Nutritional aspects of buffalo production

Introduction

Buffalo plays crucial role in rural livelihood, food security and agricultural economy. Apart from milk and meat, buffaloes contribute significantly to draught power for agricultural operations as well as through dung for fuel and soil improvement. Farmers preferred buffaloes is over cattle in many parts of the country owing to its more quantity of milk yield, disease resistance, longer productive life and higher milk productivity. Nutrition plays a vital role in livestock rearing. Optimum nutrition is of paramount importance to exploit full production potential from buffaloes. Therefore, nutritional aspect of buffalo production and management should be considered scientifically and proper attention should be given to the feeding management to achieve maximum output from this species. India possess largest population of buffaloes i.e. 108.7 million buffaloes (18 th Livestock Census, 2007) with an annual growth rate of 1.84% and shares 57% of world buffalo population and thus first in buffaloes and milk production (19 th Livestock Census, 2012, DAHD & F, Govt. of India). Buffaloes are reared for milk and meat production. India, with over 53% of the world's buffalo meat productivity, produces 39% of global buffalo meat and contribute 55% of total milk production, highlighting the untapped potential for increased production (Kondaiah, 2025) [31] . Apart from this, buffaloes termed as Black Gold, and contribute various byproducts such as leather, dung, blood, bones etc. apart from milk and meat which helps to improve the economic status of the farmers. The drought power from buffalo bullocks in field operations which have added advantage goes unnoticed. However, production performance in terms of milk, reproduction and growth rate (meat) is lower as compared to their western counterpart (Singh and Mehra, 1990; Qureshi et al., 2002; Sahoo et al., 2004; Wynn et al., 2009) [47,43,45,56] which is mainly attributed to the poor state of nutrition that accounts for the major share in cost of production. There is net deficit of 40, 57 and 36% of dry forages, concentrates and green forages, respectively in India (NABCONS, 2007). This area hence needs proper attention of researchers across the globe to enhance the production potential of buffaloes.

Use of unconventional or agro-industrial byproducts in buffaloes is promising but restricts its use due to presence of one or more anti-nutritional factors which warrants proper processing before feeding. In past, several attempts have been made to process and improve the nutritive value of feed resources by various process and treatments (Katole et al., 2011) [26] . However, there are certain constraints in adopting the methods of processing, cost being a major constraint. Improving nutritional status of buffalo is still a grey area of research to improve the productivity from buffaloes. The buffalo species is originated in India and are descendants of arni breed in wild found in north-eastern parts of India. Buffalo in Asia are also known as Water buffalo are classified under the genus Bubalus, species bubalis. Asian buffalo are classified in two subspecies i.e. River and Swamp. Buffaloes are commonly known as Asian animal and India is considered as the home tract for best buffalo breeds of the world. India stands first country in Asia for scientific and technological development in buffalo nutrition, production, reproduction, biotechnologies and genetic improvement (Borghese, and Mazzi, 2005) [10] . Moreover India has implemented the "Red Revolution" to increase meat production and strengthen the meat industry, particularly with regard to buffalo. In India, out of 10 recognized buffalo breeds, best River milk breeds are Murrah, Nili-Ravi, Surti and Jaffarabadi, which are originated from the north-western states (Table 1 &2). These breeds have high potential for milk and fat production (Sethi, 2003) [46] . Apart from this, buffaloes are used for drought power and meat production. In some Asian countries, buffaloes are used in social and cultural events. India is endowed with rich biodiversity of buffalo germplasm in the form of different recognized breeds and several local distinct types.

Table 1: Buffalo breeds of India with production traits (ICAR, 1997) [23] Name of Breed [5]

Murrah Group

Gujarat Group Uttar Pradesh Group Central India Group South India Group

In countries like Australia and Bulgaria, the meat industry is developed on the basis of meat production from buffaloes on large scale. Dressing percentage of buffaloes is 55-65% on moderate diets. Buffaloes are the good source of quality meat. Buffalo meat is leaner and less fat as compared to cattle thus there is market demand for such meat among health conscious consumers.

Buffaloes are well adapted to the hot and hot humid climate. Byproducts from buffalo production include milk, meat, skin (hide), blood, bone etc. Thus buffalo rearing may be helpful to uplift the socio-economic status of marginal or landless farmers. Apart from this, buffaloes are the financial asset and acts as an insurance against the risk of crop failure due to natural calamities (Dhanda, 2004) [15] . Indian swamp buffaloes reared for milk production, males are used for work in fields and for other agricultural operations (Faruque, 2003) [18] . On an average buffaloes are almost 3-4 times more productive than indigenous cattle. The research findings on the comparative functional rumen developments and ferementation activities suggest that the buffaloes in general are more efficiently utilizes the poor quality roughages and have better feed conversion ratio (Bartocci et al., 1997; Agarwal et al., 2009) [7,3] . Other reason might be that buffaloes are not discriminate during foraging and consume larger amount of coarse forages which are generally rejected by other animals. Thus are better at converting poor-quality roughage into high quality milk and meat. The other significant point which gives buffalo rearing upper hand is that their low cost of milk production per litre and gives better economic returns. They have almost 5% higher fibre digestibility than cattle and 4-5% higher efficiency of utilization of metabolic energy for milk production (Mudgal, 1988; Terramoccia et al., 2000) [48,33,52] . In buffaloes dry matter intake per unit body weight is also lesser than cattle.

Protein and energy

Buffaloes, from birth to death, passes through various physiological stages, thus require different nutritional allowances through different stages of life, otherwise they may be prone to various metabolic and transition disorders which limits its productivity (Katole et al., 2013b) [25] . Dry matter intake of buffaloes depends on body weight, production level, physiological stage, roughages and concentrates ratio, quality and quantity of the feeds used in ration formulation. Most important physiological stage is lactating period which need utmost nutritional attention.

Buffaloes require protein and energy to produce quality milk during lactation period. Energy and protein requirements of ruminants can be met from fermentation of feedstuffs in rumen which produces fermentation end-products such as volatile fatty acids (to fulfill energy requirement) and microbial protein (to fulfill protein demand). This depends on the amount and quality of feedstuffs offered to the animals.

The maintenance requirements of the indigenous buffaloes are somewhat lower than their western counterparts. Growth is important parameter and generally indicated by weight gain per unit time. Thus, growth phase require additional allowances as it is influenced by feeding and environmental conditions. As cost of concentrates is escalating, it is not feasible to feed buffaloes at higher level. The average growth rate in indigenous buffaloes is 400-700g/day and in crossbred buffaloes it may be even higher. Faster growing buffaloes require more energy and protein along with minerals such as calcium and phosphorus. For such animals bypass protein is recommended.

Lactation period is preceded by dry period which is of 280-310 days. During this period, reduction in nutritional allowances results in low lactation yield and poor health (Zicarelli, 2000) [58] . Nutritional requirement of foetus also increases as the dry period advances and buffaloes should be fed maintenance plus gestation rations. Therefore, provision of additional allowance is essential to maintain the reproductive performance of the animal. It is suggested that non-lactating buffaloes should be given 0.63-0.65 Milk Feed Unit (FU)/kg DM and 10-11% of crude protein (Proto, 1993; Bertoni et al., 1994) [41,8] . Bertoni et al. (1994) [8] also recommended that crude protein level in dry buffaloes should be more than 10%, otherwise lower quantity of crude protein may compromise the rumen activity and production will be lower. Energy requirements for maintenance are 97.8-188.8 kcal/kgW 0.75 in dry and lactating buffalo. For 4% fat corrected milk (FCM) energy required is 1171 to 1863 kcal/kgW 0.75 . Protein requirements for maintenance are 1.28 to 3.48 g/kgW 0.75 in dry and lactating animals. For milk production protein requirements is 126.6 to 166.34 g/100g protein secreted in milk (Banerjee, 2009) [5] . It is reported that good quality ad libitum diet should be provided to non-lactating buffaloes that provide energy content not less than 0.65 Milk FU/kg DM and crude protein content not lower than 9% (Di Lella, 2000) [16] . Diet of buffaloes should comprised of fresh green forages or good quality hay and 25-30% of dry matter form concentrates to reestablish reserves of fat soluble vitamins and minerals to normalize the rumen fermentation and hepatic functions (Terramoccia et al., 2005) [51] . The diet mentioned above causes low rumen fermentation rate, which condition the production of volatile fatty acids and favors the proliferation of cellulolytic bacteria. The nutritional requirements of the buffalo are increases after parturition, accordingly supplemental feeding should be started before parturition to avoid sudden shift in dietary regimen. Energy and protein level should be increased by about 50% and 25%, respectively (Di Lella, 2000) [16] . Low protein 7-8% in diet could be justified by the nitrogen metabolism in buffaloes that differs from cattle (Abdullah et al., 1990; Kennedy et al., 1990) [2,29] but protein should not be less than 10% of diet. More digestibility of protein is reported in buffaloes than cattle due higher carbohydrates in ration (Puppo et al., 2002) [42] . Thus carbohydrates should form the major portion of diet. This also might be due to higher protozoal population in buffaloes. Protein level in diet of buffaloes should be in range of 9-14% on DM basis (Sivaiah and Mudgal, 1978; Verna et al., 1992; Verna et al., 1994; Rai and Aggarwal, 1991) [48,54,55,44] . Quality protein if used should be bypass or rumen undegradable, otherwise it will be extensively degraded in rumen causing increased concentration of ammonia and higher levels of blood urea. Reported reduced feed intake and conception rate in cattle due to increased levels of ammonia in rumen and blood urea. Similar results in buffaloes could also be expected if more rumen degradable protein is fed to buffaloes. Protein concentrations used in lactating buffalo diets can be equal to or below 12% on DM basis, since these concentrations have little influence on the quantity and quality of milk produced (Verna et al., 1992; Verna et al., 1994). Sivaiah and Mudgal (1978) [48] suggested administration of 166 to 126 g of digestible CP/100 g of milk protein produced, while according to Rai and Aggarwal (1991) [44] , the concentration of CP on DM should be between 11-14%. [59] . Higher Ca:P alteres Ca:Mg ratio in blood resulting in excitability of the uterine-vaginal muscle fibre causing atonicity of the organ leading to prolapse (Campanile et al., 1989) [11] . Excess calcium during dry phase which causes reduce parathyroid activity at calving should be avoided. More phosphorus rich feed ingredients need to be fed to the animals to stimulate parathyroid activity (Campanile et al., 1995) [12] . To avoid the alterations in mineral balance and maintain the balance of major minerals it is suggested to offer oat hay, wheat straw and maize silage not more than 5.0-7.0 kg/head/d (Technical-Scientific Committee, 2002) [50] . These roughages being poor in minerals do not cause any variations in diet of buffaloes.

In buffaloes, lactation phase is of 270 days and peak milk yield is observed during 4-6 th week of gestation. As compared to cattle, quantity as well as quality of milk shows much variation during lactation period. Energy level in diet positively influences the percent fat in milk of buffaloes which varies from 6-12%. Similarly, protein content in milk which varies form 3.5-5.5% also depends on the protein content of the diet. According to the fat and protein percent, feed ingredients should be added to the diet of lactating buffaloes. Major minerals that should be considered during ration formulation are Ca, P and Mg.

As mentioned above milk yield increases gradually and peak is observed during 4-6 th week of gestation, nutrients requirements of buffaloes during this period are also increases accordingly. For every increase in 1 kg of milk, dry matter intake of 500 g and 0.75 milk FU need to be increased. After 150 days of parturition buffaloes raise their feed intake and accumulate excessive body reserves. This is due to shift from the catabolism to the anabolism phase of the lactation curve.

One should be careful to avoid excessive fat accumulation. This can be accomplished by using diets having low energy density, starch and more NDF. More fat tissue deposition observed in buffaloes with low yield and more lactation period (above 270 days) due to fertility problems (Terramoccia et al., 2005) [51] . Protein degradation in rumen of buffaloes is more than that in cow (Terramoccia et al., 2000) [52] which might be attributed to more retention time in buffaloes (Bartocci et al. 1997) [7] . Production of protein in milk yield, growth and body weight gain and endocrine metabolic effect should be considered while formulating protein requirement for buffaloes. It has been suggested that similar values of dairy cow of 2.5 g CP/g protein in milk are sufficient. Generally in early lactation feed intake is lower. The ration containing less than 13.5% CP is not adequate thus, CP should be increased by 10% (Campanile et al., 1995) [12] . Buffalo's milk contains 1.8-2.0 g Ca and 1.1-1.2 g P. Reported that Ca and P requirements for one kg of milk production are 5.2-5.8 and 2.1-2.3 g, respectively.

Energy and protein requirements for one kg of milk production for buffaloes are given in Table 4. Apart from energy and protein requirements, the mineral requirements and demands should also be considered, with special emphasis to the calcium, phosphorus and magnesium in ration of buffaloes. The production requirements are 6.7, 2.2 and 0.9 g for Ca, P, and Mg, respectively per kg of milk produced (Proto, 1993) [41] .

Table 4: Energy and protein requirements for the production of one kg of buffalo milk relative to the fat and protein content (Proto, 1993) [41] Energy requirements (milk FU/kg of milk) Milk Fat (%) 6.

Processing of poor quality feed resources

Dry forages are low in nutrients and rich in fibre are the main roughage source used for feeding of buffaloes in India.

Considering poor nutritional quality of dry roughages, it is necessary to process these dry roughages before feeding to improve their nutritive value. Such processing of available feed resources is helpful to exploit maximum production from buffaloes. Various methods have been tried in past to improve the nutritive value of the dry roughages (Katole et al., 2011; Katole et al., 2013a) [26,25] . Out of various methods urea treatment seems to be promising as is economic and easy to adopt by farmers. Due to urea treatment nitrogen and in turn protein content is increased in processed forage. Supplementation of chemically treated straw at 30-50% of DM of feed or 0.9 to 1.5% of live body weight may be sufficient as well as cheaper to sustain milk production and maintain optimum health status. Other supplementation such as supply of urea molasses with green fodder i.e. leguminous fodder, tree leaves of cassava, glyricidia, leucaena, water hyacinth and sweet potato etc. (Banerjee, 2009) [5] are low cost and beneficial. Feed intake, palatability, digestibility, protein and energy of dry treated forages and milk yield are reported to increase by urea treatment (Nisa et [36,30] . During shortage or scarcity of forages, these dry forages are the only source available to feed buffaloes in tropical countries. These dry forages need to be properly supplemented or combined with molasses or mineral mixture to enhance the productivity during lean periods. In India, Urea Molasses Mineral Block (UMMB) has been developed and studies were conducted on the buffaloes which showed promising results in terms of increased growth rate and milk yield (Singh and Mehra, 1990; Sahoo et al., 2004) [47, 45] . Such type of feeding provides the source of easily fermentable carbohydrates and nitrogen to the microbes. Buffaloes need to be fed green forages throughout the year to obtain milk in terms of quality as well as quantity. However, in India, as mentioned above there is always a scarcity of green forages. This necessitated the need to conserve the forages as silages or hay when sufficient green forages are available. Outlined the influence of berseem and lucerne silages on feed intake, nutrient digestibility and milk yield in buffaloes with encouraging results. Seasonal forages such as lucerne, berseem, sorghum, barley, oat, maize etc. could be conserved and to be used in lean periods. Lucerne and berseem are the fodders of choice as they are high in protein, minerals and are high yielding nutritious crops. Some studies indicated the positive influence of conserved forages on nutrient intake, digestibility and milk yield in buffaloes.

Reported that high buffering and moisture of berseem and lucerne are detrimental to ensiling and causes heavy nutrient losses. Therefore moisture content should be reduced sufficiently to avoid nutrient losses or one can ensile these high moisture and protein forages with dry forages or by supplementing them with a fermentable carbohydrate source.

Nutrition and reproduction

Reproduction includes growth of foetus and birth of young ones. The proper nutrition is pre-requisite to maintain the reproductive health of the buffaloes and maintain the cyclicity of reproduction. Proper balance of protein, energy, mineral such as Ca, P, Mn, Cu, Zn and vitamins in ration of breedable buffaloes. During first half of the pregnancy, additional allowance rarely required. However, during last trimester of pregnancy, additional allowance required as foetus growth is very fast. As a thumb rule, additional allowance of 0.15 kg of DCP and 0.8 kg of TDN is suffice during last trimester of pregnancy, which can be met by providing good quality concentrate mixture. More average daily gain and reduced age at puberty could be achieved with high level of feeding than low. Feed conversion efficiency at both the levels of feeding is similar in animals. This is also evident from various studies such as in Swamp heifers in Malaysia (Dollah et al., 1989) [17] , Nili-Ravi in Pakistan (Chaudhary et al., 1983; Asghar et al., 1983) [14,4] and Murrah in India (Kaur and Arora, 1989) [28] . However, lower fertility, seasonal estrous behavior and poor estrus, longer calving interval and delayed first calving age are reported in buffaloes (Qureshi et al., 2002) [43] . By nutritional intervention it is possible to improve all these reproductive disorder. It is also reported that feeding calves at higher plane of nutrition i.e. more energy and protein in ration buffalo help to reach age at puberty earlier than those fed on poor plane of nutrition. Supplementation of green forages during reproductive phase of buffaloes is beneficial. Green forages contain carotenes which are precursor of vitamin A which have positive influence on reproductive status of animals. Similarly, supplementation of vitamin A, E and selenium and niacin is reported to be beneficial in dairy buffaloes.

Rumen fermentation

As mentioned above lower supply of proteins causes low reproductive and productive rates in buffaloes. This deteriorates the immunity thus making animals susceptible to various infectious and contagious diseases. In order to improve the productivity, research attention should be diverted to maximize the utilization of these fibrous feed materials by improving rumen fermentation and availability of digestible nutrients at duodenal level.

There is positive correlation between increased rumen fermentation and dry matter digestibility.

Feed utilization in buffaloes vis-à-vis cattle

 Require less DM as compared to cattle.  Utilize DM and Fibre (NDF) better than cattle.  Have more microbial population which extensively degrade feed materials.  Have 10-25 times more oscillospira (protozoa) responsible for the more synthesis of microbial protein.  Have higher number of iodophylic organisms and thus have better utilization of NPN for synthesis of microbial protein which is available to the host animal.  Have higher proportion of butyric and propionic acid than acetic acid as end-product of carbohydrate metabolism.

All above mentioned benefits in buffaloes are due to larger ruminal volume than cattle, slower rate of passage of digesta results in higher recycling of sulphur and nitrogen, better pH stability, higher rate of salivation that maintain buffering capacity of rumen, retention time is more and slower rumen motility thus better mixing of digesta and more time for microbe to degrade feed materials and higher cellulolytic activity as compared to that in cattle rumen (Banerjee, 2009) [5] .

Nutritional management of buffalo calves

After calving, buffalo calves should be fed fresh colostrums (first milk) as specialized cells in lower small intestine of calf are replaced by basal nuclei which are incapable of absorbing immunoglobulins by pinocytosis. This process is known as "gut closure" and it begins as rapidly as 12 hr. after birth if no colostrum is fed. Colostrum is low in fat, high in protein, carbohydrates, vitamin A, K, riboflavin, choline, thiamine and pantothenic acid and easy to digest. It also contains ample amount of copper, iron, magnesium and manganese. Colostrum contains proteins and peptides that have potent biological activity. Most importantly, colostrum contains high levels of immunoglobulins, which transfers passive immunity to the neonates and enables them to fight off infections by increasing immunity. At birth the serum gamma-globulin levels in calves is 0.97 mg/ml which increases to 16.55 mg/ml following first colostrums feeding at 12 hr. to 28.18 mg/ml on subsequent feeding. This level is maintained till the reticuloendothelial system of the calf starts functioning to produce antibodies. Buffalo calf should be fed 2 to 2.5 liters of colostrums daily for the first 3 days of birth. As a thumb rule, colostrums should be fed at the rate of 10% body weight. A practice of not feeding colostrums to buffalo calves within 12-24 h of birth results in poor nutrition and reduced immunity. This makes buffalo calves susceptible to various diseases and even mortality (Wynn et al., 2009) [56] causing heavy losses in the buffalo production and in turn to the economy of the farmer. Proper weaning time is not followed. Male and female buffalo calves are weaned at different periods. Effect of weaning age, energy level, protein level, supplementation of yeast culture on the performance of buffalo calves is reported. To reduce the mortality buffalo calves should be weaned at birth, feeding of colostrum soon after birth should be followed by milk feeding by hand. After 3-4 days of colostrums feeding calf should be provided whole milk and that should be as far as possible from the calf's mother immediately after it is drawn. Milk feeding should be divided in 4-5 equal intervals of the day and should be continued to a week or more then reduce the frequency accordingly. After this milk feeding (app. To control calf mortality it is recommended that calves should be fed colostrums as mentioned above. Some other management strategies include clean and hygienic housing, feeding of antibiotics, feeding of milk replacer, and optimum nutritional care etc.

Bypass nutrient technology

Buffaloes are major milk producing species. However, in India as stated earlier buffaloes are not as productive as their western counterparts. During lactation period requirements for energy and protein for maintenance as well as for milk production exceed the amount of nutrient obtained from diets. Lower density of energy and protein in feed resources for animals such as crop residues and alternate oil cakes are not able to provide optimum nutrition to productive animals resulting in considerable economic losses to the farmer community. Bypass nutrient technology is a tool to provide sufficient energy and protein to the buffalo at intestinal level (post ruminal) by avoiding extensive degradation in rumen. Bypass nutrient technology involves feed management through passive rumen manipulation. Bypass nutrients are not degraded in rumen but get digested and absorbed due to low pH in the lower tract (Yadav and Chaudhary, 2010) [57] . Fat has 2.25 times more energy than carbohydrates and will be available to the animals if it bypasses the rumen otherwise most part of fat will be degraded in rumen. Energy being limiting factor for production, bypass fat thus provides optimum energy availability which can be utilized for milk production in buffaloes. Calcium salt of long chain fatty acids (Ca-LCFA) can be used in ruminants ration as bypass fat to increase the energy density of the ration without adversely affecting the DM intake and digestibility (Naik et al., 2009) [34] . Various oils such as Palm oil, soybean oil etc. is hydrolyzed and reacted with Ca to form its salts that decreases their solubility in rumen (Sukhija and Palmquist, 1990) [49] . It is reported that feeding bypass fat is an alternative to increase energy density intake and digestibility of the diet in cattle (Palmquist and Jenkins, 1982; Hammon et al., 2008) [38,22] resulting in higher milk yield. High acidity in the duodenum accompanied by detergent action of bile acids, lysolecithin and fatty acids causes saturated fatty acids to be more digestible in ruminants than in non-ruminants (Palmquist and Jenkins, 1980) [39] . Feeding of protected fat supplements increased fat, protein content and yield of milk in buffaloes (Gulati et al., 2003) [21] . It is reported that supplementation of bypass fat to buffaloes during early lactation increased milk yield and milk fat by correcting negative energy balance and increased serum triglyceride level (Barley and Baghel, 2009;Garg et al., 2012) [ 6, 19] . Supplementation of bypass fat in diet increases the proportion of unsaturated and long chain fatty acids of milk fat (Garg et al., 2008; Mahecha et al., 2008) [20,32] . Buffaloes growing or lactating demands more protein (amino acids) for tissue synthesis and milk production. Ørskov (1982) [37] and Preston (1986) [40] suggested that bypass protein sustains this higher requirement for productive animals.

Similarly, bypass protein serves as the quality source of protein and amino acids in small intestine. Positive effect of bypass protein on nutrients intake and reproduction, milk yield and its composition in buffaloes is reported (Bharadwaj et al., 2000; Chatterjee and Walli, 2003) [9,13] . Recently, improvement in average daily weight gain, body length, height and heart girth and decreased the cost of feeding per kg live weight gain is in the buffalo heifers reported (Vahora et al., 2012) [53] by feeding combination of bypass protein and fat. Increased digestibility in buffaloes fed rumen bypass protein is attributed to the more availability of nitrogen in diverse form which facilitated the synchronization of carbon and nitrogen units in rumen fluid which positively influence the microbial proliferation that results in more enzyme production (Javaid et al., 2008; Nisa et al., 2008) [35,24] leading to increase rumen fermentation activities producing more volatile fatty acid for productive purpose such as milk yield. Bypass protein increases the ammonia concentration in rumen which increases the rumen pH, as ammonia is alkaline in nature (Nisa et al., 2008) [24] . This pH changed causes the fermentation pattern shift in favour of acetate and butyrate which increased the de novo synthesis of fatty acids which accounts about 60-70% of bovine milk fat resulting in increased milk fat content.

Conclusion

In order to extract maximum production potential from buffaloes it is important to know the underlying cause to improve production, which lies in nutritional management. Thus, it is important to fed proper feeding during varying physiological stages of life of buffaloes. Lactation period and initial life of calves are critical periods in life of buffaloes and needs proper nutritional care.

Conflict of Interest Not available

Financial Support

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