. Middle mesocarps cells and
inner mesocarps cells are parenchyma cell, which contain abundant starch grains, as the middle mesocarps cells
were comparatively thick; hence they were distinguished from the inner ones.
Physiological, Biochemical And Cellular Changes…
www.theijes.com The IJES Page 4
At mature stage in the formation of inner mesocarps cells parenchyma cells were very small in size and
contained abundant starch due to cell division and cell enlargement. In the starting process of pre-ripened and
ripened stage the cell number and cell size were increased while, starch grain was decrease. However, in the last
stage the cells were collapsed and empty or contained very less amount of starch grains. During the different
developmental stages of Momordica dioica fruit more or similar histological changes was reported by
Thanki(1978) (Plate 2.2).White (2002) concluded that “Anatomically, fruit are swollen ovaries that may also
contain associated flower parts. Their development follows fertilization, and occurs simultaneously with seed
maturation. Initially, fruits enlarge through cell division and then by increasing cell volume”.
3.4.ULTRA STRUCTURE STUDIES:-
This study helps in understanding the cell wall softening or changes in cell wall. For this reason the
fruit were selected from mature stage to post-ripened stage. In the earlier development stages the cell wall
becomes very hard but when the ripening process starts the cell wall becomes soft due to presence of cell wall
degrading enzymes polygalactouronase(PG), pectin methyl esterase(PME) and cellulase. The investigations of
plasma lamella was observed at mature stage, which was cited near cell wall, while, in pre-ripened stage the cell
wall enlarged in their position, which contains granular cytoplasm, ribosome like bodies and nucleus. However,
during the ripening and post ripening stages cell wall becomes despoiled. These structural patterns were also
previously noticed by Thanki (1978) and Whitaker and Davis(1962) in cucurbitaceae fruit.
Brummell et al., (2004) concluded that cell wall softening occur due to differences in cell wall
thickness and composition, cell size, cell shape, packing, contents and turgor.
Changes in respiration rate and ethylene production:-
3.5.ETHYLENE AND RESPIRATION:-
The ethylene production was increase from pre-ripened stage to the post-ripened stage of the fruit,
while the rate of the respiration was increased from mature stage to ripened stage, their after decline nature
observed in the post ripened stage (Fig 2.6).According to Hardenuburg et al., (1986), Zong et al., (1995), Kays
& Hays (1978) and Zheng (1986) conclude that the fruits of bitter gourd (M. charantia) was climacteric fruit
due increasing rate of respiration and low production of ethylene. A more or similar observation was also
noticed in M. dioica fruit.
3.6.ANALYSIS OF VARIANCE:-
The analysis of variance revels that the changes of fruit ripening such as Chl. a, Chl. b, Total
chlorophyll, carotenoids, amino acids and the activities of amylase and peroxidase were highly significant
(Tables 2.1, 2.3 & 2.4), while less significance was observed in total sugars, starch and cellulase activity (Tables
2.2 & 2.5). However, no significance was found in case of anthocyanins, reducing sugars, non- reducing sugars,
total proteins, total phenols, nucleic acids, invertase, PG and PME activities (Tables 2.1, 2.2, 2.3 and 2.5)
REFERENCES:
[1] Ali, M. and V. Srivastava (1998). Characterization of phytoconstituents of the fruits of Momordica dioica. Indian J. of Pharm.
Sci. 60(5): 287-289
[2] Basu, P. S., S. Banerjee, and S. Das 1994. Hormonal regulation of flowering and fruit development: effects of dikegulac on
flowering, fruit setting and development of Momordica charantia L. and Luffa acutangula Roxb. Indian J. of Plant Physiol.
37(4): 282-285.
[3] Berlyn, G. P. and J. P. Miksche 1976. Botanical microtechnique and cytochemistry. IOMA State University press, IOWA.
[4] Bliss, C. I. 1967. “Statistics in biology”, (Vol-1) Statistical Methods for Research in the Natural Sciences. McGraw-Hill Book
Company.
[5] Brummell, A. D., C. D. Valeriano, C. H. Crisosto and J. M. Labavitch 2004. Cell wall metabolism during maturation, ripening
and senescence of peach fruit. J. Exp. of Bot. 55: 2029-2039.
[6] Dawes, C. J. 1971. Biological techniques in electron microscopy. Barns and Nobel Inc., New York.
[7] Dilley, D. R. 1970. Enzymes. In: A.C. Hulme (ed.), Biochemistry of fruits and their products. Academic press, London.
[8] Frankel, C., I. Klein and D. R. Dilley 1968, Protein synthesis enzymes in pome fruits. Plant Physiol., 43: 1146.
[9] Hardenburg, R. E., A.E. Watada and C.Y. Wang 1986. The commercial storage of fruits vegetables and florist and nursery
stocks. Agric. Handb., USDA.
[10] Hulme, A. C. 1970. The biochemistry of Fruits and their products. Academic Press, London.
[11] Hulme, A. C., M. C. J. Rhodes, T. Galliard and L. S. C. Wooltrotan 1968. Metabolic changes in excise fruit tissue IV. Changes
occurring in the disc of apple peel during the development of respiration climacteric. Plant Physiol., 43: 1154
[12] Huong, T. T. L. and L. C. Raymundo 1999. Biosynthesis of carotenoids in bitter melon at high temperature. Phytochem., 52(2):
275-280.
[13] Johansen, D. A. 1940. Plant microtechnique. McGraw-Hill, New York.
[14] Kays, S. J. and M. J. Hayes 1978. Induction of ripening in the fruit of Momordica charantia L. by ethylene Trop. Agric., 55: 167-
172.
[15] Kumar S. and A. K. Goswami 1985. Phenol content and peroxidase system in developing apple fruits. Indian J. Hort., 42: 184.
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[17] Luo,L., Z. Li., Y. Zhang and R. Huang 1998. Triterpenes and steroidal compounds from Momordica dioica. Yaoxue Xuebao
33(11): 839-842.
[18] Merzlyak et al., 2003
[19] More T. A. and N. M. Nayar 1998. Cucurbits. Oxford & IBH publishing Co. Pvt. Ltd., New Delhi
[20] Pratibha Devi 2002. Principles and methods of plant molecular biology, biochemistry and genetics, Agrobios publishers, India.
[21] Rodriguez, D. B., L. C. Raymundo, T.C. Lee, K. L. Simpson, and C, O. Chichester 1976. Carotenoids pigment changes in
ripening Momordica charantia fruit. Annu. Bot., 40: 615-624.
[22] Selveraj Y. 1993. Changes in enzymes activity in ripening pineapple fruit Indian. J. Hort., 50(4): 310-317.
[23] Selveraj Y. and K. Pal 1984. Changes in the chemical composition and enzymes activity of two sapodilla (Manikara zapota)
cultivars during development and ripening. J. of. Hort. Sci., 59 (2): 275-281.
[24] Selveraj Y. and M. Edward Raju 2000. Biochemical of ripening of Kagzi Lime (Citrus aurantifolia Swingle) fruit. Indian. J.
Hort., 57(1):1-8.
[25] Seymour, G. B. and W. B. McGlasson 1993. Melons. In: Seymour, G. B., J. E. Taylor and G. A. Tucker (ed.), Biochemistry of
fruit ripening. Chapman & Hall, London.
[26] Shantha, A. and M. Rada Krishanaiah 1993. Photochemical constituent of Momordica charantia L. (Cucurbitceae).Feddes
repertorium 104 (1-2): 51-52.
[27] Singh R. R. and R. M. Sharma 2000. Structure, cellular and composition of fruits and vegetable. In: L. R. Verma & V. K. Joshi
(ed.), Post harvest technology of fruit and vegetables. Indus Publishing, New Delhi.
[28] Tan, X. P., Y. Ueda, Y. Imahori, and K. Chachin, 1999. Changes in pigments in pulp and aril of Balsam pear (Momordica
charantia L.) fruit during development and storage. J. Japanese Soc. for Hortic. Sci. 68(3): 683 - 688.
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India
[30] Thimmaiah, S. K. 1999. Standard methods of biochemical analysis. Kalayani Publishers, New Delhi.
[31] Underhill, S. J. R. and C. Critchley 1993. Physiological, biochemical and anatomical changes in Lychee (Litchi chinensis Sonn.)
pericarp during storage. J. of. Hort. Sci. 68(3): 327-325.
[32] Wang and Zhong (2005
[33] Whitaker, T. W. and G. N. Davis 1962. Cucurbits. Inter Sciences, New York.
[34] White 2002. Recent advances in fruit development and ripening: an overview. J. of Exp. Bot. 53: 1995-2000.
[35] Zheng, L.1986. Primary approach on the post harvest physiology of balsam pear. M. S. thesis, Beijing Vegetables Research
center, Beijing.
[36] Zong, R. J., L. Morris and M. Cantwell 1995. Post harvest physiology and quality of bitter melon (Momordica charantia L.).
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The International Journal of Engineering and Science (The IJES)

  • 1. The International Journal Of Engineering And Science (IJES) ||Volume||2 ||Issue|| 7 ||Pages|| 01-05||2013|| ISSN(e): 2319 – 1813 ISSN(p): 2319 – 1805 www.theijes.com The IJES Page 1 Physiological, Biochemical and Cellular Changes Associated With the Ripening Of Bitter Less Bitter Gourd (Momordica Dioica Roxb. Ex Willd.) Fruits 1, Payal T. Shah and 2, T. V. Ramana Rao 1, Department Of Biosciences Sardar Patel University VALLABH VIDYANAGAR- 388 120 Gujarat, INDIA 2, Asst. Professor Shree P. M. Patel Collage, ANAND, Gujarat: INDIA -------------------------------------------------------ABSTRACT --------------------------------------------------- Fruits of Momordica dioica were selected at seven sequential developmental stages, starting from very young stage to post ripened stage and were analyzed. From the study physiological and biochemical changes it was concluded that gradual decrease was found in chlorophyll-a (5.25 fold), chlorophyll-b (13.0 fold), total chlorophyll (8.23 fold), starch (6.5 fold) and free amino acids (14.4 fold), while in case of total proteins decrease was observed from pre-mature stage to post ripened stage 67.2 percent. In case of RNA, decrease was found 55.1 percent from mature stage to post ripened stage. However, gradual increase level was found only in carotenoids (2.5 fold) and in total sugars (209 percent), non-reducing sugars (317 percent) and phenol (2.9 fold) increase was found from very young stage to pre-ripened stage. In contrast the amount of anthocyanins, reducing sugars and DNA was unstable.Hydrolytic enzymes viz., Amylase, Invertase and Peroxidase; the activity of Amylase decreased from mature stage to post ripened stage (86.73 percent). While activities of Invertase and Peroxidase was noticed in 94.3 percent decreased and 44.64 percent increased from very young stage to mature stage respectively.In contrast, the activities of cell wall degrading enzymes (such as cellulase, polygalactouronase (PG) and pectin methyl esterase (PME)) 6.38 fold increase activity was noticed in cellulase activity from very young stage to ripened stage while during ripening PG activity gradually increased with 2.37 fold, whereas the activity of PME was noticed inconsistent. These biochemical changes also reflect on cellular changes such as increasing cell number, enlargement of cell, decreasing cell content, separation of middle lamella etc., were observed during the mature stage to post ripened stage. The low production of ethylene and an increased rate of respiration indicates that the bitter less bitter gourd fruit falls under the category of climacteric fruits. ---------------------------------------------------------------------------------------------------------------------------------------- Date of Submission: 10 June 2013, Date of Publication: 10.July 2013 ---------------------------------------------------------------------------------------------------------------------------------------- I. INTRODUCTION Momordica dioica (Bitter less bitter gourd, kakrol, Teasle gourd) is a perennial, dioecious climber with thickened roots, which belongs to cucurbitaceae family occurring in Southern Asia. Bitter less bitter gourd is relatively small and oval in shape. This fruit is in demand for internal as well as external markets due to its medicinal properties and is also rich in calcium, phosphorus, iron and carotenoids. Its immature fruits are berries; softly echinate which are used as a vegetable (More and Nayar, (1998). Luo, et al., (1998) found three triterpenes and two steroidal compounds were isolated from the dry root of Momordica dioica. Their structures were elucidated by spectral analyses (MS, IR, 1HNMR, 13CNMR and DEPT) and chemical methods.Ali and Srivastava (1998) found two new chemical constituents for the first time from the fruit of Momordica dioica along with the known sterol compound and an unknown pentacylic triterpene isolated from the seeds. A more or similar study on phyotochemical constituent was previously studied by Shantha, and Radhakrishnaiah (1993) in the Momordica sps. From the above report it was derived that this plant is not only helpful for the medicinal purpose but is also commercially important. There are very few reports cited on physiological studies such as increase in respiration, ethylene production, carotenoids synthesis, chlorophyll degradation and hormonal changes was previously reported (Kays and Hayes (1978); Zheng, (1986) Tan et al., (1999), Rodriguez et al., (1976), Huong & Raymundo (1999), Basu et al., (1994), in M. charantia but not in M. dioica as a result this fruit was selected for present study.For giving idea about fruit ripening this study was alienated in five divisions (i) Changes in pigments (ii) Changes in primary metabolites (iii) Changes in hydrolytic and cell wall degrading enzymes (iv) Changes in cellular structure & (iv) Changes in respiration rate and ethylene production.
  • 2. Physiological, Biochemical And Cellular Changes… www.theijes.com The IJES Page 2 II. MATERIALS AND METHODS The fruits of bitter gourd bitter less were collected from local fields at seven sequential stages, starting from very young stage to post-ripening stage. The length and breadth were measured for the collected fruit samples before they were subjected for their physiological and biochemical analyses and their remarkable data were tabulated (Table 1, Pl. 1.1). Biochemical analysis of pigments (Chlorophyll, Carotenoids & Anthocyanins) and primary metabolites (Sugars, Starch, Total Proteins, Total Phenols & Free Amino Acids) were measured from the collected fruit samples by using different methods cited by Thimmaiah (1999). The estimation of nucleic acids (DNA & RNA) was performed by the methods of Devi (2002). Amylase, Invertase, Peroxidase, Polygalactrouronase and Pectin Methyl Esterase were extracted and assayed according to the procedure described by Selveraj and Pal (1984) & Thimmaiah (1999). Data presented at each stage were the mean of three replicates (± Standard error, S.E.). Data of the experiment were subjected to statistical analysis using Duncan’s multiple range test (DMRT) (Bliss, 1967).The histological and ultra structure studies were carried out from the last four stages (including mature stage to post ripening stage) of bitter gourd fruits. The collected fruits were fixed in Formalin Acetic Acid (FAA) (Berlyn & Miksche, 1976). Customary methods described by Johansen (6) were followed for the dehydration and embedding of these FAA fixed materials. The sections of 8 - 10 µm thick were cut with the AO Spencer's rotary microtome from the paraffin embedded materials and then they were stained with safranin - fast green combination. The stained sections of fruit were observed under Carl-Zeiss MP3 400 Microscope and photomicrographs were taken with same microscope with an Image Analyzer Facility. For the TEM studies, the preparation of fruit materials has been done as per the methods cited by Dawes (1971) and the ultra thin sections were observed under the Philips Tecnai (High Tension 200 KV) TEM. The amount of ethylene synthesis and the rate of respiration were measured from mature, pre- ripening, ripening and post ripening stages with the Perkinelmer Autosystem XL Gas Chromatography as per the procedures cited by Thimmaiah (1999). III. RESULTS AND DISCUSSION 3.1.Changes in Pigments:- The quantitative analysis of the Chlorophylls revels that the amount of Chlorophyll a (0.042 mg/gm), Chlorophyll b (0.065mg/gm) and Total chlorophyll (0.107 mg/gm) were very high in very young stage, but as the fruit proceeds towards the maturation, ripening and post ripening stages the amount of chl. a (0.008 mg/gm), chl. b (0.005 mg/gm) and total chl. (0.013 mg/gm) were found declined 5.25 fold, 13.0 fold and 8.23 folds respectively. Carotenoids (0.067 mg/gm) were found gradually increasing from very young stage to almost 2.5 folds at post ripening stage (0. 170mg /gm). Anthocyanins dose not show consistency in their quantity during the maturation, ripening and post ripening stages of fruit. The maximum amount of anthocyanins (6.365 mg/gm) found in the young stage of fruit.Merzlyak et al., 2003 found that during the analysis of pigments, amount of chlorophyll were decline, carotenoids increases due to the pigments changes at terminal stages of leaf and fruit development in many plant species. At these stages plant tissues retain certain amounts of carotenoids or carotenoids synthesis was stimulated on the background of chlorophyll degradation. The increasing level of caroetnoids and decreasing level of chlorophyll also recently noticed by Wang and Zhong (2005) during ripening of tomato fruit. The decreasing level of chlorophyll a, chlorophyll b and Total chlorophyll and increased level of carotenoids were also previously reported in the cucurbits during ripening and storage period by Tan et al., (1999), Rodriguez et al., (1976) and Huong & Raymundo(1999). Anthocyanins were found unstable during ripening which was observed by Underhill and Critchley (1993) in Litchi fruit due to polyphenol oxidase degradation. 3.2.Changes in primary metabolites:- The quantitative analyses of sugars (i.e. total sugars, reducing sugars and non reducing sugars) indicate that the amount of total sugars was gradually increased from very young stage (0.986 mg/gm) to pre-ripened stage (3.047 mg/gm) but there after their amount declines which displayed 209 percent increase. However, the quantum of reducing sugars does not show consistency. In contrast non-reducing sugars was gradually increased from very young stage (0.686 mg/gm) to pre-ripened stage (2.864 mg/gm) which displayed 317 percent increase, but thereafter declining nature was seen in remaining stages.A more or similar observation was also noticed in the apple fruit during there ripening process (Sabir et al., 2004). From the data it was concluded that increase in sugar might be due to hydrolysis of polysaccharides or due to less acidity and ascorbic acid contents. The contents of reducing sugar and non-reducing in spongy tissue may be also due to lower activities of invertase and amylase (Luiz et al., 2001).The amount of starch was observed 1.380 mg/gm at very young stage but thereafter a steady decline was found in the post ripened stage 0.211 mg/gm which displayed 6.5 fold decrease. According to Luiz et al., 2001 during development stage of mango, starch was hydrolyzed. However, as the fruit becomes over-ripe, only traces of starch were detected and amylase activity was substantially reduced.
  • 3. Physiological, Biochemical And Cellular Changes… www.theijes.com The IJES Page 3 The sugars in the pulp spongy tissue were accounted by the presence of higher content of starch, which remained unhydrolysed due the low activity of amylase.The quantitative analyses of the total proteins, free amino acids and total phenols in M. dioica reveal that initially the amount of total protein does not show consistency during their growth development and ripening process. The amount of total protein was decrease from pre-mature stage (7.644 mg/gm) to post ripened stage (2.500 mg/gm), which showed 67.2 percent decrease. The amount of total phenol was gradually increasing from very young stage (2.074 mg/gm) to pre- ripened stage (6.123 mg/gm) which showed 2.95 fold increase thereafter it declines in later stages. In, contrast of proteins and phenols the amount of free amino acids shows 14.4 fold gradual decrease.Hulme et al., (1968) and Dilley (1972) reported that in the early stages of fruit protein synthesis is stimulated and reaches a peak value of climacteric thereafter it declines slowly and gradually during ripening process. An increase in protein synthesis during early climacteric period has been measured in many fruits observed by Sharma (2000). The sharp decrease in free amino acids towards maturity and onset of ripening has been attributed to their incorporation into protein required for the synthesis of various ripening enzymes (Frunkel, Klein and Dilley, 1965). This declining trend of amino acids has been noticed by a number of workers during the approach of fruit maturity (Sharma, (2000)) According to (Dilley, 1970 and Sharma, 2000) the phenolics content of most fruits declines from high levels during early growth to low levels when the fruit is considered to be physiologically mature and therefore, susceptible to the induction of ripening. A more or similar observation was found by Kumar and Goswami (1985) that the presence of higher concentration of phenolics during early stage of development provides protection mechanism to the phytohormones like auxins, gibberellin and cytokines, which play an important role in cell division and cell enlargement. The quantitative analysis of nucleic acids (i.e. DNA & RNA) has been carried out in the seven sequential developmental stages of fruit. The amount of DNA remains inconsistent during ripening while, RNA does not shows consistency in earlier stages. RNA was found decrease from mature stage (0.261 mg/gm) to post ripened stage (0.117 mg/gm), which displayed 55.1 percent decrease. During the climacteric period the RNA was increased and there after its decline up to ripening stage, while in the case of DNA it is fluctuated during the ripening noticed by (Sharma 2000).Amylase activity was found to be gradually declining from very mature stage (0.377 activity) to the post ripened stage (0.050 activity), which displayed 86.73 percent decrease. In contrast, Invertase shows increase in first stage (0.037 activity) and declines upto mature stage (0.019 activity), which displayed 94.3 percent decrease and remains unstable for remaining stages. In case of Peroxidase was gradually increased from very young stage (0.274 activity) to mature stage (0.495 activity) that displayed 44.64 percent increase. These hydrolytic enzymes were observed very high in the early stage of development and thereafter they declined towards the ripening due to starch disappearance, hydrolysis of sugar, climacteric periods or related to growth development process respectively. The activities of hydrolytic enzymes increased from early mature stage to peel colour turning stage and declined in later ripening stage of pineapple fruit during ripening was observed by Selveraj (1993). A more or similar observation also noticed by previously Hulme (1970) and Seymour et al., (1993). The activities of cell wall degrading enzymes viz., Cellulase, Polygalactrouronase (PG) and Pectin Methyl Esterase (PME) during the ripening of fruit were presented in (Table 1.5A, Fig. 1.5B). From analyzed of cell wall degrading enzymes indicate that there was an increment noticed in the activities of cellulase from very young stage (0.013 activity) to ripened stage (0.083 activity) which displayed 6.38 fold increase. PG enzymes were gradually increase 2.37 fold during ripening. While the activity of PME was gradually decline from mature stage (1.009 activity) to post ripened stage (0.800 activity) which displayed 20.7 percent decrease.It has been widely recognized that cell wall changes are related to fruit softening and that these modification are due to the action of cell wall enzymes. The accumulation of these enzymes during ripening and their often-concerted activity will result, eventually, in degradation to many cell wall polymers leading to cell separation and associated softening. During ripening increasing activities of cell wall softening enzymes were reported by Selveraj & Raju (2000) in Kagzi Lime fruit. 3.3.Changes in cellular structure:- Histological Studies:- The development anatomy of fruit has provided information about the prepositional cell division and cell enlargement in various parts of fruit. For better understanding of histological changes the fruits were selected from the mature stage to post ripened stage. It was found that mesocarps cells were divided into three categories (i) Outer mesocarps (ii) Middle mesocarps (iii) Inner mesocarps cell. Middle mesocarps cells and inner mesocarps cells are parenchyma cell, which contain abundant starch grains, as the middle mesocarps cells were comparatively thick; hence they were distinguished from the inner ones.
  • 4. Physiological, Biochemical And Cellular Changes… www.theijes.com The IJES Page 4 At mature stage in the formation of inner mesocarps cells parenchyma cells were very small in size and contained abundant starch due to cell division and cell enlargement. In the starting process of pre-ripened and ripened stage the cell number and cell size were increased while, starch grain was decrease. However, in the last stage the cells were collapsed and empty or contained very less amount of starch grains. During the different developmental stages of Momordica dioica fruit more or similar histological changes was reported by Thanki(1978) (Plate 2.2).White (2002) concluded that “Anatomically, fruit are swollen ovaries that may also contain associated flower parts. Their development follows fertilization, and occurs simultaneously with seed maturation. Initially, fruits enlarge through cell division and then by increasing cell volume”. 3.4.ULTRA STRUCTURE STUDIES:- This study helps in understanding the cell wall softening or changes in cell wall. For this reason the fruit were selected from mature stage to post-ripened stage. In the earlier development stages the cell wall becomes very hard but when the ripening process starts the cell wall becomes soft due to presence of cell wall degrading enzymes polygalactouronase(PG), pectin methyl esterase(PME) and cellulase. The investigations of plasma lamella was observed at mature stage, which was cited near cell wall, while, in pre-ripened stage the cell wall enlarged in their position, which contains granular cytoplasm, ribosome like bodies and nucleus. However, during the ripening and post ripening stages cell wall becomes despoiled. These structural patterns were also previously noticed by Thanki (1978) and Whitaker and Davis(1962) in cucurbitaceae fruit. Brummell et al., (2004) concluded that cell wall softening occur due to differences in cell wall thickness and composition, cell size, cell shape, packing, contents and turgor. Changes in respiration rate and ethylene production:- 3.5.ETHYLENE AND RESPIRATION:- The ethylene production was increase from pre-ripened stage to the post-ripened stage of the fruit, while the rate of the respiration was increased from mature stage to ripened stage, their after decline nature observed in the post ripened stage (Fig 2.6).According to Hardenuburg et al., (1986), Zong et al., (1995), Kays & Hays (1978) and Zheng (1986) conclude that the fruits of bitter gourd (M. charantia) was climacteric fruit due increasing rate of respiration and low production of ethylene. A more or similar observation was also noticed in M. dioica fruit. 3.6.ANALYSIS OF VARIANCE:- The analysis of variance revels that the changes of fruit ripening such as Chl. a, Chl. b, Total chlorophyll, carotenoids, amino acids and the activities of amylase and peroxidase were highly significant (Tables 2.1, 2.3 & 2.4), while less significance was observed in total sugars, starch and cellulase activity (Tables 2.2 & 2.5). However, no significance was found in case of anthocyanins, reducing sugars, non- reducing sugars, total proteins, total phenols, nucleic acids, invertase, PG and PME activities (Tables 2.1, 2.2, 2.3 and 2.5) REFERENCES: [1] Ali, M. and V. Srivastava (1998). Characterization of phytoconstituents of the fruits of Momordica dioica. Indian J. of Pharm. Sci. 60(5): 287-289 [2] Basu, P. S., S. Banerjee, and S. Das 1994. 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