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CHAPTER II

REVIEW OF LITERATURE

Modern scientific research on bamboo has a history of more than 150 years. Most
research works on management and utilization of bamboo has been focused in Southeast
Asia, especially the three major bamboo bioresource available countries ie. China, Japan
and India (Sharma et al., 2008). The comprehensive utilization of bamboo was
extensively observed in China, Japan and Korea, where the industries related to bamboo
carbon and bamboo distillate has reached large scale levels. In India the main focus of
utilization of the abundant bamboo resource especially in natural forests has been for
pulp. Not much research has been reported on bamboo utilization in developed countries
such as Europe and USA since these countries lack the bamboo resource and the
corresponding bamboo industry (Nadgauda et al.,1993).Both American and African
continents have substantially lower bamboo species richness compared to Asia. The
highest species richness in Africa could be found in Madagascar and Brazil has the
greatest bamboo diversity in Latin America (Londono, 2001).

Bamboo is a group of monocotyledonous, fast growing, perennial, tall arborescent grass


that taxonomically belong to the subfamily of Bambusoideae under Gramineae Family
(Chapman, 1996 & 1997). Bamboo can thrive in hot, humid rainforests and can grow
well in cold temperature as low as -200C. It also can survive in regions of high
precipitation ranging from 80-130 cm annual rainfall (Goyal et al., 2012). Bamboo is
known to be one of the fastest growing grass in the world, with a growth rate ranging
from 30 to 100 cm per day in growing season. It can grow to a height of 36 m with a
diameter of 1–30 cm. A culm can reach its full height in about two to three months. As a
major non-wood forest product and wood substitute, bamboo is of increasing interest to
ecologists owing to its rapid growth and correspondingly high potential for mitigating
climate change. If well-managed, bamboo plantations can conserve soil and water, and
improve soil fertility and local climate as well (Troya et al., 2014). The objective of this
review is to assess the value of bamboo shoots as a potential new age health food and
identify species having prospects for promoting commercial plantation in the light of
available scientific results. The review also examines input management approaches to
realize this seemingly under exploited potential of bamboo. Hence, an updated review of
available literature pertaining to the objectives of the study and other relevant information
pertinent to bamboo shoots are presented in this chapter.

2.1 Bamboo - Distribution

Bamboo is considered to be one of the fastest growing, highest yielding renewable


natural resource. It is estimated that worldwide there are over 1200 species under 87
genera of bamboo, with a natural distribution across a broad range of environments
(humid tropical, subtropical and temperate region) (Ram et al., 2010; Hogarth and
Belcher, 2013). The major bamboo producing countries in Asia are China and India
accounting to approximately 70% of the bamboo in Asia. The American continent which
includes South America, Central America and North America, has a high biodiversity of
bamboos with approximately 400-500 species; among them about 300 species in 20
genera are woody bamboos. Though Africa has barely about 40 species, rather poor in
bamboo flora with only a few native species, it has large areas of natural bamboo forests
of Ethiopia, Kenya, Uganda and other African countries (Lobovikov et al., 2007; Ha,
2010; Yang and Hui, 2010). Due to its vast territory, and diverse climate, China has the
richest bamboo resources in the world with approximately 590 species falling under 39
genera (Yue, 2012) and hence often referred to as the "Kingdom of Bamboo" (Song et
al., 2011).

Worldwide, bamboo with over 1200 species occur in natural forests, semi-exploited
stands, and intensive plantations, covering an area of more than 14 million ha (Dransfield
and Widjaja, 1995; Fu and Banik, 1995). Eighty percent of the species and area are
confined to South and Southeast Asia and mostly in China, India, and Myanmar. Of
these, over 30% is covered by the world’s single most important and largely exploited
bamboo species, Phyllostachys pubescens, commonly known as Moso bamboo (Li and
Xu, 1997). India is the second largest producer of bamboo in the world next only to
China and also has a very high diversity of bamboos with 136 species in about 18 genera
(Sharma et al., 2008, Gulabrao et al., 2012). In India, bamboo is cultivated in about 8.96
million hectares of land, including forest land, homesteads and private plantations, which
account for nearly half the total area under bamboo cultivation in Asia. The areas
particularly rich in bamboo are the Northeastern states, the Western Ghats, Chattisgarh,
Madhya Pradesh and Andaman Nicobar Islands. The important genera are Arundinaria,
Bambusa, Cephalostachyum, Dendrocalamus, Dinochloa, Gigantochloa, Melocanna,
Ochlandra, Oxytenanthera, Phyllostachys, Pseudostachyum, etc (Jeeva et al., 2009;
Pandey and Ojha, 2013). North-east India contributes more than 66% of total bamboos
resources available in India (Sarmah et al., 2000). About 16 edible bamboo species are
reported in Northeast India (Singha et al., 2008) and some of the important edible
bamboo species are Arundinaria callosa, Bambusa nutans, Bambusa pallida, Bambusa
polymorpha, Bambusa tulda, Dendrocalamus hamiltonii, Dendrocalamus giganteus,
Dendrocalamus brandisii, Melocanna baccifera, Melocanna bambusoides,
Dendrocalamus hookeri, Dendrocalamus sikkimensis, Dendrocalamus strictus, and
Phyllostachys manni. The Western Ghats extending from Maharashtra up to Kerala is
endowed with some important genera of bamboo, including different species and
subspecies like Bambusa arundinaceae, Dendrocalamus strictus, Pseudoxytenanthera
ritcheyi, P.stocksii (D.stocksii), Ochlandra spp. (Uma Shaanker et al., 2004; Kiruba and
Jeeva, 2010; Jeeva and Sheeja, 2013).

2.2 Bamboo - Production and Market status

Bamboo has become an important trade commodity globally providing direct or indirect
livelihood support to about 2.5 billion people (INBAR, 1999). The international trade in
bamboo ranges between 5 to 10 billion USD. More than 2 million tons of bamboo shoots
are consumed annually (Kleinhenz et al., 2000) with approximately 1.3 million tonnes
produced in China alone (Shi et al., 1997). Most of the bamboo traded internationally is
exported from China ranging from 75% to 95% of total bamboo traded in the world
(Parker, 2005). Total trade in bamboo products was estimated at around 4.5 billion USD
/year (Sastry, 1998). The range of uses of bamboo for mankind is remarkable, with an
estimated annual use of 12 kg of bamboo products per capita in Asia (Recht and
Wetterwald, 1988; Sastry, 1998). Besides some minor uses such as leaves for medical
purposes (Zhang, 1997), fresh edible shoots, and culms for timber or as a raw material for
pulping are the major products from bamboo. Bamboo exploited for timber is much
greater, e.g., it was 3 million tonnes/year in India (Subramanian, 1995), and possibly
more than 20 million tonnes/year in China, and most likely about 30 million tonnes/year
worldwide. Apart from its application as raw material for various immediate uses to
humans, bamboo also serves multiple ecologic functions such as soil and water
conservation, and erosion control (Fu and Banik, 1995). Bamboo has great potential for
rapid biomass production (Pearson et al., 1994), thus acting as a significant net sink for
global CO2 (Jones et al., 1992). The annual production of bamboo in India was estimated
at 4.6 million tonnes of which about 1.9 million tonnes was used by the pulp industry.
Total number of culms at the national level has been estimated to be about 23,297 million
out of which 79% are green, 16% are dry and 5% are decayed (FSI, 2011). The value of
international bamboo market was estimated to be around 10 billion USD (Borah et al.,
2006).

The total revenue from bamboo shoots was about 1.2 billion USD while other bamboo
based products accounts for around 3 billion USD (Anon, 2003). Bamboo shoot is
consumed worldwide and annual world bamboo consumption was estimated to be over 2
million tonnes (Yang et al., 2008).In international markets, China earns 6,500 million
INR every year from export of edible bamboo shoots, with import by USA at around
44,000 tonnes accounting for 14.5% of the total world import (Lobovikov, 2003). Every
year, it was estimated that USA imports 30,000 tonnes of canned bamboo shoots from
Taiwan, Thailand and China for domestic consumption as food items (Daphne, 1996).
Dendrocalamus asper, Dendrocalamus lactiferous and Bambusa oldhami are the most
important edible species in Thailand (Fu et al., 1987) and Taiwan (Tai, 1985),
respectively. The import of shoots in Australia was estimated about 8,000 tonnes per
annum (Cahill, 1999). Consumption of shoots in Taiwan was estimated at about 80,000
tonnes of bamboo shoots annually valued at 2,500 million INR, covering 30,000 ha
under cultivation of bamboo shoots, producing total 380,000 tonnes of bamboo shoots per
year (Tai, 1985). In Japan, the reported annual consumption of bamboo shoots was 3 kg
per person in 2008, as compared to 1.2 kg per person in 1950s (Yang et al., 2008). In
2005, yearly over 2 million tonnes of edible bamboo shoots were reported to be
consumed around the world, mostly in Asian countries (Vaiphei, 2005). India is the
second richest country for bamboo production. About 26.2 tonnes, 435 tonnes and 426.8
tonnes of bamboo shoots were reported to be harvested in Sikkim, Meghalaya and
Mizoram, respectively (Bhatt et al., 2003). About 78 species of indigenous and exotic
varieties of bamboo are grown in North Eastern India (Tamang and Tamang, 2009). The
annual average consumption of bamboo shoots in the north eastern states of India was
estimated to be 1979 tonnes, 2188 tonnes, 442 tonnes, 433 tonnes, 442 tonnes and 201
tonnes in Arunachal Pradesh, Manipur, Meghalaya, Mizoram, Nagaland and Tripura,
respectively (Sarangthem & Singh, 2003a). India’s size of domestic bamboo economy
was estimated at 2,000 million INR. The market potential of bamboo in India was
estimated at 450 million INR, which was expected to increase to 26,000 million INR by
2015, thus enabling five million families of artisans and farmers, crossing the poverty
line (Farooquee et al., 2007).

Bamboo is not only an ideal economic investment with multifarious uses but also has
enormous potential for alleviating many environmental and social problems. Increasing
rates of deforestation and demand for wood and need for sustainable bioresource makes
the search for alternative natural resources more important (Zhou et al., 2005). Bamboo is
one of the most valuable natural resource due to its utilization value, wide distribution,
availability, rapid growth, easy handling and desirable properties. Also known as ‘Green
gold’, bamboo is comparatively cheap and commonly used in the daily life of local
communities in South-east Asia for a wide range of purposes. The people of Asia, Africa
and South America are largely dependent on it for their house construction and
agricultural implement. Bamboo has more than 1500 documented uses (Rai and Chauhan,
1998) and around 5000 listed uses.
(http://www.bambootech.org/tslink.asp?subsubid=71&subid=16&sname=mission&lid=2
29). Bamboo is becoming popular as an excellent substitute for wood in paper industries,
domestic commodities, cottage industries, board and charcoal (Lobovikov et al., 2007).

Bamboo is highly suited to diversified agricultural systems, constituting one of several


livelihood resources for farmers. Sustainable bamboo plantations provide direct
employment for many rural unskilled people in areas where opportunities for economic
development are low. Yet for a multitude of reasons, the market for bamboo within India
is still in its beginning. The lacunae between agronomists, financiers and potential end
users have resulted in the slow commercialization of bamboo (INBAR, 1999).
2.3 Bamboo shoots

A new emerging young culm is edible and is known as bamboo shoot or juvenile shoot. It
is actually a culm that emerges from the ground in full diameter and contains nodes and
inter nodes in a vertically miniaturized form. The young shoots are tightly clasped with
overlapping sheaths that have to be removed to extract the edible part. New culms or
juvenile shoots in bamboos usually emerge with the beginning of the monsoon season
and are harvested for edible purposes (NMBA, 2004). Bamboo shoots have been eaten as
a vegetable for thousands of years in many Asian countries and are becoming more and
more popular due to their immense health benefits (Nirmala et al., 2011).

China has a long tradition of cultivating, processing and preserving bamboo shoots as
food (Xiao and Yang, 2001). As early as in 1951, production and use of fine quality
edible bamboo shoots in eastern China and Japan was reported (Porterfield, 1951). Most
bamboo species produce edible shoots but less than 100 species out of 1200 recorded are
grown or utilized for their shoots (Midmore, 1998; Collins and Keilar, 2005). Fresh
shoots have a crispy crunchy texture and a unique taste. Bamboo shoots are eaten as
vegetable mostly in South-East and East Asian countries. Bamboo shoots are now
processed into many kinds of food, including fresh shoots, dry shoots, and canned shoots,
that are sold around the world. Some new products, such as bamboo candy, bamboo
chutney, bamboo canned juice, bamboo beer are also available. Bamboo vinegar is also
used as biofertilizer, bioinsecticide and as medicine for stomach disorders (Song et al.,
2011). Fresh bamboo shoots are delicious and healthy (Lobovikov et al., 2007). They are
brittle, tender, pollution free, low in fat, high in edible fiber and rich in mineral elements.
China has the largest bamboo industry producing approximately 1.3 million metric tonnes
of fresh bamboo. Worldwide, more than 2 million tonnes of bamboo shoots are consumed
annually of which about 1.3 million tonnes are produced in China alone (Kleinhenz et al.,
2000).
2.4 Bamboo shoots – Nutritional Composition

The nutritional value of edible shoots of different bamboo species has been worked out
by several researchers (Giri and Janmejoy, 1992; Shi and Yang, 1992; Tripathi, 1998;
Chen et al., 1999; Sharma et al., 2004; Xu et al., 2005; Kumbhare and Bhargava, 2007;
Nirmala et al., 2007, 2008). Bamboo shoots are low in calories, high in dietary fiber, and
rich in various nutrients. The main nutrients in bamboo shoots are protein, carbohydrates,
amino acids, minerals, fat, sugar, and fiber. The shoots have a good composition of
minerals, consisting mainly of potassium (K), calcium (Ca), manganese (Mn), zinc (Zn),
chromium (Cr), copper (Cu), iron (Fe), and lower amounts of phosphorus (P), and
selenium (Si) (Shi and Yang, 1992; Nirmala et al., 2007). Juvenile shoots are a good
source of thiamine, niacin, vitamin A, vitamin B6, and vitamin E (Visuphaka, 1985; Xia,
1989; Shi and Yang, 1992; Zhang and Liu, 2001) and rich source of dietary fiber
(Nirmala et al., 2009). Sucrose is the most abundant sugar in bamboo shoot. Bamboo
shoots also contain high proportion of linoleic acid. The major fatty acid in bamboo
shoots is palmitic acid; glutamic acid and lysine are the most abundant amino acids in
bamboo shoots (Soo-Jung and Sung-Ja, 1993). Bhargava et al., (1996) have reported that
bamboo shoot contains appreciable quantities of vitamin C, carbohydrates and protein.

However, bamboo shoots of many species also contain lethal concentration of cyanogenic
glucosides, which on endogenic hydrolysis, yield hydrocyanic acid. Cooking destroys the
enzymes responsible for the endogenic hydrolysis largely. Another constituent of bamboo
shoot, known as homogentistic acid is responsible for the disagreeable, pungent taste,
characteristics of bamboo shoots (Ferreira et al., 1991).

2.4.1 Protein

Nirmala et al., 2011 reported that bamboo shoots are rich in protein containing between
1.49 and 4.04 g (average 2.65 g) per 100 g of fresh bamboo shoots. In a study conducted
on 14 bamboo species, the protein content in the juvenile shoots ranged from 2.31 to
3.72 g/100 g fresh weight with the highest being in D. hamiltonii followed by B. bambos.
Similar values were also reported by other workers (Sundriyal and Sundriyal, 2001; Bhatt
et al., 2005b). Shoots contain 17 amino acids, 8 of which, serine, methionine, isoleucine,
leucine, phenyalanine, lysine, and histidine are essential aminoacids (Qiu, 1992). A
comparitive analysis by Nirmala et al., 2011 between two bamboo species (B.tulda and
D.hamiltonii) and 15 common vegetables shows that both the bamboo species are rich in
proteins and also the protein content is much higher than some of the common vegetables
like Brassica oleracea var.capitata (Cabbage); Solanum melongena (Brinjal). The
recommended dietary allowance for protein is 0.8 g/kg of body weight for adults. In
general, 1 g protein provides 4.2 kcal of energy to the body. If the diet does not contain
sufficient amounts of carbohydrate and fat, then dietary protein may be broken down to
provide energy (RDA, 2009). Since bamboo shoots have an average protein content of
2.65 g/100 g fresh weight, consuming the shoots will supply a generous amount of
protein to the body.

2.4.2 Fibre

Dietary fiber includes cellulose and lignin, hemicelluloses, pectins, gums, and other
polysaccharides and oligoscacharides associated with plants (Chawla and Patil, 2010)
and has been associated with a number of health benefits, such as bowel protection,
increasing the time taken for the body’s waste to be moved out of the body, reduced
exposure of the body to carcinogens in food and fluids, and an increase in the amount of
butyrate, the preferred energy source for cells called colonocytes. An increase in dietary
fiber reduces the blood pressure (Anderson and Strong, 1983). According to George et
al., (1982), dietary fiber is useful in the management of hypertension and obesity through
its effect on energy density of food and the extent of interference with the nutrients of
bioavailability. The recommended level of fiber for adults is 25 to 30 g a day, in
combination with at least 2 liters of fluid to ensure thorough digestion. Extracts rich in
dietary fiber obtained from plants could be used as functional ingredients as they improve
not only digestive health, but weight management, cardiovascular health, and general
wellness. The role of fiber in preventing CVDs has been well documented (Viuda-Martos
et al.,2010).

Bamboo shoots are a rich source of dietary fiber (Nirmala et al., 2009) having high
amounts of fiber, ranging from 2.23 to 4.20 g/100 g fresh weight of shoot in some
bamboo species. As a dietary fiber source, the shoots have beneficial effects on lipid
profile and bowel function. Studies conducted by Park and Jhon (2009) have reported the
beneficial effects of consuming a high-fiber diet containing bamboo shoots in lowering
blood cholesterol levels and improving bowel functions in healthy young women. Park
and Jhon (2009) recruited 8 women with an average age of 22 years and an average
bodymass index (BMI) of 20.2, and randomly assigned them to receive a dietary fiber-
free diet (control), a diet containing 25 g of cellulose, or a diet containing 360 g of
bamboo shoots. Each dietary intervention lasted 6 days and the women underwent each
segment. At the end of the study, the bamboo shoot-rich diet was associated with a 15.7
and 11.8 mg/dL reduction in total cholesterol levels as compared to the control and
cellulose diets, respectively. Moreover, LDL cholesterol was reduced by 16.1 mg/dL
following the bamboo shoot diet compared to the control diet, but there were no
differences between the bamboo group and the cellulose group. High-density lipoprotein
(HDL) cholesterol levels were unchanged by the bamboo diet but increased by 7.2 mg/dL
following consumption of the cellulose diet. Following the bamboo shoot diet, the
average number of bowel movements was 6.2 per day, compared to 4.3 and 5.6 in the
control and cellulose groups, respectively. These results suggested that consumption of
bamboo shoots, which contained high levels of dietary fiber, might help in preventing or
delaying the onset of chronic diseases. Chen et al., (1985) reported a high content of
hemicelluloses in 10 bamboo species that exists in the form of polyxylose. When
hydrolyzed, polyxylose forms xylose that is hydrogenated to produce xylitol, a product
extensively used in the food and chemical industries. Xylose is characterized by a special
flavor and can relieve cough. Bamboo shoots are reported to have anticancer,
antibacterial, and antiviral activity due to the presence of lignans, which is an important
component of fiber (Shi and Yang, 1992; Akao et al., 2004). Lignans are important
components found in cereals and other cruciferous vegetables. Although fiber itself is
invisible in food products, it is becoming one of the most appreciated ingredients in
today’s diet. In 2007, consumers ranked fiber number 5 among the top 10 functional
foods (Sloan, 2008).
2.4.3 Carbohydrates

The carbohydrate content ranges from 2.0 to 9.94% in the raw bamboo shoots was
investigated by different researchers. The carbohydrate content as reported by Kumbhare
and Bhargava (2007) was 3.3, 3.4, 2.6 and 2.9% in B. nutans, B. vulgaris, D. strictus and
D. asper, respectively. The content was found to increase after boiling, ranging from 3.1
to 5.1%, which may be due to the fibrous nature of bamboo. During processing, the
polysaccharides may be hydrolyzed into simple sugars resulting in the formation of
monosaccharides. The oligosaccharides such as stachyose and raffinose, which cause
flatulence, can be converted to monosaccharides and thus the cooking of bamboo shoots
could be beneficial (Kumbhare and Bhargava, 2007). Similarly, in another study the
carbohydrate content ranged from 4.09 to 6.91% in 14 species of freshly emerged
juvenile bamboo shoots. There was an increase in the CHO content after boiling and also
after fermentation and canning up to 72% as compared to fresh shoots (Nirmala et al.,
2008; Nirmala et al., 2011). Bamboo shoots are being suggested in diet as one of the
carbohydrate source in nutritional therapy for Type I Diabetes in Malaysia (Yap, 2015).

2.4.4 Fats

Bamboo shoots are known for low fat content and contain important essential fatty acids.
Canned shoots were reported to contain the lowest content of crude fat, i.e., 0.25% as
compared to the raw shoots of different species. As shown by Nirmala et al., (2007), fat
content increased upon keeping the shoots for 10 days by almost threefold in three
species (B. bambos, B. tulda and D. asper), and in one species, D. giganteus, the increase
was exceptionally high by six fold. The fat content in the juvenile shoots ranged from
2.64 to 3.97%, but for the 10-day-old shoots, the content ranged from 8.2 to 13.84%
(Nirmala et al., 2007; Nirmala et al., 2008; Nirmala et al., 2011). Variation in the
distribution of lipids between different portions of the shoot has also been reported. Total
lipids (TL) ranged from 800 (top) to 380 mg (base) per 100 g fresh weight and the ratio
of nonpolar lipids (NPL):glycolipids (GL):phospholipids (PL) was about 17:27:56. The
main fatty acids were palmitic, linoleic, and linolenic acids, but composition was
remarkably different among different sections of bamboo shoots (Kozukue and Kozukue,
1981).
2.4.5 Minerals

Juvenile bamboo shoots have a high content of minerals such as K, P, Na, Mg, Ca, and
Fe. The shoots are labeled as a heart-protective vegetable because of its high content of K
that helps to maintain normal blood pressure and a steady heart beat. Potassium is a
mineral involved in electrical and cellular body functions. In the body, potassium is
classified as an electrolyte. As the heart is a large muscle that is continually, rhythmically
contracting, potassium is extremely important to proper heart function. The K content in
bamboo shoots ranges from 232 to 576 mg/100 g fresh weight. Common vegetables like
Brassica oleraceavar. Botrytis (Cauliflower), Brassica oleraceavar. capitata (Cabbage),
Cucurbita maxima (Pumpkin), Phaseolus vulgaris(French bean) are also good sources of
Potassium (Nirmala et al., 2011). The content did not decrease significantly in five
different species (Bambusa bambos, Bambusa tulda, Dendrocalamus asper,
Dendrocalamus giganteus and Dendrocalamus hamiltonii) were upon keeping except in
D. hamiltonii where the content reduced to half in 10-day-old shoots (Nirmala et al.,
2007). Potassium content examined in fresh bamboo shoots (533 mg/100 g) decreased to
450 mg/100 g on boiling for 5–10 min. The content further decreased to 300 mg/100 g
when the shoots were soaked for 2 h and boiled at 50°C for 5–10 min (Tsaltas, 1969).
People prone to high blood pressure are often advised to increase K intake and decrease
Na consumption. The daily recommended intake of K is 2.0 to 5.5 g/d (Belitz and
Grosch, 1999). Hence consuming shoots regularly may be a good way to maintain normal
blood pressure and a healthy heart.

Sodium is essential to maintain the Na–K pump and osmotic balance within the cells. Not
many studies are available on the sodium content in raw bamboo shoots. The content
was found to decrease drastically, i.e., three to fourfold in 10-day-old shoots as compared
to the raw in all the five species tested (Nirmala et al., 2007). However, the content
remained almost the same in 10-day-old shoots (3.64 mg), fermented (3.62 mg), and
canned shoots (3.24 mg) for D. giganteus species (Nirmala et al., 2008). Amongst all the
species, mixture of leaves and shoots of Bambusa arundinacea was found to be the
richest source of sodium (Bhargava et al., 1996).
Calcium is an important mineral required for the growth of bones. Calcium content in
shoots in some early reports varied from 0.01 to 0.03% (Ferreira, 1992; Duke and
Atchley, 1986). Recently, 30–400 mg/100 g calcium was reported for bamboo
shoots(Kumbhare, 2003). The calcium content in the mixture of leaves and shoots was in
the range of 320–560 mg/100 g for different species (Bhargava et al., 1996). High values
of calcium ranging from 1,200 to 1,900 mg/100 g were also estimated in species like
B.balcooa, M.baccifera, D.hamiltonii commonly found in North-eastern regions (Bhatt et
al., 2003; Bhatt et al., 2005b). The change in calcium content upon keeping the shoots
was examined and reported that the calcium content decreased in 10-day-old shoots.
Drastic reduction took place in D. giganteus from 6.80 to 1.48 mg/100 g (Nirmala et al.,
2007). In a recent study, values ranging from 21.17 to 180.69 mg/100g, highest being in
B. polymorpha and lowest in B. pallida have been reported (NMBA, 2009).

Phosphorus is another macro element that is required along with calcium for the growth
and maintenance of bones and teeth. The work done during the early 1980s reported a
higher value ranging from 40 to 65 mg/100 g (Yamaguchi, 1983; Gopalan et al., 1978;
Duke and Atchley, 1986). Another report showed values ranging from 150 to 1,000
mg/100 g (Bhatt et al., 2003; Bhargava et al., 1996; Bhatt et al., 2005b). Processed and
dried shoots have shown to possess a very high phosphorous content, i.e., 1,049 mg/100 g
(Rajyalakshmi and Geervani, 1994). Significant reduction in the phosphorous content
upon storage was not observed except in D. asper where the content decreased from
40.95 to 29.08 mg/100 g. (Nirmala et al., 2008).

Magnesium, although a macroelement, is yet an overlooked mineral in case of bamboo


shoots. It is required for contraction and relaxation of muscles and for the functioning of
certain enzymes in the body. The content ranged from 5.38 to 140 mg/100 g according to
various reports. (Bhatt et al., 2003; Bhargava et al., 1996; Bhatt et al., 2005b; Nirmala et
al., 2007). The content did not decrease significantly upon keeping, fermenting, and
canning (Nirmala et al., 2007).

Many essential microelements like cobalt, copper, cadmium, lead, nickel, selenium, iron,
and zinc have been determined in different bamboo species (Nirmala et al., 2007).
Among these, iron holds the more prominence followed by zinc and selenium as far as
nutritional importance of these elements is concerned.

Iron, though not a macroelement, is an important element for the formation of RBCs and
prevents diseases like anemia. Not many researchers have tried to estimate the iron
content in bamboo shoots. The iron content ranged from 0.1 to 3.37 mg/100 g in different
species (Yamaguchi, 1983; NMBA, 2009; Nirmala et al., 2007; Gopalan et al., 1978). A
higher value (22 mg/100 g) was reported in processed and dried shoots (Rajyalakshmi
and Geervani, 1994). The iron content was found to reduce in 10-day-old shoots in all the
species (Nirmala et al., 2007).

Selenium is essential for normal growth, fertility and for the prevention of a wide variety
of diseases in animals. Selenium is a constituent of the enzyme glutathione peroxidase
(GPx), a selenoprotein. This enzyme is the protective agent against the accumulation of
H2O2 (hydrogen peroxide), and organic peroxides within cells. It is involved in immune
mechanisms, ubiquinone synthesis, and mitochondrial ATP biosynthesis. RDA of
Selenium is around 55 mcg per kg body weight of an adult (Dietary Guidelines for
Indians, 2011). Selenium has close metabolic relationship with vitamin E for curing
certain diseases (Anon, 2009). Selenium was found in trace amounts (0.0003 mg/100 g)
in the juvenile shoots (Nirmala et al., 2007).

Approximately 15 mg of zinc is required by an adult in a day. Like iron, zinc is absorbed


according to the body’s needs. The mineral is required for normal growth and sexual
maturation, as part of an enzyme that transfers carbon dioxide from tissues to the lungs,
the production of insulin by the pancreas, etc. The content of zinc ranged from 0.57 to
1.01 mg/100 g shoots in five species of shoots (Nirmala et al., 2007).

2.4.6 Vitamins

Not many studies have reported the vitamin content in fresh shoots, although bamboo
shoots form an important delicacy in many Southeast Asian countries. Yamaguchi (1983)
reported the vitamin A (20 IU), B1 (0.15 mg/100 g), B2 (0.07 mg/100 g), B3 (0.60
mg/100 g), and C content in bamboo shoots in general. Rajyalakshmi and Geervani
(1994) reported that thiamine (0.05%), riboflavin (0.01%), and niacin (0.03%) contents in
the processed and dried shoots were found to be lower as compared to raw shoots. The
content of nicotinic acid ranged from 3.87 mg to 14.92 mg/100 g in the shoots of four
species of bamboo, highest being observed in B. nutans (Kumbhare, 2003). Earlier work
reported lower nicotinic acid content ranging from 0.20 mg to 0.80 mg/100 g in bamboo
shoots (Duke and Atchley, 1986). The amount of pyridoxine in raw bamboo shoots
ranged from 0.53 mg to 1.70 mg/100 g (Kumbhare, 2003). Edible shoots have been
believed to contain considerable amounts of vitamin B12. However, it turned out that
they do not contain appreciable amounts of vitamin B12; but, certain compounds
showing vitamin B12-like activity (known as the alkali-resistant factor) were found in
shoots (Watanabe, 2007). Many researchers have focused on estimating ascorbic acid,
one of the important water-soluble vitamins and antioxidant. Vitamin C content was
found to be as high as 23 mg/100g in a few species (Bhargava et al., 1996) as compared
to 4 mg/100g, which was reported much earlier than that (Yamaguchi, 1983; Duke and
Atchley, 1986). This kind of wide variation could be because of the mixture of shoots and
leaves taken in the first study. In another report, vitamin C content for a number of
bamboo species ranged from 3.0 to 12.9 mg/100g, highest being in D. hamiltonii and
lowest being in D. sikkimensis (Bhatt et al., 2003, Bhatt et al., 2005b).

The ascorbic acid content has been reported to increase upon fermentation of bamboo
shoots into a product called Soibum, commonly consumed in North-east India, from
0.46% (raw) to 0.55% (5th day fermented shoot) (Devi and Singh, 1986). On the other
hand, the ascorbic acid content decreased upon storage from the day of harvesting
(Nirmala et al., 2008). Juvenile shoots of B. bamboos, B. tulda, D. asper, D. giganteus
and D. hamiltonii contained 1.90, 1.42, 3.2, 3.28 and 2.45 mg of vitamin C/100 g,
respectively, as compared to 10-day-old shoots containing 1.31, 1.00, 2.12, 2.15 and 1.79
mg/100 g shoots, respectively. Further reduction in the content was observed in the
canned shoots (1.8%) of D. giganteus followed by fermented shoots (1.09%) (Nirmala et
al., 2008). Only one researcher (Yamaguchi, 1983) has determined Vitamin A content
(20 IU), which is much less as compared to other vegetables. Vitamin E has been well
known for its antioxidative properties, which play a role in preventing metabolic
disorders by acting as a free radical scavenger. Very few studies involved in the
quantitative analysis of this vitamin in bamboo shoots have been reported. Vitamin E
content ranged from 0.61 to 0.91% and kept on reducing with older and fermented
shoots. The content in juvenile shoots for B. bamboos, B. tulda, D. asper, D. giganteus
and D. hamiltonii was found to be 0.61, 0.61, 0.91, 0.69 and 0.71, respectively, but it
decreased to 0.24, 0.24, 0.42, 0.24 and 0.31, respectively, in 10-day-old shoots (Nirmala
et al., 2007). In a recent study, compounds of nutritional importance like α-tocopherol
(0.26 mg/100 g),c-tocopherol (0.42 mg/100 g), β-carotene (1.9 lg/100 g), and lutein
(35.6lg/100 g) have been reported in raw bamboo shoots (Kim et al., 2007).

2.4.7 Bioactive compounds

Bioactive compounds are essential and nonessential compounds that occur in nature, are
part of the food chain, and benefit human health (Kris Etherton et al., 2002; Biesalski et
al., 2009). Plant food contains many bioactive compounds in addition to those that are
traditionally considered nutrients. These physiologically active compounds, referred to as
“phytochemicals,” are produced by way of secondary metabolism in relatively small
amounts (Rodriguez et al.,2006).

Bamboo shoots, both fresh and fermented, are a good source of phytosterols that are the
precursors to many pharmaceutically active steroids found in plants (Srivastava, 1990;
Sarangthem and Singh, 2003a) and act as nutraceuticals (Miettinen, 2003). An antitumor
agent has been prepared from moso bamboo (P. pubescens) that has an antitumor effect
(Hiromichi, 2007). Rich sources of phytosterols include grain legumes such as sesame,
chickpeas, lentils, and peas; cereal grains such as wheat, corn, millet, rye, and barley
(Ryan et al., 2007); vegetable oils including corn oil; and nuts such as pecans, pinenuts,
pistachio nuts, peanuts, cashew nuts, and almonds (Ryan et al., 2007), and bamboo
shoots (He and Lachance, 1998; Lachance and He, 1998; Sarangthem and Singh, 2003c;
Lu et al., 2010). Predominant sterols in bamboo shoots have been identified as β-
sitosterol, campesterol, and stigmasterol (He and Lachance, 1998; Lachance and He,
1998). The phytosterol content and composition in shoots of 4 bamboo species,
Pleioblastus amarus, P pubescens, D. latiflorus, and P. praecox, were evaluated using
ultra-performance liquid chromatography with atmospheric pressure chemical ionization
mass spectrometry (MS) in order to facilitate dietary recommendations and
comprehensive utilization of the shoots (Lu et al., 2009). The major sterols present were
β-sitosterol (24.6%), campesterol (2.2%), stigmasterol (1.2%), er-gosterol (0.2%),
cholesterol (0.6%), and stigmastanol (<0.1%); to-tal phytosterol content was 28.7% (Lu
et al., 2010). Bamboo shoot oil (BSO) is a phytosterols-rich extract from the bamboo (P.
pubescens) shoots obtained by supercritical carbon dioxide extraction and its protective
effects and its mechanism on nonbacterial prostatitis (NBP) were investigated by Lu et
al., (2011). The level of total phytosterols in bamboo shoots ranges from 0.12% to 0.19%
on a dry weight basis in different species of bamboos (Sarangthem and Singh, 2003a).
The concentration of total phytosterols from the initial stage of fermentation (0 d) up to
35 d increases, but later on more or less of a decline in the level of the phytosterol
contents occurs. The increase in concentration in fermented shoots may be almost up to
double as in B. balcooa (increases from 0.18% to 0.61% dry weight) and in D.
hamiltonii(increases from 0.19% to 0.44% dry weight) (Sarangthem and Singh, 2003a,
2003b). The increase in the fermented samples is due to anaerobic digestion by
microorganisms that caused degradation of organic matter and resulted in the enrichment
of phytosterols (Stahl, 1969).

Phytosterols are precursors of many pharmaceutically important steroidal products such


as corticosteroids, oral contraceptives, anti-inflammatory agents, synthetic anabolic
steroids, and estrogenic hormones. Thus, succulent bamboo shoots, which are easily
available in large quantities, can be used as a source of phytosterols. Plant phenols are
bioactive compounds of interest present in all plants because they are multifunctional and
can act as free radical terminators, metal chelators, and singlet oxygen quenchers (Kris
Etherton et al., 2002). Phenolic compounds, commonly referred to as polyphenols, are
secondary metabolites and their distribution is almost ubiquitous (Pereira et al., 2009).
Most research studies have investigated the functional activities of bamboo leaves and
stems. It has been determined that the antioxidant capacity of bamboo leaves is due to
their high polyphenol content (Lu et al., 2006). However, only a few studies have
reported on the functional properties of bamboo shoots. Wang and Ng (2003) reported the
isolation of an antifungal protein, Dendrocin, isolated from the bamboo Dendrocalamus
latiflora. Bamboo shoots can be a good dietary source of natural phenolic oxidants and
dose-dependent inhibitory activity on angiotensin-converting enzyme as determined in 2
species of Phyllostachys pubescens and P. nigra (Park and Jhon, 2010). Antioxidants and
antimicrobial compounds have also been isolated from P. heterocycla. (Quitain et
al.,2004). Phenolic acids present in the tender shoots have mild anti-inflammatory
properties and are potent antioxidants that prevent cancer and blood vessel injury that can
start atherosclerosis (Belitz and Grosch, 1999). Kim, Y.N. et al., (2007) estimated the
tocopherol and carotenoid contents of Korean vegetables including bamboos. In bamboo
shoots, they reported α-tocopherol and γ-tocopherol contents to be 0.26 mg/100 g and
0.42 mg/100 g, respectively.

2.5 Health benefits of Bamboo shoots

The main nutrients in bamboo shoots are protein, carbohydrates, amino acids, minerals,
fat, sugar, fiber, and inorganic salts. As a dietary fiber source, the shoots have beneficial
effects on lipid profile and bowel function. An increase in dietary fiber from shoot
consumption may also be useful in the management of hypertension and obesity through
its effect on energy density of food and nutrient bioavailability (Anderson and Strong,
1983; George et al. 1982). People prone to high blood pressure are often advised to
increase K intake and decrease sodium consumption. Bamboo shoots fill that need
completely. The shoots are also reported to have anticancer, antibacterial, and antiviral
activity due to the presence of lignans, which is an important component of fiber (Shi and
Yang, 1992; Akao et al., 2004). Its also claimed that eating bamboo shoots imparts a
freshness to the skin and makes the skin fairer and smooth (Shi and Yang 1992). The
survey conducted by Kalita and Dutta (2012) showed that some ethnic tribes of Northeast
India used bamboo shoots to control high blood pressure and cardiovascular ailments.

In South Asian countries, bamboos have been utilized for traditional medicine treatments
to relieve hypertension, sweating, and paralysis. However, little scientific evidence has
supported such claims until now. Modern research has revealed that bamboo shoots have
a number of health benefits, from cancer prevention and weight loss to lowering
cholesterol level, improving appetite and digestion. It is also low in sugar and therefore
can be used by persons on sugar-restricted diets. Bamboo shoots suggested in diet as a
part of carbohydrate source in nutritional therapy for diabetes in Malaysia (Yap, 2015).
The shoots also contain anticarcinogenic agents, and making them a regular part of a diet
effectively reduces the free radicals that can produce harmful carcinogens. There are
instances of using locally available bamboo shoots by Karbi tribes in Karbi Anglong
autonomous region of Assam state in India to control early stage of cancer (Kalita and
Dutta,2012). The anticancer property of bamboo shoots might be attributed to the
presence of lignans and phytosterols.The production of carcinogens, growth of cancer
cells, cell invasion, andmetastasis are inhibited by phytosterol (Meric et al., 2006). It is
believed that bamboo extract may have antioxidant activities and provide
antiinflammatory effects (Hu et al., 2000; Lu et al., 2006). Furthermore, bamboo-derived
pyrolysates have been proposed to have antimicrobial and antifungal activities (Fujimura
et al.,2005) and to protect neurons from oxidative stress (Akao et al., 2004). Hong et al.,
(2010) studied the effects of pyrolysates derived from 3 bamboo species, Phyllostachys
bambusoides(Siebold et Zucc.), P. nigra ([Lodd. Ex Lindl.] Munro), and P. pubescens
(Mazel ex J. Houz.), and indicated that pyrolysates may have antiapoptotic effects and
can be useful as a supplement for ischemic injury treatment. Several antimicrobials and
antioxidants have been isolated by supercritical CO2 and subsequent hydrothermal
treatment of the residues from moso bamboo, including an ethoxyquin, a sesquiterpene,
and a cyclohexanone derivative (Quitain et al.,2004). Bamboo shoots are also reported to
have anticancer, antibacterial, and antiviral activity due to the presence of lignans that are
an important component of fiber (Shi and Yang, 1992; Fujimura et al.,2005). Its
relatively high content of even up to 4% cellulose increases the peristaltic movement of
the intestines and helps digestion. It also prevents constipation and decreases body fat.

Composition of nutrients like carbohydrates, proteins, vitamins and dietary fibers may
vary considerably among different species and also on agroclimatic conditions. Thus it
becomes imperative to understand the nutrient composition of a particular bamboo
species growing in a particular region to exploit its edible potential. A study by Nirmala
et al., (2011) on 14 species of edible bamboo shows that the carbohydrate (CHO) content
is also found to vary between bamboo species with highest in B.tulda and lowest in
D.asper and D.brandisii. A comparitive analysis of two species with common vegetables
shows that both the bamboo species are rich in proteins and also the protein content is
much higher than the common vegetables. Apart from proteins and CHO, some species
(B.vulgaris, D.hamiltonii) were found to have high concentration of vitamin C and
vitamin E also. Shoots also contain fairly high amounts of minerals like iron, manganese
and zinc as compared to some commonly used vegetables like Brassica oleraceae var.
Botrytis (Cauliflower), Solanum tuberosum (Potato), Abelmoschus esculantus (Ladies
finger) and Solanum melongena (Brinjal). Selenium, another very important mineral
known for antioxidant properties, is also present in bamboo shoots unlike in other
vegetables. The Recommended Dietary Allowance of an adult person per day for
protein, dietary fibre and minerals is more or less met from around 100gms of bamboo
shoots. Hence, daily consumption of shoots can be expected to be beneficial to the human
body in so many ways besides helping in balanced nutrition.

2.6 Shoot processing

2.6.1. Significance of shoots processing

Bamboo shoots contain very high concentration of cyanogenic glucosides, which on


endogenic hydrolysis, yield hydrocyanic acid lending a bitter taste to the bamboo shoots.
Several studies have reported that bamboo shoots contain cynogenic glycoside, which can
prove hazardous to health if not processed properly and if consumed in excess.
Taxiphyllin (4-hydroxy-(R)-mandelonitrile-b-D-glucopyranoside), a cynogenic glycoside
(Conn, 1969) has been found to be the potential toxic component present in the different
species of bamboo shoots like Dendrocalamus latiflorus, D. giganteus, D. hamiltonii,
Bambusa vulgaris and B. guadua (Schwarzmaieer, 1976, 1977; Vetter, 2000). Cooking
largely destroys the enzymes responsible for the endogenic hydrolysis.

Another constituent of bamboo shoot, homogentistic acid, is responsible for the


disagreeable, pungent taste, characteristic of bamboo shoot (Ferreira et al. 1995).
Cyanogenic glycosides which are phytotoxins, occur in at least 2000 plant species, of
which a number of species are important food items in many parts of the world. Cassava
(Mannihot esculenta) and sorghum (Sorghum vulgare), important staple food in many
parts of India, also contain cyanogenic glycosides. There are at least 2650 species of
plants that produce cyanogenic glycosides. Unlike linamarin and lotaustralin which are
the cyanogenic glycosides found in cassava plants, taxiphyllin in bamboo shoots is highly
unstable and is easily decomposed when treated with boiling water (Haque and
Bradbury,2002). These plants also possess a corresponding hemolytic enzyme (beta-
glycosidase), which combine together when the cell structure of the plant is disrupted by
a predator or as a result of physical damage. This compound subsequently breaks down
to a sugar and a cyanohydrin, and then rapidly decomposes to hydrogencyanide (HCN)
and an aldehyde or a ketone. This combination of cyanogenic glycosides and hydrolytic
enzymes is primary deterrent or bioshield against predators. HCN thus formed, if present
in very high concentrations is toxic. A fairly high concentration of HCN is found in
juvenile bamboo shoots which could be mainly to deter predation.

The cyanide (HCN) content varies in different parts of a plant and also between the same
parts of different portions of the same species (Jones, 1998). The HCN content varies
from species to species and also within the juvenile shoot. The major cyanogenic
glucoside found in edible part of bamboo shoots is taxiphyllin. This compound is unusual
amongst the 60 or so known similar compounds in that it degrades readily in boiling
water. Any normal cooking of bamboo shoots should be able to remove this problem
though. Consumption of improperly prepared or unprocessed bamboo shoots may
produce symptoms like rapid respiration, drop in blood pressure, dizziness, stomach
pains, headache, vomiting convulsion, and coma (FSANZ, 2004). Hydrogen cyanide
interferes in proper functioning of cytochrome oxidase inhibiting normal cellular
respiration. According to FSANZ report, the HCN present in concentration of 0.5–
3.5mg/Kg body weight is considered as lethal which may cause serious health damage
leading to death. However, FAO/WHO Codex Alimentarius has defined a safe limit for
human consumption, which is 10 mg HCN equivalent per kg dry weight
(FAO/WHO,1991). HCN is a respiratory poison which when present in high
concentration may lead to serious health predicament to individual concerned. Pramod
Kumar et al., (2011) reported a case of cyanide poisoning in JSS Hospital,
Mysore,India,when a 14-year-old female patient was brought to the Hospital in
unconscious state with a history of consumption of bamboo shoot extract (juice). The
patient showed the symptoms of convulsion, vomiting, respiratory distress, and loss of
consciousness but regained consciousness two days after proper medical treatment. This
case might be an isolated one as there have not been any reports of extreme cyanide
poisoning because of bamboo shoot consumption as fresh or fermented shoots are
presoaked and boiled in water during the preparation of most bamboo based ethnic
dishes. Haque and Bradbury (2002) observed total cyanide content in shoots of Bambusa
arundinacea (1010ppm) to be much higher than other plants like flax seedmeal
(390ppm), cassava root (27ppm), sorghum leaf (750ppm), apricot stone (785ppm), and
giant taro leaf (29ppm).

Bamboo shoots need to be processed before consumption to remove bitterness.


Communities which traditionally have been using shoots may have evolved their own
local techniques for doing so. Processing of bamboo shoots after harvest varies from
region to region depending on culture and tradition. Traditionally in rural areas of
peninsular India, the shoots are processed by soaking in water for more than 24 hours
before cooking. In North-east India, processing is slightly different and shoots are
generally allowed to ferment before consumption. However different processing methods
have varied impact on the nutrient content of the edible shoots(Choudhury et al,. 2011).
A study on D.giganteus, a commonly used species in the sub-Himalayan and North-
eastern parts of India, and also in Nepal and Bhutan in the fermented form, suggests that
fermentation considerably reduces the content of various nutrients like carbohydrates,
proteins, vitamins and also minerals like sodium when compared to nutrients in the fresh
shoots. It is possible that some other processing methods like boiling, steaming, treatment
with common salt can also reduce the HCN content in the bamboo shoots (Ferreira et al.,
1995; Bindu et al., 2010; Sopade, 2000). The impact of these treatments on the nutrient
composition needs to be worked on to standardise an effective technique since there is
not much published literature available on this aspect.

2.6.2 Shoot processing methods

Fermentation, a chemical change produced through the break-down of carbohydrates and


proteins by yeast, bacteria, or molds, is a process used for centuries in order to make and
preserve certain types of food. Not only is fermentation a process to preserve food, it also
adds to the nutrient value and enhances flavor. Fermentation of bamboo shoots involves
several steps. Sheaths from the freshly harvested shoots are peeled off, inner creamy
white portion cleaned in water, and cut into small slices . Earthen pots or baskets made of
bamboo culms slivers are used as containers for fermentation. When bamboo baskets are
used, the inner surface is layered with banana leaves or perforated plastic sheet to drain
off the liquid during the fermentation process. The chopped bamboo shoots are put into
the container tightly by pressing and are covered with banana leaves or polythene. The
shoots are subjected to heavy weights like stones or wooden logs to keep them under
pressure. If using pots, sun-dried pots are filled tightly with shoots up to the neck and
covered; the bottom is perforated for draining out the acidic fermented juice produced
during fermentation. The shoots are kept in this condition for 6 to 12 months. Major
species used for fermentation in North-east India include Bambusa tulda, B. vulgaris, B
balcooa, B. pallida, Dendrocalamus hamiltonii, D. hookeri, D. flagellifer, D.
giganteus,and Melocanna baccifera (Giri and Janmejoy, 2000; Bhatt et al.,2005b;
Jeyaram et al., 2009). No preservative is used and after complete fermentation, the shoots
can be stored 1 to 2 years before marketing, thus making them available throughout the
seasons. The microorganisms involved in fermentation of bamboo shoots are Bacillus
subtilis, B. licheniformis, B. coagulans,and Micrococcus luteus (Sarangthem and Singh
2003a). It has been shown that the fermented succulent shoots of B. tulda and D.
giganteus are an enriched source of phytosterols and can, therefore, be used as starting
material in the production of steroidal drugs (Srivastava, 1990; Sarangthem and Singh,
2003a).

Traditionally, various fermented bamboo shoot products are consumed in the world since
ancient times. Ethnic people living in sub Himalayan regions, Nepal and Bhutan prepare
and consume a variety of domesticated and wild bamboo shoots and their fermented
products (Sharma, 1989; Tamang and Tamang, 2009; Tamang et al.,1988; Tamang, 2000;
Vaiphei, 2005). A traditional fermented bamboo shoot product of the eastern hills of
Nepal and Bhutan is mesu (Tamang and Sarkar, 1996). Use of mesuas, a pickle and as a
base in curries is a conventional dish among the Nepalis, Bhutias and the Lepchas of the
Darjeeling hills and Sikkim (Tamang, 2005). Delicacies in Indian cuisines include ushoi,
soidon, soibum, soijim, iromba, ekung, eup, hiring, lung-siej,tuairoi, tuaithur and syrwa
(Giri and Janmejay, 1987; Tamang, 2000, 2009;Mao and Odyuo, 2007; Agrahar-
Murungkar and Subbulaskhmi, 2006; Jeyaram et al., 2009; Chakrabarty et al., 2014).
Ushoi, a fresh bamboo shoot, is one of the popular food products among the Manipuris
and Apa Tanis of Arunachal Pradesh. Soibum, a fermented bamboo shoot, is an
exceptional delicacy of the Meities of Manipur, eaten as pickle and curry mixed with
fermented fish. Similar fermented bamboo shoot product called naw-mai-dong or nor-
mai-dorngis consumed in Thailand (Phithakpol et al.,1995). Soidonis, another fermented
bamboo shoot product in Manipur, is prepared from the tip of matured bamboo shoots
and consumed both as a curry and pickle. Soijim is another type of fermented bamboo
shoot product developed by submerged fermentation in Manipur. Iromba is a fermented
or boiled bamboo shoot taken with fish and other vegetables by Khasi tribes in
Meghalaya (Agrahar-Murungkar and Subbulaskhmi,2006). Bhardwaj et al., (2005) also
reported the use of banana leaves by Adi tribe of Arunachal Pradesh for removing toxic
cyanogen during bamboo fermentation. The bamboo shoots were pressed hard under big
stones and kept for 3-4 months to lower down the bitter taste of fermented bamboo
shoots. In central India, the young shoots are grated and fermented to prepare
kardioramil, a sour vegetable soup. In the region, the bamboo shoots are fermented, dried,
ground into powder and used as a garnish called hendua, which is commonly preferred
among the tribal people (Panda and Padhy, 2007; Bal et al.,2009). In Nepal, bamboo
shoots are fermented with turmeric and oil, and cooked with potatoes to prepare an item
called alu tama (Tamang, 2009).

People consume fresh bamboo shoots in various forms. Bamboo shoots can be eaten fresh
after boiling. In Indonesia, bamboo shoots are eaten with thick coconut milk and spices,
which are called gulei rebung; sometimes also mixed with other vegetables, called sayur
lade (Bhatt et al.,2003). Sweet pickles, chutney and candies are prepared from the pith of
the bamboo shoots in the region. The sap of young culms tapped during the rainy season
is simply made into a soft drink in China (Yang et al., 2008). In Manipur, the fresh
bamboo shoots are taken with dry fish (Tamang and Tamang, 2009). The edible bamboo
species in Western Ghats of India are also used as snacks, fried food stuffs, and curries
(Bhatt et al.,2003). Tama, a non-fermentated bamboo shoot curry is very familiar among
the people of Sikkim (Tamang, 2009). In Singapore, people consume bamboo shoots in
canned or frozen form (Pan,1995). The various traditional processing methods and
consumption pattern across India and world is listed in table 2.1.

Cooking of raw shoots is usually done to enhance its palatability. Various cooking
methods may also have different degrees of impact on the nutrient composition and
functional ingredients as well. The actual nutrient value of the cooked bamboo shoots
may differ from that of the fresh raw bamboo shoots. A comparitive study on the impact
of different cooking methods ( boiling, steaming and stir-frying ) indicates that the
nutritive content of bamboo shoots are influenced by these methods. Boiling of bamboo
shoots resulted in the loss of crude proteins, soluble sugars, ash, free amino acids, and L-
ascorbic acid and reduction of the antioxidant capacity. However in steaming, the basic
nutrient components, Total Free Amino Acids (TFAA), and antioxidant capacity of the
bamboo shoots changed very little, but each free amino acid had been modified to varied
extents and the L-ascorbic acid content was considerably reduced. The addition of peanut
oil during stir-frying increased the crude fat content, while both the soluble sugar and
TFAA contents decreased. High temperature and short time stir-frying promoted the
occurrence of the Maillard reaction which gives a unique flavor and generates antioxidant
substances. Also, L-ascorbic acid was preserved at maximum extent as compared to the
other two methods of cooking (Sopade, 2000).

The presence of toxic compound and bitterness in bamboo shoot decreases its consumer
acceptability. Haque and Bradbury (2002) reported that tip of the bamboo shoots had the
highest HCN content (920-1600 mg/kg), followed by the mid portion (620-1140 mg/kg),
and the least HCN content was reported in the basal portion (114-440 mg/kg). Bhatt et
al., (2005b) reported 0.01 to 0.02 per cent HCN in edible bamboo shoot. The nutritional
implications of cyanogens and process to reduce their levels in products to a safe limit
have been extensively studied. Codex Alimentarius Commission Standard (CACS) has
defined the maximum safe limit as maximum of 10 mg HCN equiv. per kg fresh shoot.
Cyanide content, naturally, is reported to decrease substantially following harvesting
(Nirmala et al., 2007). Different indigenous methods of reducing acidity/bitterness from
fresh bamboo shoots has been reported and some of them include chopping of tender
shoots into small pieces, partial drying of fresh shoots, boiling in water/salt water and
draining or keeping shoots in hot water for 10 – 15 min or in water for a week at ambient
temperature, etc. Bhatt et al., (2005b) reported unique traditional processing of bamboo
shoot fermentation to reduce the cyanide percentage. Ferreira et al., (1995) reported the
optimum cooking conditions that resulted in 97% reduction of HCN were 98 - 102°C for
148 - 180 min. Subsequently, Tripathi (1998) mentioned that removal of HCN can be
done by steaming bamboo shoot. Sopade (2000) reported reduction of 40-70 per cent of
cyanogenic glycosides in bamboo shoots as a result of steaming. Steaming is reported to
be as effective as baking in detoxifying shoots but both are less effective than boiling.
Toxic constituent taxiphyllin from shoots can also be removed by several changes of
water during cooking or by presoaking for a long time in several changes of a solution of
2 per cent salt. (Bhargava et al., 1996). Wongsakpairod (2000) reported superheated
steam drying under low temperature removes HCN from bamboo shoot as Taxiphyllin
decomposes at around 116°C. Commercial processing of bamboo shoots involves
washing followed by peeling of the skin, shredding, slicing, and cutting into cones. Next,
they are boiled for about 10 min to remove bitterness. This is then followed by storing in
brine solution with 5% salt and 1% citric acid. Finally, the processed shoots are vacuum
packaged in 100 g packets containing 25 ml brine solution (Nimachow et al.,2010).
However, the tribal people of the northeast have evolved different indigenous techniques
of processing: slicing, shredding, fermenting, pickling, and drying (Nimachow et
al.,2010; Singh et al.,2007).

Even though fresh shoots are far superior in taste and texture, bamboo shoots are
commonly available in the canned form. Canning has been observed to be effective in
abating rancidity and preventing the growth of micro-organisms in bamboo shoots (Fu et
al.,2002).The Japanese were the first to develop and use modern technology for the
processing of fresh shoots. The shoots are cut in various ways into halves, thin sliced,
stripped, diced, or canned as whole after processing. The processing technique to produce
canned bamboo shoot involves crucial steps that must be followed in order to obtain good
quality products. This involves boiling the sliced bamboo shoots in water for about 4 h or
for 40 to 60 min at 120°C, cooling, and then storage in brine solution containing usually
around 5% to 10% NaCl or more, depending on the species used, and 1% citric acid.
Products so processed may be in cans or retort pouches. Cans contain processed fresh
materials, while retort pouches contain processed salted or cured materials. Most tinplate
canned products are used for cooking, and the pouch products are like ready to eat form
(Midmore, 1998). The drawback of canned shoots are poor shape, loss of aroma and
crunchiness, and unknown source of shoots are questionable in quality and require
prompt usage within 3 to 4 days after opening the can lid. In some countries, good
recognized species are cultivated to produce shoots of low fiber, and good texture, color,
and taste. However, except for the dietary fiber, there is an over-all decrease of the
nutrient components, especially vitamins and minerals, during canning (Nirmala et
al.,2008). There is a growing demand for processed and packaged bamboo shoots in the
national and international markets. Shelf life of freshly harvested bamboo shoots is 9 and
23 days in water and brine, respectively (NMBA, 2009). During storage, a bitter taste
develops in the bamboo shoots. This is also noticed if stored for a longer period of time
or exposed to sunlight. It has been reported that bamboo shoots, preserved in plastic bags
have a risk of contamination by the materials present in the plastic bags (Chiangthong
and Chayawat, 2009). Bamboo shoots dried, marinated and sautéed can be converted to
powders or nuggets and used as medicines.

It has been reported that there is a 95.1% decrease in the moisture content of dried
bamboo shoots (Satya et al., 2010). Water content of fresh bamboo shoots is 92.6/100 g
of fresh weight compared to 4.6/100 g in dried bamboo shoots (Muchtadi and Adawiyah,
1996). Cheng (2006) reported a multi-stage vacuum pressure preservation technique
combined with hydro-cooling which could reduce the temperature of bamboo shoots
efficiently. The author suggested that the process has the advantage of minimizing the
energy consumed by the vacuum system. A comparative study of various methods for
drying of bamboo shoot along with their advantages and disadvantages has been
systematically reported by Majumdar (2006) and Satya et al., (2010). Bal et al., (2010)
studied the effect of microwave power on drying kinetics of bamboo shoots. The authors
suggested that by increasing microwave output power, the effective moisture diffusivity
values increased from 4.153×10−10 to 22.835×10−10m2s−1. Production of bamboo being
seasonal with high demand throughout the year, especially in China, Japan, USA,
Canada, Thailand, Nepal, Bhutan, Australia and India, there is a need to develop effective
process technologies to preserve the bamboo shoots in consistent and imperishable forms
to be used during the off-seasons (Choudhury et al.,2011).
Table 2.1: Different processing methods/consumption pattern of bamboo shoots from India and other parts of the world

S.No. Method / Consumption Location Reference


1 Mesu - fermented bamboo shoot product Eastern hills of Nepal and Tamang and Sarkar, 1996; Tamang, 2005
Bhutan, Lepchas of the
Darjeeling hills and Sikkim
2 Ushoi,a fresh bamboo shoot Manipuris and Apa Tanis of Giri and Janmejay, 1987
Arunachal Pradesh
3 Soibum – sliced bamboo shoots Manipur Tamang, 2000
fermented in clay pot or bamboo basket
by Meitei tribes in Manipur
4 Soidonis - fermented bamboo shoot, Manipur Agrahar-Murungkar and Subbulaskhmi, 2006
prepared from the tip of matured bamboo
shoots and consumed both as a curry and
pickle
5 Soijim is another type of fermented Manipur Agrahar-Murungkar and Subbulaskhmi, 2006
bamboo shoot product developed by
submerged fermentation
6 Iromba is a fermented or boiled bamboo Meghalaya Agrahar-Murungkar and Subbulaskhmi, 2006
shoot taken with fish and other
vegetables byKhasi tribes in Meghalaya
S.No. Method / Consumption Location Reference
7 Hikhu is bamboo shoot chopped pieces Arunachal Pradesh Bhardwaj et al., 2005
and wrapped in banana leaves and kept
for 6-8 days by Adi tribes and Apatani
tribes of Arunachal Pradesh for
removing toxic cyanogen
8 Hiring is sliced bamboo shoots kept in Arunachal Pradesh Bisht et al., 2012
bamboo cylinders and covered with
leaves for 7 days by Apatani tribes of
Arunachal Pradesh
9 Hithyi is sliced bamboo shoots sun dried Arunachal Pradesh Bisht et al., 2012
and stored in bamboo basket by Apatani
tribes of Arunachal Pradesh
10 Ekung is sliced bamboo shoots kept in Arunachal Pradesh Bisht et al., 2012
basket and covered with ekkam leaves
for 5-6 days by Adi tribes of Arunachal
Pradesh
11 Edung is bamboo shoots fermented in Arunachal Pradesh Bisht et al., 2012
bamboo cylinder by Adi tribes of
Arunachal Pradesh
S.No. Method / Consumption Location Reference
12 Lung Siej is bamboo shoots fermented in Meghalaya Bisht et al., 2012
plastic/glass bottles by Khasi tribes of
Meghalaya
13 Eishie is bamboo shoots fermented in Nagaland Bisht et al., 2012
plastic/glass bottles by Aao tribes of
Nagaland
14 Mautuai is bamboo shoots fermented in Mizoram Bisht et al., 2012
plastic/glass bottles by Mizo tribes of
Mizoram
15 Khorisa-tenga, Kardi, Amil, Hendua – North-east India Tamang, 2000; Tamang and Tamang, 2009
other fermented bamboo shoots in North-
east India
16 Kardioramil - young shoots are grated Central India Panda and Padhy, 2007
and fermented to prepare a sour
vegetable soup
17 Hendua - bamboo shoots are fermented, Central India Bal et al.,2009
dried, ground into powder and used as a
garnish, which is commonly preferred
among the tribal people
S.No. Method / Consumption Location Reference
18 Fresh bamboo shoots in traditional Western Ghats in Karnataka and http://www.bambootech.org/files/ERG%20report
cooking Konkan region %20on%20shoots.pdf Accessed on 19th January
2015
19 Bamboo shoots as Pickles Western Ghats in Karnataka and http://www.bambootech.org/files/ERG%20report
Konkan region %20on%20shoots.pdf Accessed on 19th January
2015
20 Bamboo shoots in traditional cooking Hilly regions in Gujarat, http://www.bambootech.org/files/ERG%20report
Karnataka, Maharashtra, Kerala %20on%20shoots.pdf Accessed on 19th January
and Tamil Nadu 2015
21 Bamboo shoots as pickles and papads Kashmir http://www.bambootech.org/files/ERG%20report
%20on%20shoots.pdf Accessed on 19th January
2015
22 Alu tama - bamboo shoots are Nepal and Bhutan Sharma, 1989; Tamang and Sarkar, 1996;
fermented with turmeric and oil, and Tamang, 2000;Tamang, 2009; Tamang et al.,
cooked with potatoes 2005
23 Tchang and Mesu – preparations from Nepal and Bhutan Sharma, 1989; Tamang and Sarkar, 1996;
bamboo shoots Tamang, 2000;Tamang, 2009; Tamang et al.,
2005
S.No. Method / Consumption Location Reference
24 Gulei rebung - fresh bamboo shoots are Indonesia Bhatt et al.,2003
eaten with thick coconut milk and
spices
25 Sayur lade - fresh bamboo shoots Indonesia Bhatt et al.,2003
sometimes also mixed with other
vegetables
26 Canned or frozen bamboo shoots Singapore Pan,1995
27 Naw-mai-dong or nor-mai-dorngis - Thailand Dhavises, 1972; Phithakpol et al.,1995
fermented bamboo shoot
28 Labong, Ginataang Labong, Philippines Phithakpol et al.,1995
Dinengdeng na Labong– preparations
from fermented bamboo shoots
29 Bamboo shoots used as vegetables in Vietnam Jeyaram et al., 2005
stir-fry dishes
30 Canned bamboo shoots China Angau et al., 2003
2.7 Cultivation and management of bamboo for shoot production

2.7.1 Cultivation of Bamboo

Different bamboo species have different requirements for humidity and temperature.
Normally, bamboo thrives at temperature range of 8.8 to 36°C, but some species can even
grow in cold climate with temperature of about -20 °C. Rainfall plays a very important
and dominating role in the distribution and growth of different species. Bamboo prefers
regions of high rainfall ranging from about 1270 mm to about 6350 mm or even more
(Ha, 2010; Ram et al.,2010; Song, et al.,2011). Precipitation affects distribution and
limits growth of bamboo more than any other component of climate, except temperature.
The maximal annual evapotranspiration rate of bamboo has been estimated to be about
3300 mm (equivalent to 33 million per ha-1). In India, 61% of all bamboo species can be
found in areas with 1,500-4,000 mm rainfall per year (Biswas, 1988).

The growth pattern in bamboo is known as exponential, i.e., each year a fresh set of
vegetative shoots originate from the base of the clump soon after monsoon showers. An
emerging culm reaches its full height within 2-4 months (McClure, 1966; Ueda, 1960).
This is achieved by elongation of internode in a telescopic manner and increase in
diameter. Subsequent hardening occurs through lignification (McClure, 1966). Growth of
the new shoots in bamboo clumps occurs as a result of transfer of energy accumulated in
culms through photosynthesis in the previous year (Magel et al.,2005). Every year in a
clump only 10%-40% of the shoots grow into mature bamboo culms while 60%-90%
abort or remain small. The higher the yield of a bamboo clump, the more dead and small
shoots it will tend to have. It is impossible to wipe out all dead shoots; it is natural for the
clump to have big and small, strong and weak shoots every year. In production, therefore,
strong shoots require to be preserved for propagation but the thin and small shoots dug
out for food (Yang and Hui, 2010). Sympodial bamboos produce shoots from May to
October with most production in July to August. Edible shoots should be harvested
before they become fibrous and tough. Any delay will result in loss of quality and
quantity. Generally, the emerging initial shoots and most of those produced in the
summer are harvested, but those produced towards the end of the shooting period will be
selected and retained as mother culms. (Troya et al., 2014). In Japan, bamboo shoots are
forced to grow under mulch heated with electric cables placed 6–8 cm below the soil.
Mulched bamboo shoots can be harvested earlier than those growing in the natural
environmental conditions (Yamaguchi, 1983)

Despite a wide and divergent range of management objectives and environmental


conditions across the tropics, a remarkably small number of genera dominate tropical
plantations as well as agroforestry projects and trials tend to be dominated by very few
bamboo genera. Nonetheless, little is known about their suitability and management in
the context of bamboo based agroforestry. The use of a greater variety of species in
reforestation may improve the resilience of ecosystems, decrease sensitivity to pest and
diseases, and increase functional diversity (Benayas et al., 2009; Rodrigues et al., 2009).
From a socio-economic perspective, increasing species choice may preserve a range of
management options, enhance adaptability to changing market opportunities, and provide
a wider variety of products and services (Aronson et al., 2007). Introducing new species,
however, is not without risks. Many reforestation projects fail due to inappropriate
species choice, a consequence of inadequate knowledge about the potential of species and
their growth and survival rates under different site conditions (Rodrigues et al., 2009).
The importance of testing large numbers of species in screening trials is paramount and
has long been recognized. Yet, few large trials have actually been established across the
tropics (Michiel et al., 2011; Shono et al., 2007; Wishnie et al., 2007).

2.7.2 Management schedule

Several studies have systematically tested the benefits of nutrient application during
different times of the year on bamboo productivity; however, no single, most important
stage was identified. Nutrient application earlier in the season, i.e., shortly before and
during the shoot season, was found to be most appropriate time of fertilizer application
for Phyllostachys nidularia in China and Phyllostachys pubescens in Korea (Shen et
al.,1993; Jeong et al.,1995). Hong (1994) concluded that nutrient demand of
Phyllostachys pubescens cultivated in China is greatest during periods of rapid growth,
i.e., of shoots and rhizomes. Many researchers have noted the generally positive effects
of greater availability of water on bamboo production and how lack of water limits its
productivity. Lin (1995) reported that irrigation promotes shoot and culm growth when
rainfall is insufficient in Dendrocalamopsis oldhamii. Koyama and Uchimura (1995)
highlighted the high water requirement for culm elongation in Phyllostachys
bambusoides, and Huang et al., (1993) showed that in locations with little available soil
moisture, biomass accumulation is largely partitioned below ground in P.pubescens in
China. It is known that wet summers increase shoot production of sympodial bamboo
(Pearson et al.,1994), and that bamboo is not a suitable commercial species for areas
where sufficient water is usually not available (Siddiqui, 1994). Introduction of irrigation
could, therefore, improve production of bamboo products in areas without adequate
amount and distribution of rainfall.

However, only very few studies have attempted to quantify water usage of bamboo under
field conditions or to systematically test irrigation rates. Kleinhenz and Midmore (2000)
estimated the maximal annual water dissipation rate of bamboo to be about 3,300 mm
rainfall equivalent. Midmore et al., (1998) reported that successful cultivation of P.
pubescens in semiarid parts of Australia depends primarily on frequent but non-excessive
additions of water through precipitation and irrigation (>2,000 mm per year). Not only
general demand for high water availability but also availability of water during specific
growth stages may affect bamboo productivity. The detrimental effects of drought but
positive effects of irrigation during the shoot and culm growth phase for species
commonly cultivated in China were outlined by Chu and Xu (1988), Li and Zhang
(1987), and Lin (1995). Fu and Banik (1995) reported that irrigation was required for
intensively managed shoot stands after 10 days without rainfall during the shoot season
for P.pubescens. Midmore et al., (1998) stressed the need to pay attention to water supply
just before and during shoot production for the species cultivated exclusively for shoot
production in Australia.

In natural bamboo forests and in stands under low-input conditions, the nutrient-
supplying capacity of soils is usually the most important soil property governing bamboo
growth and yield (Chen et al.,1996). Soil physical factors, such as slope of land, texture,
bulk density, moisture-holding capacity, and temperature, however, are among the
nonchemical properties of soil which influence bamboo productivity. In contrast to other
soil physical and chemical properties, nutrient availability can be managed comparably
easily through fertilization. It is generally agreed that fertilization can have dramatic
impacts on bamboo productivity under “poor” site conditions and under minimal
management.

Oshima (1931) reported that as with vegetable cultivation, moso bamboo in Japan
requires abundant fertilizer, but there are few studies to show that excessive fertilizer
application reduced its yield and quality. Hong (1987) recognized the particularly
beneficial effect of fertilizer application on P.pubescens production on poor soils in
China. Widjaja (1991) described fertilization as the most important management
technique for commercial cultivation of species like D.asper in Indonesia. Ahmad and
Haron (1994) recommended regular fertilization for optimum bamboo growth and
performance in Malaysia. Meanwhile, Qiu et al., (1992) concluded that productivity of
bamboo forests under minimum management in China could be significantly increased
by fertilizer application. Fertilizers dramatically increased yield of D.strictus in India
(Lakshmana, 1994) and P.edulis in Japan (Suzuki and Narita, 1975). Nitrogen fertilizer
application in B.blumeana plantations in Philippines has been reported to enhance both
shoot number and culm yield, although there was no significant increase in culm diameter
or culm height (Tiongco, 1997). Fertilizers have been reported to increase shoot
production in B.blumeana and B.vulgaris in Philippines (Decipulo et al., 2009).

As for many other agricultural and horticultural crops, nutrient application rates, ratios
between nutrients, schedules of nutrient application, form of fertilizer, and nutrient
placement are equally important considerations in bamboo production. Since bamboo is a
perennial crop, however, nutrient management schemes that have been developed for
annual crops may not apply to it. Moreover, bamboo is grown for several products, and it
is understandable that optimal fertilization will vary with purpose of cultivation. Due to
perceived scarcity of petroleum resources in the future, there is a need to match efficient
fertilizer use to sustained productivity, and to sustain favorable soil conditions over the
short and long term. Soil N availability is of great importance and fertilizer application
can have dramatic impacts on bamboo growth and production of bamboo products .
Bambusa oldhamii, cultivated in Australia for edible shoots has been reported to have
marginally responded to increasing rates of fertilizer application with notable response
when it was applied in the inorganic form and combined with the higher irrigation rate
(Kleinhenz et al., 2000). Despite this knowledge, there is little information available on
the effects of N supplements on bamboo shoot production under different rates of water
availability. Nutrient management must not only satisfy requirements for yield but also
for quality of harvested parts. Very little research has been conducted on the effects of
fertilizer application on the many quality parameters of fresh edible shoots and bamboo
timber. There is, however, a general trend that with more N, P, and K applied to
P.pubescens stand, total yield of shoots and/or culms were higher due to the greater
number of shoots and/or culms harvested, and longer culms, but with reduced diameter
(Hong and Jiang, 1986; Hong, 1994).

Fertilization experiments were conducted by Shanmughavel and Francis, (1996c) in five


year old plantations of B. bambos at Kallipatty, in RBD with four treatments (0, 100, 200
and 300 kg/ha). The results showed that the 200 kg/ha of fertilizer was sufficient for
increasing the productivity of the species. However, it was reported in Japan that tissues
of shoots and culms harvested from plantations abundantly supplied with nutrients are
usually “softer,” resulting in lower shoot quality and less durable culms with poorer
mechanical properties (Ueda, 1960). For edible shoots, higher N decreases but higher P
increases, sugar content in P. pubescens in China (Hong, 1994). For the same species,
Zhu et al., (1991) quantified the effects of N, P, and K on shoot quality of P. pubescens
in China and reported that N, P, or K enhanced amino acid content, N decreased sugar
content, P increased sugar content, N increased hydrolytic acids and N and/or P lowered
free tyrosine of canned shoots.

Silicon has a special role in the nutrition of bamboo, because it is associated with cell
wall constituents and is present in xylem cell walls and fibers of plants. There is a general
agreement that Si should only be applied to bamboo stands for timber production. For
production of edible shoots, Si may prevent shoot development and reduce quality by
increasing fiber content (Hamada, 1982; Hong, 1994; Fu and Banik, 1995). As for many
other crops, bamboo responds more favorably to split application of nutrients throughout
the year than to a single annual dressing (Raina et al.,1988).
Suzuki and Narita (1975) reported that the number of sprouts of P.edulis from fertilized
plots was 1.7 to 1.9 times that of control in Japan. Also fertilizer experiments conducted
on Thrysostachys siamensis, Dendrocalamus asper, Bambusa spp.and D.strictus in three
year old plantation at Dong-larn in Japan showed that the use of 15:15:15 N:P:K fertilizer
at 100 kg ha-1 is sufficient to increase the yield (Suwannapinunt and Thaiutsa, 1990). In a
study conducted on growth performance of Dendrocalamus asper and Gigantochloa levis
stands fertilized and studied over a period of four years in Philippines, it was reported
that the influence of the organic fertilizers on average number of shoot emergents,
number of culms per clump, average height and culm diameter was only significant
during the first year of plantation establishment. However, no statistically significant and
consistent improvement in establishment of the clumps or growth and quality of the
culms could be assigned to the effects of organic fertilizers studied over four years of
study (Fernandez et al., 2003). In another experiment, (Azmy et al., 2004), the influence
of three organic fertilisers (chicken dung, palm oil mill effluent and peat materials
specially made for fertilisers) on the number of new shoots, the diameter at breast height,
the height of the culms and the number of culms in Gigantochloa scortechinii in
Malaysia was studied. The number of new shoots and the diameter at breast height were
positively influenced, which is promising for commercial purposes.

In Kerala, six bamboo species viz, Bambusa bambos, B. tulda, Dendrocalamus


hamiltonii, D. brandisii, D. longispathus and D. strictus produce shoot during June to
September. If regularly irrigated, the duration of shoot emergence can be increased. In B.
bambos and D. hamiltonii, the average number of shoots and production were 23 and 30-
50 kg and 53 and 20 – 40 kg respectively per clump in four years. A good clump
structure for plantation devoted to shoot production is in the ratio of 4:4:2 for 1st, 2nd
and 3rd year respectively and 50-60% of the shoot can be harvested every year. At
harvesting, a shoot may contain 80-90% moisture and the edible content is typically
around 30%. Since bamboo is a perennial, all culm management practices must also aim
to sustain long-term productivity of the stand (Virtucio, 1996). The intensity of removal
dictates the total number of culms per unit area remaining, and the choice of culms
harvested based upon maturity status governs the age structure of culms (i.e., the relation
between younger culms and older culms). Being a biological material, bamboo is
subjected to greater variability and complexity due to various growing conditions such as
moisture, soil and competition. Many factors, such as culm height, topography and
climate, affect the properties of bamboo to a great extent and thus its utilization. The
production of shoots is far less than the area covered by bamboo in India would indicate,
as bamboo is currently underexploited, being used primarily as a source of timber and
fibre and not for its vegetable shoots (Farrelly, 1984).

2.7.3 Effect of harvest on shoot emergence

Production of new culms in B.bambos was found to depend mostly on the degree of
congestion, culm age and rainfall of the previous year (Shanmughavel et al.,1997).
Induction of new shoot increases upto a certain age and gradually decreases which also
depends on soil and climatic condition (Banik, 2000, Shanmughavel and Francis,
2001).Studies on culm dynamics of plantation grown Bambusa bambos revealed that the
rate of production of new culms was linearly related to the clump size (Das and
Chaturvedi, 2006). Population structure of Bambusa cacharensis, B.vulgaris and
B.balcooa under traditional harvest regimes exhibited preponderance of younger age
class than older ones (Nath et al.,2006). A study on growth of Dendrocalamus
longispathus Kurz. in Bangladesh indicated that on aging of the clumps, the culms
emerging in subsequent years were gradually taller with larger diameter and more
internodes. In a culm in its third and fourth years, leaf production and growth were low,
but in the fifth and sixth year these sharply declined and ultimately the culm died. Thus
harvesting of culm within three to four years of age could be an ideal management
practice in the species (Banik and Islam, 2005). Also, the removal of culms will promote
the growth of new shoots, in time for the incoming rainy season. The cut should be made
as close as possible to the base (Bareja, 2010).

It has been recorded that Bambusa vulgaris, B. balcooa and B. tulda produced only 5
culms per clump in the first year. The number of culms gradually increased in the
subsequent years and became maximum (17 culm) in the 6 th year of clump age. The
number of culm production then slowly decreased from the 7 th year and was only 6 in
the 9 th year of culm age (Shanmughavel, 1995; Banik, 1988). The crowded and
congested condition in most bamboo species has been reported to create a scarcity of
room for the new emerging culms and also increased competition among them for
survival (Kondas, 1981, Banik, 1993). Probably due to this reason culm production
subsequently decreased from the 6 th year in the Bambusa species. Removal of dry culms
increased the productivity of bamboo (Singh, 2002).

2.8 Biomass and productivity

2.8.1 Biomass studies on bamboo – World wide and in India

Biomass is the total quantity of organic matter per unit area present in an ecosystem at a
given time and may relate to a particular species or community as a whole. It is expressed
on the basis of oven dry weight. Such biomass studies are important to know the
biological productivity and at times to understand the economic productivity also. The
relative allocation of biomass to various above ground parts is a decisive factor that
reflects the success of an organism in an environment (Gadgil and Solbrig, 1972).
Aboveground biomass production in bamboo has two components: growth of newly
emerging culms and biomass increase in older culms. The culm biomass and relative
allocation of various fractions to total biomass varied markedly among the species. Total
biomass in an area depends on stocking level. Therefore the standing stock is one of the
important measure for site productivity, overstocking and/or understocking may retard
the growth and development of the bamboo (Shi et al.,1993). With regard to the
percentage contribution of biomass components of the above ground biomass the
plantation age was observed to be an important factor. The percentage contribution of
leaf (7% to 1 %) in B.bambos, in Tamil Nadu, India, was found to decrease with
increasing age (Shanmughavel et al.,1997). Kigomo and Kamiri(1987) also found that the
relative contribution of various components to the standing biomass of Oxytenanthera
abyssinica, grown in Kenya, was in the order : bamboo culm> Bamboo branch> bamboo
leaves.

Above ground biomass production in growing bamboo (Bambusa bambos) in Tamil


Nadu, India from two days old (0.12 m height) to maturity (28.5m height) were studied
over five months (Shanmughavel and Francis, 1996a). There was a linear increase in the
total above ground biomass for all components with the percentage contribution of culm
being maximum followed by branches and leaves. Singh and Singh (1999) estimated the
biomass of Dendrocalamus strictus in dry tropical region in India. The results showed the
total biomass of 46.9 t ha-1 in 3-year old to 74.7 t ha-1 in the 5-year old plantation with
35% of biomass remaining below ground.

The biomass production in an age series of Bambusa bambos plantations estimated in


Tamil Nadu (Shanmughavel et al., 2001), showed a linear increase of the biomass of all
the compartments with the age of the plantation. In the above ground biomass, the
percentage contribution of culms (81%), branches (14%) and leaves (1%) was 96%,
whereas in the below ground, rhizome contribution was only 4%. The total biomass
ranged from 2.3 t DM ha-1 (1st year) to 297.9 t DM ha-1 (6th year). The mean annual
biomass production was 49.6 t DM ha-1 up to 6th year period and there was gradual
decrease in biomass accumulation after the 6th year (Shanmughavel and Peddappaiah,
2001). The biomass accumulation in G. angustifolia in Columbia found a different
pattern from B. bambos. Riano et al., (2002) worked on growth and biomass distribution
on G. angustifolia in Columbia in relation to age. The results on samples from complete
clumps with different ages showed the accumulated biomass of 54.3 t ha-1. The aerial
portions which include culm, branches, foliage, leaves and caulinear leaves accumulate
80.1% of the total biomass and CO2 fixation, and the rhizome 19.9% of these. Biomass
contribution of G. angustifolia to soil biomass in the sixth year was around 10
tons/ha/year, and dry matter accumulation was found to reach 76.6 tons ha-1 including
culms, branches, foliage leaves and cauline leaves.

However in case of D. asper the pattern of biomass accumulation was different. The
diameter and basal area growth, biomass accumulation and net production were all
substantial increased with the increasing stand ages in D. asper in Taiwan. The
maximum net annual production of 41.4 ton ha-1 was recorded at eighth year after
planting. An average of 9.3 ton ha-1 of biomass was obtained between the age of 12 to 15.
The plantation reached its highest productivity at 26.3 ton ha-1, one year after removing
the culms older than four years old. (Kao and Chang, 1989)

The biomass accumulation of monopodial bamboos are different from sympodial


bamboo. Isagi et al., (1997) have studied the net production of Phyllostachys pubescens
in China and the standing culm density and average DBH in 1991 were 7100 ha-1 and
11.3 cm, respectively. The above-ground biomass was 116.5 t ha-1 for culms, 15.5 t ha-1
for branches, 5.9 t ha-1 for leaves and 137.9 t ha-1 in total. The total above-ground
biomass was one of the largest among the world’s bamboo communities. The biomasses
of rhizomes and fine roots were 16.7 t ha-1 and 27.9 t ha-1, respectively. Oli (2005)
prepared biomass tables for Bambusa tulda grown in Eastern Nepal. The total of 153
culms from 59 clumps were selected for growth measurements and samples were oven
dried to estimate the biomass. A regression model was developed on the basis of oven dry
and green weight. The model used was W = a + b ×(D2L). Based on the oven dry weight,
the R2 values were obtained for culm, branch and foliage components, which were 92, 81
and 83%, respectively. Similarly, R2 values for culm and foliage components on the basis
of green weight were 92 and 82%, respectively. The R2 values obtained for branch and
foliage components were slightly lower as compared to the culm. This equation could be
useful in estimating bamboo biomass of managed natural stands or plantations in similar
site conditions. Krishnankutty and Chundamannil (2005) also developed a commercial
weight tables for Bambusa bambos from forests of Kerala for different clump-diameters
based on the measurements of 610 clumps and culms enumerated. Using the data on
commercial weight of green culms and clump-diameters, different weight prediction
equations were estimated. Based on the best fitting equation, a commercial weight table
for green culms as well as dry culms in a clump was generated for different clump
diameters. These tables are ready to use and facilitate making quick estimates of bamboo
growing stock in forests.

The Masha natural bamboo forest, Ethiopia was stocked with 8840 clumps ha-1,
uniformly distributed with a mean height of 16.8 m and diameter of 7.6 cm. Culm
contributed 82%, branch 13% and leaf 5% to the 110 t ha-1 total above ground biomass,
while clumps mature for harvest (>3 year) made up 73%. The culm component of the
mature clumps was 60% of the above ground biomass, whereas the biomass of current
shoots (<1 year) constituted only 7% (Embaye et al., 2005).

Das and Chaturvedi (2006) estimated the culm recruitment, standing biomass and net
production at 3, 4 and 5 years in B. bambos plantation in Bihar, India. The number of
culms produced per clump varied from 22 to 31. The total biomass was 170.8 Mg ha -1 in
3 year old to 257.3 Mg ha-1 in 5 year old of which culms contributed (65%), branches
(21%) and leaves (4%). The standing stocks of rural bamboo (Bambusa bambos) in
Palakkad and Malappuram districts were assessed (Kumar, 2008) through a two-stage
stratified random sampling scheme in which 600 homegardens in 20 panchayaths were
surveyed. Total growing stock of thorny bamboo in Palakkad district was estimated at
61,196 clumps or 47, 87,041 culms, having a total dry weight of 1, 24,389 Mg. The
corresponding figures for Malappuarm were 44, 215 clumps, 33, 99, 855 culms, and 86,
267 Mg dry weight. Average cash income generation per farmer through bamboo sale
during the period from 2000 to 2005 was, however, low with Rs. 4002 and 1834 per
garden per year for Palakkad and Malappuram respectively. In a similar study, Tian et al.
(2010) estimated the biomass and carbon storage for Phyllostachys makinoi in Taiwan
with a stand density of 21191±4107 culms ha-1. The results showed that bamboos of all
ages shared a similar trend, where culms displayed a carbon storage of 47.49–47.82%,
branches 45.66–46.23%, and foliage 38.12- 44.78%. Inspite of the high density of the
stand, the diameter distribution of makino bamboo approached a normal distribution and
aboveground biomass and carbon storage were 105.33 and 49.81Mgha-1, respectively.

Rai et al., (2013) studied the growth and biomass accumulation in six year old Meloccana
baccifera plantation in Mizoram, India. The result showed that the number of culms
varied from 2165 to 4190 per ha.; the biomass ranged between 8.212 to 9.025 kg/culm
and195.391 t ha-1 to 362.56 t ha-1.

Nath et al., (2009a) reported that above ground stand biomass of village bamboos of
Barak valley, North East India was 121.51 t ha-1 of which 86 per cent was contributed by
culm component followed by branch (10 per cent) and leaf (4 per cent). B.balcooa
plantation raised under precision farming has shown annual yields of over 100 t ha-1 and
stabilization at 150 t ha-1 from the fifth year onwards (Barathi, 2010). Agarwal and
Purwar (2010) reported that the total biomass produced was 30 per cent more on fresh
weight basis than dry weight in six year old plantation of five species viz.
Dendrocalamus strictus, Bambusa vulgaris, B.multiplex, B.bambos and Phyllostachys
nigra in Uttarakhand, India. Average biomass produced was D.strictus – 581.8,
B.vulgaris – 832.4, B.multiplex – 15.9, B.bambos – 703.1 and P.nigra – 72.5 kg ha-1 pole-
1
on dry weight basis.

Biomass can vary substantially within individual species, even when cultivated at the
same site. Due to their high genetic variability, species of Dendrocalamus such as D.
latiflorus are well known for such differences. Although D. latiflorus, in China, may
have a yield potential of up to 41 t/ha of edible shoots per year (Pan, 1986), Lü et al.
(1997) found great variations in growth between plants of D. latiflorus in Taiwan. Kiang
et al. (1976) measured yields of edible shoots ranging from 7.4 to 20.3 kg/clump/year in
different strains of D. latiflorus growing in China.

The North-east region of the country holds more than 67% of the growing stock of
Bamboo in India. The 83% of the total growing stock comprises of three main species
viz., D. strictus (53%), B. bambos (15%) and M. baccifera (15%). The estimated annual
harvest which is put to different uses is about 13.47 million ton. The average
productivity of Indian bamboos averaged at 1.5 t ha-1 yr-1 which is very poor in
comparison to that of China (3.79 t ha-1yr-1) (Gupta, 2008). In most part of the country,
bamboo resource base has been under managed and is commonly over exploited.
However, stand type, species and region wise productivity varies.
Table 2.2. Reports of Biomass of six bamboo species (in Megagram per Hectare)
taken up for the current study across the world

S.No. Species Country Age Total Biomass Reference


-1
Mg ha
4 122.0
Shanmughavel &
6 225.0
Francis, 1996a
8 287.0
1 Bambusa bambos India
3 170.8
Das and Chathurvedi,
4 206.7
2006
5 253.7
3 46.9
Dendrocalamus Singh and Singh 1999
2 India 5 74.7
strictus
3 207 Singh et al., 2004
3 Guadua Columbia 6 76.6 Riano et al, 2002
angustifolia
Taiwan 8 41.4 Kao and Chang, 1989
Dendrocalamus
4 India 2 0.21 Agarwal, A.& Purwar,
asper
J.P. 2009.
5 150 Barathi, 2010
5 Bambusa balcooa India
6 69.06 Jijeesh, 2013
6 Dendrocalamus India 8 66.47 Rane et al., 2014
stocksii
In Assam, the annual yield of M. baccifera and B. tulda is 3.1to 5.0 t ha-1yr-1 while for D.
strictus and B. bamboos in Western Ghats, the figures are 3.0 and 6.0 t ha-1yr-1
respectively. It is seen that in case of some species, intensive cultural operations and
scientific management can produce much higher yield. In Taiwan, intensive cultivation
with fertilization in D. giganteus resulted in 20-30 t ha-1 yield. (Singhal and
Gangopadhyay, 1999, Tripathi, 2008). With regard to biomass production by bamboos,
there is a wide variation depending on species and region of cultivation. Even within the
same region, production potential varies due to clump densities, mode of plantation and
silvicultural management adopted. Total above ground biomass in both monopodial and
sympodial bamboo types generally varies between 7 to 165 tha-1 as recorded from 26
bamboo species worldwide (Kleinhenz and Midmore, 2001). Total biomass production
from natural bamboo forest (D. strictus) of Chatra and Latehar in Jharkhand has been
recorded from 1.30 to 43.63 t ha-1 due to varying clump density. In plantations under
same edapho-climatic conditions in Australia, the above ground biomass production by 6
yr old different species was found to be : B. balcooa – 17.125 tha-1, B. bambos- 17.116
tha-1, B. nutans - 16.39 tha-1, B. vulgaris- 12.905 tha-1, D. strictus- 12.156 tha-1. On an
average allocation of total above ground biomass to culms, branches and leaves was 82,
12 and 6% in monopodial species and 77, 13 and 10% in sympodial bambos (Kleinhenz
& Midmore, 2001).

Rane et al., 2014 reported that aboveground biomass of evaluated clumps distributed
amongst various elevation zones along the Western Ghats varied for individual culm
biomass and total biomass of old culms. Individual culm biomass of the clumps located
between 101 to 650 MASL was comparatively higher. Elevation gradients create spatial
heterogeneity in Western Ghats resulting in variation in culm growth and biomass
parameters of Dendrocalamus stocksii. Culm biomass production in D. stocksii was
optimum between 101 to 650 MASL elevation in Central Western Ghats as this area
receives maximum rainfall and is endowed with favourable edapho-climatic conditions.

2.8.2 Physiological studies using IRGA

Studies have been conducted on various agricultural species like sorghum, rice etc on the
correlative impact of irrigation, fertilization and photosynthesis on the productivity and
total biomass of the plant. Measurements of the photosynthetic process are useful for
understanding how it might be controlled and how productivity can be increased by
the phytomonitoring of plant development. Measurements of photosynthesis are
essential for comparing and understanding productivity (biomass accumulation) of
vegetal systems at the leaf, plant or community level as well as their response to
environmental stresses. Evaluation of the behavior of certain physiological parameters
can effectively be used to assess the performances of a species under given
environmental conditions. Leaves usually exhibit a remarkable capacity to adjust their
morphology and physiology in response to different light conditions due to their
phenotypic plasticity (Bond et al., 1999; Valladares et al., 2007; Yoshimura, 2010).

Net photosynthesis rate (Pn), transpiration rate (Tr) and total leaf area per plant are the
important factors that determine the biomass production and Water Use Efficiency
(WUE) of a species. Variation in Pn, has been reported as determinant of plant
productivity of the species (Nataraja and Jacob, 1999). Reports state that photosynthetic
rate varies among the plants belonging to different taxa and also among the varieties
within the same species (Arora and Gupta, 1996). The water molecules lost per molecule
of carbon fixed by the plant during photosynthesis, is referred to as water use efficiency
(Ellsworth, 1999). Intrinsic water use efficiency (Pn/gs) implies the inherent ability of the
plant to assimilate CO2 (Ares and Fownes, 1999). A higher value indicates better ability
of the plant for carbon assimilation (Rekha Warrier et al., 2013).

Some earlier reports indicate that there is poor correlation between photosynthetic rate
and biomass production in crop plants as reported by Evans and Dunstone (1970),
Heichel and Musgrave (1969), and Rhodes (1972). However, later reports suggests that at
least 90% of the dry matter of higher plants is derived from CO2 assimilated by
photosynthesis (Zelitch, 1982). Total biomass accumulation is a function of the rate
of biomass production (associated with leaf area production, leaf area retention,
and net photosynthetic rate) and growth duration (Austin, 1981). Zelitch (1982)
stated that the lack of a positive relationship between leaf photosynthesis and
production in most studies is due to instantaneous photosynthetic measurements
conducted at a single period of development under ideal laboratory conditions,
rather than seasonal measurements conducted under field conditions. In addition,
single leaf measurements fail to account for canopy leaf area and architecture
differences which influence light interception and whole canopy CO2 assimilation
(Wells et al., 1986, Peng et al., 1991). Positive relationships between photosynthetic
rates at the canopy level and plant productivity have been reported for wheat
(Puckridge, 1971), barley (Biscoe et al., 1975), sorghum (Kanemasu and Hiebsch,
1975), maize (Vietor and Musgrave, 1979), soybeans (Wells et al., 1982) and
cotton (Wells et al., 1986). The products of photosynthesis are the primary
determinants of plant productivity, and increasing photosynthesis has been widely
recognized as a key trait to increase yields (Long et al., 2006; Zhu et al., 2010; Parry
et al., 2011; Raines, 2011).

In common with other crops, the total photosynthesis of bamboo stands is determined by
the leaf area and by the photosynthetic capacity of the canopy. The photosynthetic
capacity of an individual leaf depends on its age, its nutritional status, the age of the culm
it is growing on, its position in the canopy, and on climatic conditions such as
temperature and photon-flux density which vary with season. The photosynthetic
capacity of the bamboo canopy is not only determined by the photosynthetic capacity of
leaves, but also depends on the life span of leaves which influences the leaf-age structure
of its canopy. The life span of leaves is substantially different between monopodial and
sympodial bamboos. (Qiu et al., 1992; Huang et al., 1993; Isagi et al., 1993; Fang et
al., 1998; Li et al., 1998). Understandably, the leaf area index (LAI) of a complete
bamboo canopy increases with leaf biomass across bamboo species and ranges from 5 to
12. It is generally recognized that photosynthetic activity of leaves steadily declines with
age after full expansion. Huang (1986), Huang et al., (1989), and Qiu et al., (1992)
noted photosynthesis rates up to three times higher in new (<1 year old) leaves compared
with those of older (>1year old) leaves. For example, the net photosynthetic rate of new
leaves (2 months old) was 2.5 times greater than that of 1-2-year-old leaves in P.
pubescens (Huang, 1986). This was attributed to greater metabolic activity of tissues
(Huang, 1986) and higher nutrient concentrations (Zhou and Wu, 1997) in younger
leaves. For growth and production of commercial bamboo, it is particularly important to
know the sources of energy and nutrients that sustain the rapid accumulation of biomass
in young culms. Since shoots and later young culms have essentially no photosynthetic
leaf area to provide energy and sustain their growth, net-import of energy is required at
least for their early growth (Pearson et al., 1994; Liese, 1995). The photosynthesis rate
of bamboo leaves not only depends on their age, but also independently on the age of the
culm upon which they are found. In young (<7-8 years old) bamboo stands, leaves on
older culms are shaded more than those on younger ones since younger culms are taller
than older culms until the stand reaches its maximum height. In mature (>7-8 years old)
stands, transmission of light decreases rapidly from the top of the bamboo canopy
towards the bottom (Yang et al., 1988).

During January and February, the photosynthetic rate of leaves is generally low due to
low air temperature (Huang et al., 1989: Koyama and Uchimura, 1995). In studies
conducted by Koyama and Uchimura (1995) with P. bambusoides, net photosynthesis
increased until air temperature was 27°C, but decreased rapidly thereafter. The
compensation point at which the respiration rate equals that of photosynthesis was a little
below 40°C. Net photosynthesis rates (Pa) reached saturation at 12 µmol C02m-2s-1 when
the PAR was 1,200 µmol-1m-2s-1. Therefore, P, in P. bambusoides peaked around midday.
It is probable that “critical” levels especially of temperature and perhaps of PAR will be
much higher for tropical and typically sympodial bamboo (Kleinhenz and Midmore,
2001).

Gratani et al., (2008) studied the growth pattern and photosynthetic activity of different
bamboo species (Phyllostachys viridi-glaucescens Rivière et C. Rivière, Phyllostachys
pubescens Mazel ex Lehaie, Phyllostachys bambusoides Siebold et Zucc., and Bambusa
ventricosa McClure) growing at the Botanical Garden of Rome. Among the considered
species, P. pubescens had the highest mean culm height (14.3±0.6 m) and diameter
(10.7±1.5 cm), while B. ventricosa had the lowest mean culm height (6.0±0.2 m) and
internodes number (35±1). The highest net photosynthetic rates (NP) of the considered
species were measured in early autumn, while the lowest ones in spring (30% of the
maximum in the genus Phyllostachys), in the period of vegetative activity, and in winter
(10% of the maximum in B. ventricosa).
Net photosynthesis rate (Pn) and stomatal conductance (gs) are often strongly related
(Kundu et al., 1998). In most of the species, photosynthesis rate increases when stomatal
conductance increases (Heber et al., 1986; Bunce, 1988). Reduced stomatal conductance,
coupled with increased photosynthesis may also improve the water use efficiency (WUE)
of many plants, which is of particular importance for productivity in arid and semi-arid
regions (Polley et al., 1992; Polley et al., 1993; Sage, 1995; Field et al., 1995; Owensby
et al., 1999; Smith et al., 2000)

Transpiration is one of the major gas exchange related traits associated with plant growth
and productivity. In perennial species stomatal transpiration contributes more than 90%
of total transpiration (Taiz and Zeiger, 2002). Transpiration and photosynthesis are two
major gas exchange parameters which determine WUE of plants. Studying the yield
potential or complex traits like Water Use Efficiency (WUE) of any perennial plant
requires long gestation period experiments. To overcome this problem, Nataraja and
Jacob (1999) suggested to look into the behavior of growth related physiological
parameters such as photosynthesis to evaluate the performance on field. Positive
correlation between photosynthesis and biomass has been reported in different species
(Mythili and Nair, 1996; Gerik et al., 1996; Hampson et al., 1996; Atkin et al., 1997;
Thomas et al., 1998, Orlovic et al., 1998). Arun Kumar et al., 2009, inferred that in a
perennial species like Santalum album, clones having high photosynthesis might have
high growth rate and biomass production ability. Instantaneous WUE has been reported
to vary among clones of the same perennial species thereby serving as a parameter for
selection of potential parents for tree improvement for WUE (Kannan et al., 2007;
Nataraja and Jacob, 1999) . Li (2000) reported that measurement of WUE may be a
useful trait for selecting genotypes with improved drought adaptation and biomass
productivity under different environmental conditions. Combined with reduced stomatal
opening, conductance and transpiration rates, elevated CO2 concentration depresses dark
respiration rates leading to increased water use efficiency (Wullschleger et al., 1992;
Murray, 1995). Increase in water use efficiency has been found to increase drought
tolerance in many plant species, which may allow increased plant distributions (Tyree
and Alexander, 1993; Huxman et al., 1998). Whether future increases in plant
distribution are realized is dependent upon whether increased water use efficiency will be
greater than enhanced transpiration, as a result of global warming (Houghton et al., 1990
& 2001; Crowley, 2000). Net photosynthesis and related gas exchange parameters have
been suggested as early selection criteria to improve the efficiency of breeding and
improvement (Balasubramanian and Gurumurthi, 2001).

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