Weak Neck of Musa sp. cv. Rastali: A Review on its Genetic, Crop Nutrition and Post Harvest
Weak neck is the physiological weakening at the pedicel of Musa sp. fruit that causes individual fruits to be released prematurely from the hand. The level of resistance to weak neck is different among groups of Musa sp. with A and B genomes, diploid, triploid and tetraploid. Musa sp. cv. Rastali is AAB genome, but this cultivar is sensitive to weak neck. Sensitivity to weak neck comes from the two A genomes. There is high probability that application of Magnesium (Mg), Boron (B) and Silicon (Si) may solve weak neck on Musa sp. cv. Rastali. Magnesium application increases chlorophyll synthesis and photosynthesis rate. Assimilate accumulation on fruit cells triggers cell elongation and cell wall thickening, especially in the neck zone. Boron plays an important role in regulating hormone and enzyme levels in plants. Sufficient boron in plant tissue impedes synthesis and activities of Pectate Lyase (PL) and pectinmethylesterase (PME), thus decreasing cell wall hydrolysis. Plant cells and tissues become stronger with Si application. This is caused by the formation of silica-cuticle double layer on the epidermis. Low temperature along with low relative humidity during storage and ripening decreases the activities of PL and PME. The reduction of PL and PME activities result in a decrease in pectin degradation rate and cell wall hydrolysis, thus reducing the occurrence of weak neck.
May 02, 2010; Accepted: May 21, 2010;
Published: June 14, 2010
Banana is the worlds fourth most important food crop after rice, wheat
and maize. More people in the tropic and sub-tropic region consume this fruit
on a daily basis (FAO, 2008). Southeast Asia is the centre
origin of bananas. This region comprises of Malaysia, Indonesia, Thailand and
the Philippines. These four countries are important banana fruit producers and
exporters in the world.
Banana is one of the main fruits exported and it is very popular as compared
to other fruit. In particular, Malaysia is one of the worlds major exporters
of bananas. Since 1990, Malaysia has been gaining recognition for its export
of local fruit to overseas market. In 2007, export of bananas reached 33838
tons (FAO, 2008). The cultivation areas for bananas had
therefore increased from 23,266 ha in 1985 to 38,678 ha in 1994, 54,138 ha in
2000 and 72,254 ha in 2007 (Department of Agriculture Malaysia,
2008). Musa sp. cv. Rastali is one of the common dessert bananas
in Malaysia. Total cultivation areas for Musa sp. cv. Rastali was 7.14%
of the total area for banana in 2007. That was approximately 5,158.94 ha, with
the total production of approximately 57321 t. This cultivar has AAB genome
(Ploetz et al., 2007).
Musa sp. cv. Rastali has a good potential of being exported to overseas
markets (Hassan, 2002). However, there are many obstacles
faced by the farmers that are interrelated with the development of Musa
sp. cv. Rastali as a superior commodity, particularly on productivity and post
harvest handling. Two of the main obstacles faced by farmers are packing and
dispatching the fruits to the market. Long distance transportation cause fruits
to be released from its hand. This is called weak neck. Weak neck is the physiological
weakening at the pedicel of the fruit that causes individual fruit to be released
prematurely from its hand (Saengpook et al., 2007).
However, weak neck is not prevalent to all Musa sp. cultivar. One of
the cultivars that is susceptible to weak neck is Musa sp. cv. Rastali.
Weak neck decreases the quality of Musa sp. cv. Rastali fruit. The fruits
thus become unacceptable and the farmer suffers financial losses (Daddzie
and Orchard, 1997). Besides that, the development of Musa sp. cv.
Rastali as a potential export commodity is undermined. These factors caused
Musa sp. cv. Rastali to be merely used for local consumption.
A holistic approach is needed in order to solve the weak neck problem of Musa
sp. cv. Rastali. The three aspects that were reviewed were plant genetics, plant
nutrition and post harvest. This study discusses recent information and possible
solutions to solving weak neck in Musa sp. cv. Rastali.
After harvesting, there are many factors that can decrease the quality of Musa
sp. cv. Rastali fruits. Generally, the obstacles can be classified in to two
groups, i.e., mechanical injury and physiological disorder. Mechanical injury
is considered as a type of stress that occurs during the post harvest handling
of fruits. Mechanical damage is defined as plastic deformation, peel ruptures
and tissue damage caused by external factors (Del Aguila
et al., 2007). There are three main sources of mechanical injury
to banana; these are impact, pressure (compression) and cut (Daddzie
and Orchard, 1997). Impact is generally caused by a collision between the
fruit with a solid surface and between the fruit during harvest, handling and
transportation. Compression injuries are caused by the variable pressure exerted
by the adjacent fruit or fruit with a container. Meanwhile, cuts injuries are
generally caused by collision between the surface of fruit with a sharp object
that caused epidermal ruptures, or because of pressure caused by uneven surfaces,
like the edge of the container of fruit (Del Aguila et
Physiological disorder occurs on fruit tissue and is not caused by pathogen.
Physiological disorder develops as plant response to environmental conditions
such as temperature and lack of nutrition (Wills et al.,
1989). Most of the physiological disorders affect discrete areas of the
tissue. Some disorders may affect the skin of the fruit but may leave the underlying
flesh intact; others affect only certain areas of the flesh.
The most important physiological disorder affecting Musa sp. cv. Rastali
is weak neck. Weak neck has been defined by Baldry et
al. (1981) as the physiological softening and weakening which cause
individual fruits in a hand to separate from the crown. Hands of banana with
missing fingers cant be sold to consumers and individual fruits that have
dropped have no pedicel and cant be marketed (Imsabai
et al., 2006). For this reasons, weak neck has become a major problem
for Musa sp. cv. Rastali (Putra et al., 2010).
Weak neck reduces the quality and selling value of Musa sp. cv. Rastali,
resulting in financial loss (Daddzie and Orchard, 1997).
There are three possible causes of weak neck: genetics (Semagn
et al., 2006), nutrition (Williams, 2002)
and post harvest handling (Sairam et al., 2008).
An understanding concerning the relationship among Musa sp. cv. Rastali
genetics, nutrition and post harvest handling with weak neck is needed in order
to solve weak neck on Musa sp. cv. Rastali.
Genetics is one of the factors that determine the occurrence of weak neck on
Musa sp. cv. Rastali. The information concerning the relationship between
genetics and weak neck on Musa sp. cv. Rastali however is still limited.
Information available is still based on the morphological character of Musa
sp. (Hicks, 1934; Marriott, 1980;
New and Mariott, 1983; Semple and
Thompson, 1988; Pereira et al., 2004).
The level of resistance to weak neck is different among Musa sp. with
A and B genomes. Generally, the Musa sp. with B genome are more resistant
to weak neck than those with A genome. The level of resistance to weak neck
is also different among Musa sp. group of diploids, triploids and tetraploids.
The diploid group of Musa sp. is more resistant than triploids and tetraploids,
while triploids are more resistant than tetraploids. Musa sp. within
the tetraploid group are the most sensitive to weak neck (Pereira
et al., 2004).
Hicks (1934) and Semple and Thompson
(1988) reported that triploid Cavendish AAA group was the most sensitive
to weak neck. Marriott (1980) reported that all of the
Musa sp. tetraploids were sensitive to weak neck. However, Pereira
et al. (2004) showed that all of the diploid Musa sp. with
BB genome were resistant to weak neck. Some triploid Musa sp. were also
resistant to weak neck. Triploid Musa sp. that possessed intermediate
resistance to weak neck were Cavendish (AAA) and Grande Naine (AAA). Similar
to the findings of Marriott (1980), all of the tetraploid
Musa spp. experienced weak neck, i.e., Pioneira (AAAB), YB42-21 (AAAB),
Buccaneer (AAAA) and Calypso (AAAA) (Pereira et al.,
2004). Based on previous studies Musa sp. that were resistant to
weak neck contained the B genome, either in diploids and triploids. Thus, it
can be concluded that the carrier of resistance to weak neck is the B genome.
Musa sp. cv. Rastali is a triploid with an AAB genome, but this cultivar
is sensitive to weak neck. Sensitivity to weak neck apparently comes from the
two AA genomes present in Musa sp. cv. Rastali.
There was a strong relationship between weak neck and fruit morphological characteristic
mainly fruit firmness, peel and pulp colour, peel and pulp thickness, pulp to
peel ratio and pedicel. A morphological characteristic of the fruit is the interaction
between genetics and environmental factors (Pereira et
al., 2004). Some time, fruit morphological characteristic of crops under
limited conditions were better compared with normal conditions.
Molecular markers-assisted selection is already helping breeders to improve
stress-related traits (Tuberosa and Salvi, 2006; Niknejad
et al., 2009). The best marker for Musa sp. is microsatellites
because it is highly polymorphic, show a co-dominant mode of inheritance and
easy to interpret (Semagn et al., 2006). DNA
microsatellites are polymorphic elements which are abundant in nuclear genomes
and consist of a succession of a small repeat-unit, usually less than four nucleotides
long. Fragments containing microsatellites can be amplified through Polymerase
Chain Reaction (PCR) (Creste et al., 2003). The
use of molecular markers makes possible to find a solution to the weak neck
problem of Musa sp. cv. Rastali.
Ferreira et al. (2004) said that several characteristics
of Musa sp. cv. Rastali depends on their genes, for example, their resistance
to diseases, height of crop, number of finger/bunch and length of fingers. Even
so, those characteristics can be influenced by environmental conditions. One
of the environmental conditions mentioned is nutrient supply from the growing
media. Nutrient supply for plant depends much on nutritional management. Optimal
nutritional management is essential to produce adequate nutrient supply to the
Growth and yield of Musa sp. cv. Rastali get the maximum if the mineral
nutrient in sufficient quantities. Some ways can be done to meet the needs of
the mineral nutrients are fertilizer, inoculation of Plant Growth-Promoting
Rhizobacterial (PGPR) and increase the efficiency of nutrient uptake by roots
(Mia et al., 1998, 2005;
Zakaria, 1998; Zakaria et al.,
1998). Inoculation of PGPR on Musa spp. grown under hydroponic conditions,
combined with 33% fertilizer-N of the total N requirement, could produce better
growth and yields than the Musa sp. that get 100% of N-fertilizer. It
shows that the application of PGPR can reduce the supply of N-fertilizer, because
PGPR play a role as bioenhancer and biofertilizer. Application of PGPR also
increases the uptake of Phosphorus (P), Potassium (K), Calcium (Ca) and magnesium
(Mg), thus increasing its concentration in tissue (Mia et
al., 1998, 2005; Zakaria
et al., 1998). On the other hand, the efficiency of mineral nutrient
uptake can be enhanced through root system improvement, which can be achieved
through plant breeding programs (Saleh and Gritton, 1994).
The application of several macro and micro elements such as magnesium (Mg)
and boron (B) are able to increase the availability of those nutritions for
Musa sp. cv. Rastali. The availability of Mg and B was reported to increase
the productivity of Musa sp. cv. Rastali and decrease finger drop (Matoh
and Kobayashi, 1998; Mostafa et al., 2007).
It is related with the important roles of Mg and B in plant physiological activities
such as photosynthesis and synthesis of enzyme, hormone and protein (Jones
et al., 1991; Mostafa et al., 2007;
Singh et al., 2007; Rashid
et al., 2007). Moreover, B also has the role in strengthening cells
and tissues. Boron is also an important inorganic component in the cell wall
(Matoh and Kobayashi, 1998).
On the other hand, Silicon (Si) is a beneficial element for monocots. Application
of this element had a positive effect on plant growth, yield and plant resistance
to pest and plant disease (Basagli et al., 2003;
Ma et al., 2004; Almeida
et al., 2009; Gorecki and Danielski-Busch, 2009).
Silicon has a positive effect in metabolism, i.e., improvement of nutrient imbalance,
reduction of mineral toxicities, improvement of mechanical properties of plant
tissues and enhancement of resistance to other various abiotic and biotic stress
(Iwasaki et al., 2002; Kim
et al., 2002; Henriet et al., 2006,
2008). In such cases, the positive effect of Si addition
to the plant under optimum conditions is generally not too significant. The
positive effect of Si will appear significantly when the plant is subjected
to stress (Epstein, 1994; Henriet
et al., 2006).
Henriet et al. (2006) discovered that bananas
planted on vertisol are more resistant to fungi. This resistance is related
to Si because vertisol has high Si availability. Results from Hinsinger
et al. (2001) and Rufyikiri et al. (2004)
also showed that the roots of banana were able to induce Si dissolution so as
to increase the availability of Si in the rhizosphere. This information shows
that banana plant benefits from Si accumulation, but the mechanism is still
Application of nutrition can also be used in for disease control. Controlling
of plant disease by managing the nutrition application pattern gave a better
yield than using pesticides. Fawe et al. (2001)
reported that Si given to plants cultivated through hydroponics had a positive
effect in controlling several diseases. As an example, application of Si on
melon, cucumber and strawberry that were planted through hydroponics increased
plant resistance to powdery mildew disease (Fawe et al.,
Datnoff et al. (2001) said that the application
of Si can decrease the main disease aggression level on paddy, i.e., blast,
brown spot, sheath blight, leaf scald and grain discoloration. These diseases
were caused by fungi aggression. Controlling diseases that are caused by fungi
with Si application gave better yield than fungicide application. This condition
will decrease the intensity of fungicide application and help achieve agriculture
Weak neck is caused by physicochemical changes during fruit growth and maturation.
During fruit maturation, content of glucose, fructose, sucrose and total sugar
increase but the starch decreases (Osman et al.,
2000). This condition causes the rigidity of cells in peels of Musa
sp. cv. Rastali fruit to decrease rapidly, which eventually leads to weak neck.
Ultrastructural study on six cultivars of Musa sp., i.e., Rastali, Awak,
Tanduk, Abu, Berangan and Lemak Manis showed that cells of Musa sp. cv.
Rastali collapsed completely after ripening and cells of Musa sp. cv.
Awak became thinner making these two cultivars prone to weak neck. The cells
of the other four cultivars became elongated and the cell wall became thicker
after ripening, thus not showing weak neck symptom. Cultivars with elongated
cells at the neck zone and thick cell walls after ripening are more resistant
to weak neck. Cell elongation and the thickening of cell wall after ripening
can be modified by nutrient application of Mg, B and Si during planting (Putra
et al., 2010).
Marschner (1986) and Williams (2002)
reported that some elements are important in structural integrity, i.e., P,
Ca, Mg, B and Si. Deficiency in these elements decreases structural integrity
of cells and this condition is a possible cause of weak neck. Adequate Mg, B
and Si in plant tissues can increase the strength of cells, tissues and organs
by increasing lignin in tissues (Salerno et al.,
2004). Lignin is a polymer of phenolics, especially phenylpropanoids and
is primarily a strengthening agent in the wall. It also defends against fungal/pathogen
attack (Salerno et al., 2004). Besides lignin,
other organic materials in cells that are important to cell strength are suberin,
wax and cutin which are associated with wall waterproofing (Richard,
2008). Depositions of Mg, B and Si in the plant tissue increase the thickness
of cell wall and size of vascular bundle (Williams, 2002).
Thicker cell walls and bigger vascular bundle increases the strength of the
Magnesium is a component of chlorophyll hence, magnesium application on Musa
sp. cv. Rastali may increase chlorophyll synthesis. An increase in leaf chlorophyll
content result to an increase in the photosynthesis rates and also Net Assimilation
Rate (NAR). When assimilate concentration on plant tissue increases part of
it is be translocated to fruit cells. Assimilate accumulation on fruit cells
induces cell elongation and the thickening of the cell wall, especially at the
neck zone (Smithson et al., 2004).
There is a high probability that application of B may resolve weak neck on
Musa sp. cv. Rastali. Boron plays an important role in regulating hormone
and enzyme levels in plants (Rashid et al., 2007).
For example, sufficient B concentration in plant tissue will impede synthesis
and activities of Pectate Lyase (PL) and pectinmethylesterase (PME), where PL
and PME are enzymes that stimulate pectin degradation and cell wall hydrolysis.
Reduction of PL and PME decreases pectin degradation rate and cell wall hydrolysis
and finally reduce weak neck on Musa sp. cv. Rastali.
Silicon is another beneficial element for monocots including Musa sp.
Accumulation of Si during stressed condition will impede destruction of plant
cells and tissues. Plant cells and tissues become stronger when the formation
of silica-cuticle is doubled on the layer of epidermis (Fawe
et al., 2001; Datnoff et al., 2001).
Apart from the role of Mg, B and Si individually, application of Mg, B and Si concurrently appears to be interrelated in plant. Balanced nutrition is essential for optimum crop growth and maximum quantity and quality of yield. Studies concerning nutrition management for resolving weak neck on Musa sp. cv. Rastali have never been reported, therefore, this aspect is necessary for further investigation.
Information on post harvest of Musa sp. cv. Rastali aids in determining the occurrence or absence of weak neck. Several treatments are often applied to Musa sp. cv. Rastali which had been harvested in order to reduce the risk of weak neck. Several post harvesting techniques which were applied for reducing weak neck are usually applied during ripening and post harvest. Compared to the other aspects, i.e., genetic and plant nutrition, modification of environmental condition during post harvesting and ripening for eliminating of weak neck is the best to be done.
Several studies have shown that weak neck occurred because of the relative
humidity and the extreme temperature in storage and during ripening (Semple
and Thompson, 1988). High relative humidity increases the activity of Pectate
Lyase (PL) and pectinmethylesterase (PME). High PL and PME activity increases
pectin hydrolysis. The concentration of water soluble pectin in the peel and
pedicel is usually high and this can decrease fruit firmness. The peel of Musa
sp. fruit also become soft and easy to breaks. This condition is also observed
on the pedicel. The pedicel becomes weak thus weakening the neck region or abcission
layer (Imsabai et al., 2006; Saengpook
et al., 2007).
Another factor is ethylene where weak neck decreases if the concentration of
ethylene in the storage and ripening process increased. High ethylene concentration
accelerates ripening and water loss. This condition was likely the cause of
weak neck (Paull, 1996), which is similar to the mechanism
during leaf abscission (Sexton and Roberts, 1982).
According to Seymour (1993), softness of Musa
sp. cv. Rastali fruit peel is the same as in the other tissues. This is because
depolymerization of pectin in the primary cell wall and middle lamella. Besides
that, the softening of peel on Musa sp. cv. Rastali fruit also involved
cell wall hydrolysis.
Weak neck on Musa sp. cv. Rastali is not related to the activities of
endopolygalactoronase (endo-PG) and exo-PG, but it relates with the activities
of PL and PME. The fact shows that weak neck is not caused by abscission but
it is more likely caused by the breaking of the peel at the junction of the
fruit and the hand. Previous research from Putra et al.
(2010) showed that weak neck in Musa sp. cv. Rastali was caused by
disintegration, collapse and thinning out of cells along the neck region. Imsabai
et al. (2006) had determined that weak neck was not caused by fruit
weight, peel thickness and water content of the peel at the pedicel area.
Based on the above explanation, more experiments on post harvesting should
be done to reduce of the activities of PL and PME. Result showed that the low
relative humidity and temperature during storage and ripening of Musa
sp. cv. Rastali were able to decrease the activities of PL and PME. The reduction
of PL and PME decreases pectin degradation rate and cell wall hydrolysis. Slow
pectin degradation and cell wall hydrolysis reduces weak neck (Saengpook
et al., 2007). Ethylene application during ripening will speed up
fruit ripening and water loss. Musa sp. cv. Rastali which has ripened
quickly and underwent drastic reduction in water content would not experience
to weak neck (Paull, 1996).
The most important problems related to the physiology of Musa spp. are weak neck, breakage of peel and destruction caused by cold temperature. Among those, weak neck is the main problem of Musa sp. as a commodity. Weak neck is the physiological weakening at the pedicel of Musa sp. fruit that causes individual fruits to be released from its hand prematurely.
The level resistance to weak neck is different among groups of Musa spp. with A and B genomes, diploid, triploid and tetraploid. Musa sp. group for B genome is more resistant to weak neck than A genome. The diploids group of Musa sp. is more resistant than triploids and tetraploids, while triploids are more resistant than tetraploids. Musa sp. cv. Rastali is AAB genome, but this cultivar is sensitive to weak neck. Sensitivity to weak neck is thought to come from the two A genomes.
There is high probability that application of Mg, B and Si can resolve weak neck on Musa sp. cv. Rastali.
Magnesium application increases chlorophyll synthesis and photosynthesis rate. Assimilate accumulation on fruit cells induces cell elongation and the thickening of cell wall, especially in the neck zone. Boron plays an important role in regulating hormone and enzyme levels in plants. Sufficient amounts of B in plant tissue impede the synthesis and activity of PL and PME. Plant tissue and cells become stronger with Si application. This causes the formation of silica-cuticle double layer on the epidermis.
Low temperature along with relative humidity in storage as well as the process of ripening can reduce the weak neck problem because these conditions are able to decrease the activities of PL and PME. The reduction of PL and PME result to a decline in pectin degradation rate and cell wall hydrolysis. Slow pectin degradation and cell wall hydrolysis can reduce weak neck on Musa sp. cv. Rastali.
This study was funded by Universiti Putra Malaysia under the Research University Grand Scheme (Project No. 01/01/07/0004/RU). The authors would like to express their gratitude to Mr. Mohd Shahril Ab. Rahman and Mr. Daud Mustam for their assistance in the study.
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