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    WATERBUFFALO NUTRITION

    Luigi Zicarelli1

    RESUMO

    Este trabalho se refere a ingesto de materia seca e da necessidade de energia, proteina, Ca eP dabfala em lactao.

    Assim como na vaca leiteira nos primeiros 50 dias de lactao, a ingesto de materia seca baixa e contudo determina perda de peso. Depois da fase catabolica aumenta a ingesto desubstancia seca e ento observa-se o recupero de peso em torno do 85 dia de lactao. Estaevidencia somente observada se na dieta contem 0,9 MUF x kg de substancia seca.Com taldensidade energetica no final da lactao o animal aumenta de peso, porm uma condionecessaria para se obter um leite com alto teor de gordura (> 8.5%) e proteine (> 4.7%). Umadieta com baixo teor do enegia diminui a produo de leite e o percentual de gordura/proteina

    baixo, sendo assim algumas vezes o leite inadequado a produo de mozzarella. A novilhaaumenta 90kg de peso no final da lactao, atingindo o peso adulto. A bufala adulta que alimentada com uma dieta que contm 0,9 MUF x kg de substancia seca engordam se aproduo inferior a 2700 kg. Ingordam tambm a bufala vazia que apresenta uma lactaomais longa de 270 dias.

    Com uma dieta com 0.9 e com mais de 0.8 MUF/kg x kgde substancia seca respectivamentedepois da desmama e depois dos 6 meses de vida as novilhas vo parir com menos de 30meses. Recordamos que nos paises em que se cria bufalos, as novilhas quando vo parir somais velhas (cerca de 40 meses) que as femeas bovinas.

    SUMMARY

    In this paper we refer on the DM intake and energy, protein, Ca and P requirement of buffalothroughout different phase of productive life.

    As in dairy cows, in the first 50 days of lactation roughly, DM intake is low and entails a weightloss, also present in buffaloes. After these catabolic phase the DM intake increases and therecovery body weight is performed at 85 d of lactation. These performances are obtained withdiets that content 0.9 MUF/kg DM. This energetic density increases the body weight at end oflactation but is necessary for obtaining a milk with high fat (> 8.5%) and protein (> 4.7%)contents. The diet whit low energy decreases milk production and percentages of fat andprotein and is often not able for cheese manufacturing. The primiparous at end of lactationincrease of 90 kg the body weight and reach the body weight of adult subjects. In thepluriparous with a diet that contents 0.9 MUF/kg DM the fattening is present in subjects thatproduce lesser than 2700 kg of the milk and in empty cow that hence present a lactation length

    more than 270 days.

    The diet with 0.9 and more than 0.8 MUF/kg DM respectively after weaning and after 6 monthsold is useful to obtain the first calving at less than 30 months old. We empathize that incountries in which the buffalo species is more present the heifers at first calving are older (40months old ) than bovine.

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    1 Dipartimento di Scienze Zootecniche e Ispezione degli Alimenti DISCIZIA - Universit "Federico II" Napoli. Fax:+39 081 292981; e-mail: [email protected]

    mailto:[email protected]:[email protected]
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    INTRODUCTION

    Most buffaloes (over 90%) are reared in their tropical areas of origin where relative humidityand temperatures (unlikely to fall below 18C) are high, the winter-spring period is dry whilesummer and autumn are usually hot and rainy.

    Some authors (Kamal et al, 1993) assume that buffaloes can develop better than bovine undertropical conditions because buffalo can better acclimate, have a higher resistance in swampzone, and most importantly, have a greater capacity to use forages with high crude fiber (CF)content (Devedra, 1972; Johnson, 1982). These assumptions are not always confirmed bynutritional studies (Abdullah et al, 1989; Raghavan, 1982).

    In particular, research carried out in Italy has shown that only with diets which have a highcontent (35.5%) of ADF (70% forage on DM.) the digestibility coefficient of organic matter andfibrous fractions is greater in buffaloes than in sheep. The above differences between the twospecies are not only cancelled when diets are less ADF-rich (< 24%) but sheep also makebetter use of the CP (Di Lella T., 1997).

    According to Bartocci et al. (2000), when diets have a different forage/concentrate ratio, bovine

    always digest crude fibre better than buffalo despite the latters greater ruminal capacity (DeGirolamo et al.) corresponding to a shorter intestine.

    It is likely that as milk production increases and high nutritive value diets with low CF contentare used, the difference between buffalo and other ruminants regarding gross food digestibilitydecreases.

    The greater numerical increase in buffaloes over bovine in the Tropics is thus due not, as wasclaimed in the past, to the fact that buffaloes digest forage fibrous fractions better althoughtheir welfare is not affected in areas where high temperatures cause rapid deterioration (a fewdays or even in a few hours) in the nutritive value of pastures. In our opinion, the comparisonbetween buffalo and bovine cows must take account of their adaptation to environmentalcharacteristics and other factors such as the coincidence of photoperiod -breeding - peak

    growth of pastures, especially wild forage.In tropical countries north of the Equator, due to the sensitivity to the positive photoperiod, thebovine calving season coincides (Shah, 1990) with shortage of pasture while the dry periodcoincides with forage availability. Shortages chiefly concern the failure to cover the protein andmineral requirement, which means a delayed recovery in the reproductive cycle andconsiderable weight loss.

    The diametrically opposite case occurs in buffalo (species with negative photoperiod) in thatlactation coincides with forage availability (Zicarelli, 1994, Zicarelli, 1997; Zicarelli, 1999a) whilethe dry period coincides with the dry season (Shah, 1990). This enhances the advantage of thebuffalo over the cow.

    In tropical countries south of the Equator, the phenomenon is inverted in that the calving

    season occurs between November and February in the semi-range zeb and between Februaryand May in buffalo, which may use reserves accumulated between November and February.

    The shorter lactation and the longer dry period (one month less and two months more,respectively) allows the buffalo more recovery time and hence the accumulation of bodyreserves which may be used in subsequent lactation (Zicarelli et al., 1977). What stated aboveare likely to hold for both species only given the same dietary conditions and length ofintercalving length.

    In brief, if we consider the average chemical composition of pasture during the year in tropicalareas, due to the high energy/protein ratio, forage in such areas satisfy buffalo requirementsbetter than those of bovine. Given the same energy output with milk diets with protein deficitscause a greater weight loss in bovine than in buffalo. The corrected buffalo milk (ECM) in factusing the formula ((((((fat (g) - 40)+(protein (g) -31))*0.01155)+1)*kg of milk) of Di Palo(1992) has a lower protein (28.3 vs. 31.0 g, respectively) and P (0.9 vs. 0.7) content thanbovine 4% FCM (tab. 1). On the basis of the same energy content (ECM) Brazilian buffalo milk

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    shows higher protein and lactose percentage values, that are in any case lower than bovinemilk. Fat percentage is higher than bovine milk like ECM Italian buffalo milk.

    Table 1. Bovine and buffalo milk composition and nutritional requirements.

    Bovine Buffalo

    4% FCM Milk Equivalent

    corrected milk ECM

    Milk composition ITALY BRASIL ITALY BRASIL

    kcal /kg 740 1227 1066 740 740

    MJ /kg 3,13 5,19 4,51 3,13 3,13

    proteins (g/kg) 31 47 42 28,3 29,16

    lipids (g/kg) 40 83 68 50 47,27

    lactose (g/kg) 49 49 50 29,6 34,85

    Ca (g/kg) 1,1 2 1,2

    P (g/kg) 0,9 1,2 0,7kcal/g proteins 23,9 26,1 25,4 26,1 25,4

    CP/Ca 28 23,5 23,5

    Nutritional requirements/ kg of milk

    CP (g) 85 129 115 78 80

    PDI * (g) 48 73 65 44 45

    MUF** 0,44 0,73 0,63 0,44 0,44

    NEL (MJ) 3,13 5,19 4,51 3,13 3,13

    Ca (g) 3,5 6,36 3,84

    P (g) 1,7 2,27 1,37*PDI = True Protein truly digestible in the small intestine.

    ** MUF = NEL 1700 kcal o/or NEl 7.11 MJ

    DRY MATTER INTAKE

    In order to perform a correct rationing it is important to know the voluntary dry matter intake.

    Data concerning dry matter intake in dairy buffaloes are somewhat conflicting. This is due tothe fact that almost always the intake level is not referred to the days in milk, productive leveland live weight of the buffaloes and to the level of fibrous fractions which regulate dry matterintake or the BCS at the beginning of the observation period.

    We should not therefore be surprised that alongside 2.2 - 2.6% intake levels (Ranjhan, 1992;Bertoni, 1992; Verna et al., 1993) we find others equal to 2.7 - 3.4% (Di Palo, 1992) and 2.9 -

    3.1% of live weight (Di Lella, 1998).In trials carried out by Campanile et al. (1997a 1997b) it emerged that the intake wasconditioned negatively by the content in the cellular walls (ADF, NDF, ADL) and positively bymilk production. With diets characterised by 27 - 28 % ADF, 49 - 45% NDF and 46 - 55%forage on DM, dry matter intake per kg of ECM amounts to 275 g in addition to what requiredfor the maintenance (91 g per kg of metabolic weight). Despite a greater presence of cellularwalls, the above DM intake values, in relation to the ECM yield, do not deviate much from thosenormally recorded in dairy cows. Unlike the latter, the buffalo ingests more quickly, thanks tobetter developed incisive and ruminates for longer (Campanile et al., 1977b). This allows it tofeed on more bulky diets which undergo a higher rumination time.

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    In a recent trial (tab. 2) carried out on 390 primiparous producing 2.325 kg of milk in 270 dayswe have found that the dry matter intake (DM), the percentage of DM/L.W. (fig. 1a) andDM/ECM is related to the days in milk (DIM) and the milk production (fig. 1): in fact theyincrease from the delivery until 121 - 150 DIM and decrease successfully, while ECM productiondrops after 61 - 90 days. It is possible to distinguish, like bovine, a catabolic phase ( until 55days) and an anabolic phase. Through out the catabolic phase the energy intake is less than

    the requirement and the cow utilize the body reserve for the production. During the anabolicphase occurs the opposite: the body reserve are reconstituted and the body weight increases.

    Table 2. Dry matter intake (DM kg), ECM production at different days in milk (DIM),percentage of DM on live weight (L.W.), DM and MUF intake for kg ECM, live weightand coefficient correlation (r) in primiparous.

    DIM D.M. ECM % L. W. D.M./ECM MUF/ECMLive

    weight

    1 5301 - 30 12.29 14.61 2.34 0.08 0.40 523

    31 - 60 13.50 17.98 2.60 0.15 0.37 52061 - 90 16.72 17.85 3.18 0.32 0.54 53491 - 120 17.27 16.87 3.19 0.34 0.59 551121 - 150 17.34 16.41 3.10 0.32 0.60 567151 - 180 16.17 14.31 2.82 0.26 0.61 576181 - 210 15.22 12.84 2.63 0.20 0.61 583211 - 240 15.82 11.53 2.68 0.24 0.72 598241 - 270 16.15 9.28 2.66 0.29 0.96 619271 - 300 11.61 6.41 1.88 -0.38 0.67 612301 - 330 13.06 5.62 2.13 -0.16 1.07 615331 - 347 13.81 4.33 2.23 -0.04 1.48 621

    0 - 60 12.89 16.30 2.47 0.12 0.39 52361 - 150 17.11 17.04 3.16 0.33 0.58 542151 - 270 15.84 11.99 2.70 0.25 0.73 588271 - 347 12.83 5.45 2.08 -0.19 1.07 61661 -210 16.54 15.66 2.98 0.29 0.59 556

    r coefficients

    dim -0.158 -0.880 -0.453 -0.370 0.670 0.976DM kg 0.430 0.944 0.882 0.258 -0.174ECM kg 0.667 0.509 -0.817 -0.883

    % DM/L.W. 0.884 -0.101 -0.480DM/ECM 0.224 -0.366

    MUF/ECM 0.872

    We underlain that the diet of above trial have 0,9, 15,7%, 38,5% and 50% respectively ofMUF/kg D.M., Crude Protein, N.D.F. and roughages/concentrate. The diet cause the energydeficiency until 55 DIM but allow to recovery body weight at 85 days DIM and reach the adultweight at end of lactation (tab. 2, fig. 1).

    The above observations, confirmed repeatedly in the field, allow us to estimate, at leasttheoretically, dry matter intake in the herd. With diets consisting of lower cellular wall contents,intake levels are little changed and in some cases are lower, especially when the increase inenergy density is obtained with highly fermentable foodstuffs which involve an increase inglycemia. Glycemic values above 70 mg/100 ml are likely to be detrimental on the appestat.

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    In the diet used in the above trial when the differences between the density energy of DM (0,9MUF/DM) intake vs. energetic density of DM required or MUF/DM are higher than 0.6 and 0.2respectively, the dry matter intake for kg of ECM decreases and becomes negative after 270days of lactation (fig. 2). In other words, it is production itself and the energetic density of DMwhich regulates intake.

    As with dairy cows, in the early lactation (roughly in the first 55 days) DM intake is low andentails a weight loss also in buffaloes. In our opinion this phenomenon is an ancestralthrowback to life in the wild which has become a behavioural characteristic that is observedmore or less markedly in all domestic animals. Thanks to the above behaviour, species have

    been able to survive: to be able to protect their offspring from predators, it was necessary torestrict their movements and thus to search for their food just when post-partum nutritionalrequirements increased day by day. Over time, this has favoured the survival of thoseindividuals able to activate catabolic processes more efficiently, which maximise the use of bodyreserves. In selecting subjects with greater milk-yield capacity, man indirectly chose animalsthat managed to better use their reserves accumulated during the dry period.

    This concept is extremely important to be able to perform rationing which takes account of thecharacteristics peculiar to each species. The high-producing dairy cow, for example, is able toactivate the above catabolic mechanisms efficiently. Indeed, it is well known that in the periodin which the intake capacity is limited, the dairy cow can draw on its reserves, losing more than1 kg in weight per day, and cope with a 20-30% deficit in its requirements without penalizingproduction. In the buffalo, the catabolic capacity is lower and, where requirements are not

    covered, this entails a reduction in milk production due to deficits which the high-producingbovine is able to tolerate much better. In other words, the buffalo possesses a catabolichabitus which is usually less pronounced than that of the high-producing dairy cow. Its lowerlipid metabolism designed to store reserves for the periods of low forage availability (typical ofanimals originating in extremely hot or cold climates), seems unlikely to favour its milk-producing capacity to the detriment of its welfare. However, it should be added that buffaloesproducing over 40 q of milk possess more enhanced catabolic capacities and, given the samediet, are distinguished within the group by their lower BCS values.

    However, much remains to be ascertained on dry matter intake: inexplicably, given the sameseason, the same yield, live weight, number of primiparous, calving interval and dietarycharacteristics, field observations show that some herds are particularly voracious while others,albeit more productive, have moderate intake levels. This often stems from the fact that

    consumption also depends on previous feeding regimes which cannot always be ascertained:

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    subjects are led to modify their intake in order to reach a body score which is more in keepingwith the physiological phase in question.

    Therefore it is always worth ascertaining the herds actual intake capacity so as to supply thecorrect dry matter composition. As regards the season, it should be pointed out that, unlikewhat occurs in cows milk, summer temperatures do not have a markedly negative effect on theintake level provided buffaloes have shade and pools at their disposal (fig. 3; unpublisheddata). From these observations spread out that the low daily temperatures decrease the drymatter intake.

    Mean calving distance of the herd, however, merits separate treatment: it varies during theyear both in out-of-season herds and in those characterised by typical seasonality. In theformer case, calving distance is on average 200 days in January, decreasing to 100 days in Julyand subsequently lengthening until December, a month in which the herd once again reaches,and in many cases exceeds, 200 days. In the latter case, highest values occur between Juneand July while lowest values between January and February (Zicarelli, 1999).

    Average milk yield per head and milk chemical composition thus continually vary in herds withgood fertility and a concentration of primiparous calving into only two months. In these herdsdry matter intake continually changes and feed characteristics often need to be modified.

    However, there are very few herds with a homogeneous calving distance during the year, asoccurs with dairy cow herds.

    Buffalo herds with this characteristic are those in which there is a high incidence of primiparousor where there is a more hypofertility: the average calving distance is longer, milk chemicalcomposition is more homogeneous during the year but production tends towards low levels.

    ENERGY REQUIREMENTS

    A) MAINTENANCE

    In the practice of rationing of Italian buffalo, for maintenance we use the requirements

    suggested by the French for dairy cows, that is MUF 1.4 + MUF 0.6 per 100 kg of live weight

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    (Zicarelli L., 1990, Proto, 1993, Di Lella T., 1997). These values have to be increased by aquantity of MUF that depends

    1. on the energy required for walking which varies according to the space available;

    2. the hierarchy existing in the groups and hence competitiveness and aggressivenesswhich some animals use over others (the greater the age difference in the herd, the

    more accentuated this will be);

    3. on the use or otherwise of pasture feeding;

    4. management and hence the man-animal relationship.

    In this regard, according to our experience, the less space is available and the greater thenumber of hours with empty feeding troughs, the greater is the herd's aggressiveness. Hence,rations which do not theoretically cause energy waste have the indirect effect of increasingrequirements inasmuch as the group becomes "more lively". This is especially true whenbuffaloes are not dehorned, as happens in Italy.

    Such factors are difficult to quantify although, to be on the safe side, energy supplements of atleast 20% more than maintenance requirement, are suggested. Another important factor is the

    nutritional state of the subjects (BCS): at the same weight, the higher is the animals BCS, thegreater is the maintenance requirement, in that in the composition of live weight, fatspredominate.

    Thermoregulation requirements take on an important role for the buffalo. Its tropical originmakes it more sensitive to low than to high temperatures. Its capacity to thermoregulate duringwinter in Italy, especially in colder areas, entails considerable variations in the endocrinesystem. This is confirmed by higher hematic values of the thyroid hormones recorded in winter.In this season, the buffaloes raised in the mountain foothills of Campania (Campanile et al.1993) show higher hematic values of thyroid hormones compared with those in coastal areas.

    At the beginning of colder spells, when animals have not yet adapted to low temperatures,abnormal titratable acidity values (SH) are often recorded in milk, above all when subjects areexposed to wind and energy requirements are not covered.

    From these remarks, it is evident that in some environments, sound design of cowsheds isimportant, without which there might be adverse effects on the quality and quantity of milkproduction. In tropical areas where the climate is more favourable to the buffalos endocrinesystem, we believe that such phenomena occur less often.

    At the start of the cold season, almost always the rather acrid smell of acetone is noted, whichtestifies the intense catabolism of the stored fats to cope with the increased requirementscaused by temperatures which the species finds uncongenial. Such phenomena are not alwaysaffected by the lactation phase: increases are often observed in the hematic level of -hydroxybutyrrate (Bertoni et. al., 1993; Campanile et al., 1995) even in the final phase duringthe transition from two daily milkings to one. The phenomenon is more intense in herds inwhich the transition is accompanied by a reduction in the nutritive value of the diet.

    Evidently, even when requirements are low, the high milk fat content and reduction in intakecapacity (fig. 2), observations of the terminal phase of the lactation curve, suggest soundformulation of the diet.

    B) MILK PRODUCTION

    Irrespective of the buffalos capacity to use different forage types, it must be acknowledgedthat the net energy required to ensure production of 1 kg of standard milk (740 kcal) is equal tothe caloric value of the milk produced.

    However, field trials (Di Palo, 1992, Bertoni et al., 1993, Di Lella, 1997, Di Lella, 1998) haveshown that about 0.5 MUF are required to obtain 1 kg of ECM. This figure is understandableinasmuch as it includes, if calculated in relation to energy consumption of a group:

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    1. the recovery and possible increase in the animals weight;

    2. the energy surplus which is required to cope with various environmental conditions(climate, more accentuated hierarchical factors in some herds compared with others,etc.), the fattening of hypofertile animals or those with protracted low productivelevels;

    3. the feeding of bulls in the herd. The latter usually account for at least 4% of the group.

    In dairy cows, standard milk has a quantity of lipids, proteins and lactose equal to 4.0%, 3.1%and 4.9% respectively. With regard to the above values, we encounter oscillations whichdepend on the genetic, the lactation phase, the type of diet and so on. Lipids may vary, forexample in the Italian Friesian, from 2.8% to 4.2% and proteins from 2.7% to 3.5%. In thebuffalo milk, the fat percentage may oscillate from 5.5% to 13% and proteins from 3.8% to5.5%. Therefore the energy requirement for a kg of cows milk varies from 0.35 MUF to 0.45MUF (= 28.6%) while in buffalo (tab. 5) it is between 0.53 MUF and 1.02 MUF (= 92.5%).

    The greater variability found in the buffalo entails rationing difficulty: the buffaloes which areable to produce milk rich in fat use their energy surpluses for this purpose while those withoutthe above capacity accumulate body reserves. Often in the former, any energy deficit fails to

    result in the production of milk with a lower fat content while determines a lower milkproduction. The same phenomenon is recorded for other nutrients: if not enough precursors aresupplied to the animals to produce a secretion with almost fixed characteristics, the productionwill lower with little change in its composition.

    Given that in the dairy cow the increase in the milk fat content usually does not exceed 29%while in buffalo it often go above 92%.

    Then excess rationing leads to fattening in the cow (fat cow syndrome) rather than the buffalo,which uses the excess more beneficially to modify milk chemical composition and, in certainways, limits the amount of waste. However, with the same diet, it may be noted that the mostproductive buffaloes in the group are thinner than the least productive.

    All this may give rise to some perplexities, given that, at least in Italian livestock farming, it is

    easy to observe excessively fat buffaloes. However, it is worth pointing out that theconformation of a buffalo differs from that of a dairy cow. A fat cow which has a BCS of 5 isfatter than a buffalo which is given the same rating. Indeed, if we believe that a cow is at itsbest when it has a BCS equal to 3.5, the same cannot be said for a buffalo with the same scoresince the constitution (the constitutional habits used by old endocrinologists) of the species isdifferent. Indeed, the welfare of a dairy cow with a chiefly catabolic habits cannot beidentified with that of a buffalo in which selection for milk production is still in its early stages.Moreover, we should consider that under the same BCS, which evaluates the roundness of theshape, a buffalo is thinner than a cow. Given the same subcutaneous fat deposition, the formerhas less lipid infiltration into the muscles. This is another reason why we prefer to use the 1 to9 scale which is more suited to animals for meat production.

    C) WEIGHT RECOVERY

    The weight loss which occurs in the first phase of lactation has to be made good in subsequentmonths. The amount of body weight loss varies with the energy density of the diet fed to theherd according to its productive level and the weight of the animals. As stated above, theweight loss that a buffalo can sustain without compromising the persistence of the lactationcurve is, owing to its moderately catabolic habits, lower than that of cows. In any case, weightlosses also occur in buffalo and they should be offset, bearing in mind that if the catabolism of1 kg of body weight supplies energy equivalent to 3.5 MUF, for its reconstitution 4.5 MUF arerequired in the anabolic phase of lactation.

    Further proof of the actual weight loss of the buffalo in the early stages of lactation is given bythe fact that, while the MUF required to produce a kg of ECM milk amounts to 0.35 in the first

    40 days (Di Palo R., 1992), it is over 0.5 during the whole of lactation. This demonstrates thatthe difference existing between 0.35 and 0.44 in the first 40 days represents the amount ofstored energy that the buffalo makes available for udder synthesis to overcome the gap

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    between energy intake with the diet and that synthesised in milk. It can be easily shown that ifthe buffaloes of a livestock farm in the first 40 days produce an average 12 kg of milk, theweight loss will be at least 300 g per day (MUF 0.44 0.35 = 0.09; kg 12 x 0.09 = MUF1.08/3.5 = g 0.309).

    D) GROWTH OF PRIMIPAROUS

    Late puberty and age at first calving are a feature of the buffalo. Indeed, in Italy, as in manyother countries, the first calving occurs at about 41 months, compared with 30 months inbovine (in the Friesian or in the Regian the age is brought forward to 29 months). This suggeststhat the buffalo is actually less precocious than the bovine; this is further confirmed by thepresence of gingival mucus which covers the incisors at birth, despite gestation lasting over 10months. The late age at which the species reaches puberty, in our opinion, is considerablyaffected by the raising technique used. On farms where weaning is carried out at a weightexceeding 90 kg and appropriate rationing is adopted (Italy) or natural suckling is practised andextensive pasturage is available (Brazil), the first calving occurs between 28 and 33 months andin many cases at 24-28 months (independently of whether the out-of-season technique isadopted, which postpones the first conception in pubertal animals which are separated from the

    male at the beginning of the summer). Recently we found that the use of feeding programmesfrom weaning, with an energy density equal to/higher than 0.8 MUF/kg DM and a CP DMpercentage of 12.5%, recorded after one, two and three years of adoption a first calving agebelow three years respectively in 20%, 42.2% and 65.3% of animals.

    It was confirmed that if weaning and growth in the prepubertal phase are not optimal,corrective interventions carried out after 18 months are unable to maintain the start of pubertybelow three years (7). This shows that the species is unable to rapidly recover the damageexperienced in the first growth phase. It should be noted that, unlike what happens in the dairycow thanks to the effect of selection, age at first calving in the Italian Mediterranean Buffaloand in other buffalo breeds/populations is still very variable. Indeed, the weight of adultfemales varies from 550 to 900 kg, which also indirectly conditions the pubertal age.

    In rationing of primiparous there should be a surplus for the growth. Normally a buffalo whichweighs 650 kg when adult, weighs about 550 kg at the first calving. During the first lactation, itshould theoretically grow by 1/3 of a kg per day, for which a daily energy supplement 1.166MUF (4.5 x 0.333 g/die) is required. This value, borrowed from the requirements of primiparousbovine of dairy breeds, does not take account of the optimal conformation of the primiparousbuffalo, which has a higher fat/muscle ratio. Hence, until in-depth research proves the opposite,it is worth considering a surplus of 1.5 MUF per day (4.5 x 1/3) in the rationing. Moreover, itshould be borne in mind that the age at first calving of the Italian Mediterranean buffalo in1999 (AIA data) was 3 years, 4 months and 4 days, hence on average 8 - 10 months behindthe italian dairy cow. As age increases, food conversion efficiency is known to lower; hence theenergy required for weight increase after the first calving in the buffalo must be held to behigher than that attributed to the dairy cow until further experiments will be carried out.

    The proposed energy supplement should obviously be added to requirements, and in herdswhere there is no age-based separation of animals, the above energy surplus should be addedto the group ration taking due account of the number of first calving buffaloes.

    ENERGY RATIONING

    As we have so far referred theoretical aspects, we should now deal with the field application ofsatisfying energy requirements. If technology is not used (transponders), in field tests grouprationing is implemented usually by using - at least in Italy - the total mixed ration.

    As stated above, at least in southern Italy, diets for buffaloes are formulated withapproximately 50% of DM consisting of forage and an NDF percentage exceeding 40%.Depending on such factors, the quantity of energy administered may be varied. It may be

    diluted or concentrated in actual quantities of dry matter ingested by the group, always taking

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    account of the herds BCS. The above tables allow us to calculate the ration for standard dailyproduction.

    As dry matter consumption increases, the energy density of the diet rises. As production rises,the energy density of the diet necessarily increases, also because there is a limit to the DMintake capacity insofar as it cannot exceed the quantities that are conditional upon live weight.

    Thus the increase in the herds production level is possible if, as the genetic potential increases,there is a rise in animal size and their capacity to transform feed into milk.

    In support of the data referred to above, we report (fig. 4) the pattern of milk production perday and the fat percentage according to the DIM recorded on data collected over three yearson the farm which is linked to our department, where animals produce more than 3 thousandkg of milk per lactation and fed a diet with 0.93 MUF/kg DM. As milk production is reduced, thefat content rises and, conversely, the latter decreases as production rises (especially between90 and 100 days from calving when the herd exceeds 14 kg of milk). This prove that in theabove lactation phase, even when the animals receive a diet of 0.93 MUF/kg DM, they cannotcover their requirements. Indeed, the milk fat content is lower than the average of the wholelactation (8.73%).

    After the recovery phase (over 150 days of lactation), the animals ingest above theirrequirements and tend to increase body reserves to an extent which is proportionally inverse toproduction. In this phase, it is necessary to lower the energy density of the diet and drasticallyreduce the percentage of NSC, especially starch, which should be kept below 18% as it is themain cause of fattening and shortening of lactation. It is advisable to move less productiveanimals and feed them a diet with more roughage and/or cellular walls. This strategy which wehave employed in the past was not very beneficial in that the buffalo showed great sensitivity tothese changes both due to hierarchical factors and because it responds more negatively thanthe dairy cow to nutritive restriction. Indeed, it produce milk losses which, economicallyspeaking, exceed the benefit derived from nutritional cost savings (Campanile et al 1996). Whathappens in Italy, where a kg of milk is worth approximately six times the price of MUF, mightnot hold for other areas where the above relation is different.

    The fattening of animals is therefore in economic terms a necessary evil insofar as the buffalo,

    having not yet acquired a milk-bearing habits, manages to persist in lactation only if it receivesa diet greater than its own real needs. Excessive fat deposition occurs markedly in animalswhich extend lactation beyond 270 days due to fertility problems (it is hypofertility, at least in

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    pluriparous, which fattens animals and not excessive fatness which causes hypofertility) or inless productive animals. In others, fattening is less evident and the waste during lactation mayeffectively be recovered in the dry period if two groups are set up according to the BCS reachedat the end of lactation.

    However, feeding restriction applied to fatter subjects during the dry period should preservesome of the fat (BCS 3.5 - 4) accumulated during lactation in that this is most effectivelymobilised after calving. Thus the economic waste occurring in lactation would be recovered,while protein, mineral and vitamin requirements would be covered.

    The energy density suggested should come as no surprise as buffalo ingest a quantity of dry

    matter which is proportional to their production in corrected milk and their live weight.Fattening occurs if the diet is particularly rich in starch or there is an imbalance between starchand protein intake. In the past, buffaloes were definitely thinner, had lower glycemia andcholesterolemia, but were less productive and gave milk with a different chemical composition.In 1978, the year of the first functional control, the average milk production amounted 1641 kgbut fat content was 6.42%. In several trials carried out in DISCIZIA (figg. 5 and 6) it is possibleto observe the positive trend among the years of fat and protein contents of buffalo milk inItaly.

    PROTEIN REQUIREMENTS

    In the first part of this paper we showed that, due to its milk characteristics, the buffalos

    protein requirement, given the same energy (740 kcal) produced (ECM), is lower than that ofthe cow (tab. 1).

    This holds under extreme conditions of survival and the buffalo, thanks to efficient urearecycling, can cope with protein deficiency. What has stated above cannot always beextrapolated in the practice. One of the characteristics which marks the buffalo is that proteindegradation in the rumen occurs much more quickly than in the cow (Terramoccia et al., 2000).Compared with the latter, the feed stays longer in the rumen but less time in the intestine(Bartocci et al., 1997) This ensures that few proteins bypass the rumen or may be directly used(absorbed) in the intestine (Di Lella et al., 1995). However, this is not detrimental, as it is in thecow, in terms of damage from protein excesses. In spring, buffaloes at pasturage in Italy ofteningest even 30% more protein than is required, without evidencing the imbalance which occursin cows (damage to hooves, mastitis, alkalosis and hypofertility). In a recent experience

    (unpublished) we found that two buffaloes groups undergone A.I. and fed with different proteinpercentage on D.M. show the same fertility.

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    In the light of these considerations, we believe that it is difficult to specify the buffalos proteinrequirements in that, unlike other ruminants, the calculation becomes complicated by thebuffalos metabolic state and especially its capacity to save available nitrogen.

    In the field rationing, until a truly reliable method can be set up, we believe that in order toestimate requirements the use of CP, which has provided acceptable results to date, issufficiently valid. We suggest a quantity of degradable proteins in the diet at less than 70%when the dietary protein content is less than 14% of DM and even at lower percentage than67-64% for more protein-rich diets.

    In rationing, we consider a maintenance requirement of 700 g of CP for a 650 kg buffalo, whichis equivalent to about 400 gr of PDI of the French system (3.25 g of PDI x kg of P.M.). Forweight recovery we estimate, again according to the French system, a PDI requirement of 300gr which amounts to roughly 500 g of CP to be added to other requirements (177 g of CP x 333g of live weight gain/die). As regards milk, because of its variability in protein content, weprefer to consider a requirement of 2.742 g of CP per gram of protein synthesised in milk sincefor cows 85 g of CP are advised per kg of milk with 3.1% protein (85/31 = 2.742).

    Also in the case of protein rationing, the same applies as in energy rationing. As productiondecreases, the percentage of CP on DM should decrease. This would suggest that, with falling

    production, animals should be moved to a group subjected to a lower energy and protein diet.This technique entails milk losses which fail to compensate the saving achieved by rationing.

    CONSIDERATIONS ABOUT PROTEIN RATIONING

    As regards protein rationing, it should be remembered that besides having a plastic function,proteins are also likely to have metabolic effects as yet unascertained. In our experience,buffaloes exceeded a production per die of 20 kg only when the amount of protein in DMexceeded 15%, without there being any side effects upon the state of health of less productiveindividuals.

    At the beginning of lactation, due to lower intake levels, we suggest increasing the protein

    supply by 10%, on the basis of trials carried out by our department. Buffaloes which produceover 20 kg ECM and ingest diets with 13.5% CP/DM have low values of azotemia between the11th and 32nd day post-partum. These values rise between 70 and 110 days when dry matterintake increases. These data suggest that between the day 11 and 70 post-partum, DM intakecould vary between 13 and 16 kg and hence the protein requirements are not met if the dietaryCP is less than 13.5% (Campanile et al., 1995).

    Table 3. Live weight, MUF and crude protein per kg of dry matter for different production in270 days in order to cover the requirement s for milk, live weight and recovery bodyweight in primiparous buffaloes.

    Live weight MUF per kg DM Crude protein (g) per kg DMq buffalo milk (270

    days) 23.3 30 40 23.3 30 40 23 30 40DIM

    1 530 530 5301 -30 526 532 519 0.97 0.89 0.97 150 133 15031 -60 514 526 497 0.99 0.94 1.04 156 153 17461 - 90 516 519 474 0.86 1.01 1.09 133 152 18491 - 120 530 514 454 0.84 1.03 1.11 127 146 181121 - 150 549 509 434 0.75 0.94 1.03 114 145 177151 - 180 568 508 424 0.75 0.90 0.99 112 132 168181 - 210 582 511 418 0.76 0.88 0.96 112 126 153211-240 599 515 414 0.70 0.86 0.93 101 120 143241-270 623 523 415 0.63 0.82 0.89 88 110 128

    271-300 646 536 424 0.77 0.76 0.81 103 96 107301-330 655 552 436 0.66 0.73 0.78 87 92 101331-347 674 569 450 0.58 0.70 0.74 75 85 93

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    On farms which administer higher protein concentrations which we consider more suitable formeeting daily protein requirements at the beginning of lactation, low azotemia does not occur(Campanile et al., 1997c and 1997d). The greater availability of keto acids, which usually arisefrom protein metabolism, favours glucose synthesis. Glycemia is therefore higher in subjects fedwith protein excess. High glucose and low insulin values guarantee the udder a greateravailability of glucose for lactose synthesis. The metabolic processes triggered in subjects with

    protein excess thus contribute to reducing negative glucose output at the beginning oflactation. Spires and Clark (1979) have noted that high levels of NH3 in steers cause poor useof glucose, contributing to rapid glycogenolysis.

    On the basis of the above considerations for energy, protein and recovery body weightrequirements we also referred (tab. 3) the live weight trend of primiparous buffaloes producing23.3 q of buffalo milk, fed with 0.9 MUF per kg of D.M throughout 270 days of lactation.Furthermore we report the theoretical live weight of primiparous buffaloes characterized byproductions of 30 and 40 q and the energy density and the grams of CP per kg of DM needed inorder to satisfy the requirements for 23.3, 30 and 40 q of buffalo milk in 270 days of lactation.

    E) CA AND P REQUIREMENTS

    Buffalo milk has 1.8 2.0 g of Ca and 1.1 1.2 g of P per kg.

    As regards maintenance, see the bovine requirements of INRA (Jarrige, 1988). For milkproduction we calculate requirements taking into account that while for a kg of cows milk 3.5 gof Ca are required (1.2 g/kg of milk), buffalo milk will require 5.25 - 5,8. The same holds for Pwhose requirement is 1.7 (0.9 g/kg of milk) for cows, equivalent to 2.1 2.3 g for buffaloes.

    The quantities of Ca and P in the diet must be proportional to the energy and protein contentso as to cover production requirements assured by the latters.

    Mineral dietary supplementation is particularly important in the dry period. Due to dietarycharacteristics in Italy, Ca and/or P deficiencies often occur (Zicarelli et al., 1982). P deficiencyis one of the most frequent causes of vaginal and/or uterine prolapse in this species. Indeed,

    the buffalos long dry period compared with that of the dairy cow amplify any rationing errors,however small, made during this period. A daily 10-gram deficit of P or Ca causes, in a 120-daydry period, a reduction of 1,200 g which is equivalent to 24% of P and 13.3% Ca contained inthe bones.

    It is always worth knowing the percentage of the two minerals in forage so as to ensure at least45 g of Ca and 47 g of P. Ca excesses during the dry period cause parathyroid hypoactivitywhich leads at birth to the persistence of low calcemia values. In the dairy cow this causespuerperal collapse sometimes accompanied by uterus prolapse. In the buffalo, only the latterevent occurs with greater frequency (Zicarelli, 2000).

    GENERAL COMMENTS ON RATIONING

    We should take account of the fact that in the herd are animals with different potentials andthat, as production rises, those with greater milk-producing aptitudes are more heavilypenalized when DM intake corresponds to the theoretical intake. In formulating rations thetechnician will evaluate the cost/benefit ratio and percentage of possible high yielding animalsin the herd.

    From our experience, the subdivision of animals into several production groups in which torotate the animals or the adoption of artifices, which complicate the management of herds withover 150 adults, solves the problem of waste but also penalizes the herds productivity.

    The best solution would be to subdivide animals into several groups after calving and fed thema uniform diet throughout lactation according to their different productive potential (EBM).

    Adoption of this technique promotes the accumulation of reserves only in hypofertile animals

    which, their lactation being longer, will spend more time at a higher nutritional level.Subdivision of animals in the dry period into at least two groups in which to adopt a diet

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    according to the degree of fattening reached at the end of lactation is, in our opinion, thesoundest practice.

    As stated above, if the ration does not allow the buffalo to synthesizes milk with a fat contentconsonant with its genotype and lactation phase, production is not optimal.

    In terms of energy requirements, the buffalo aims to match its milk fat content to its lactation

    phase. With an energy deficiency, there is little change in the fat percentage and productiondeclines. However, it is more likely that as it cannot produce milk with the characteristics of itslactation phase, both the fat content and production decline. Clearly other factors come to playin this mechanism, not least of which is the interaction between genotype and rationingcharacteristics.

    Figures 7 and 8 shows the trend in production and milk fat content in two successive years(1997 and 1998) for buffaloes raised on two farms. One of the farms keeps the buffaloes in asingle lactation group (az. A) up to a production of 4 - 5 kg and the other (az. B) moves animalsinto several groups as production varies.

    The data were obtained from the functional controls conducted by APA (Provincial FarmerAssociation). In both years, the buffaloes in Farm A showed a constant increase in fat content

    according to calving distance, while those of Farm B delayed the above increase to the end oflactation. The persistence of milk production, irrespective of the genetic level of the two herds,is decidedly more favourable in Farm A buffaloes. By contrast, those of Farm B show a similartrend in production and milk content to that of buffaloes raised in Campania in the 1970s orthat of Brazilian buffaloes kept wild which are affected by seasonal pasture production (Macedoet al., 1997). Comparison between the two farms indirectly supplies the evidence for ourassertion.

    In rationing, it is essential to reconcile theoretical and actual needs, ascertaining the herds BCSand dry matter intake in particular.

    In buffaloes in mid-lactation (132 days) we noted (tab. 4) that rationing deficits in CP (-30%)and/or PDI (-21%) altered the cryoscopic index. Restoration of the protein balance led tonormal cryoscopic values in a few days and also increased milk protein content (Campanile et

    al., 1998).

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    formulated with a higher protein content (12%) and a lower NSC/CP ratio (3.04 and 2.69respectively at 132 and 164 days).

    At the beginning of both trials, besides the changed cryoscopic index, we noted a blood ureavalue higher than theoretically found according to diet protein contents. In other words, after aprolonged period of protein deficit, circulating urea remained at such levels as not to suggestthe above deficit. Moreover, after the protein percentage in the diet was increased, urea valueswere no different from starting values. However, a clear decrease in azotemia (and urea in themilk at the same time) was observed when there was a shift from

    normal to low protein content in the diet.

    These results suggest that the failure to cover protein requirements and/or long-term diets witha low protein content, especially those with a high NSC/CP ratio, do not cause low azotemia ormilk urea values but alter the cryoscopic index.

    However, the urea level in the milk and blood only declines after abrupt falls in dietary proteincontent, causing abnormal values in the cryoscopic index. As the latter is used for milkpayment, it is better in formulating rations not to use protein contents lower than 12% evenwhen lower values would ensure the coverage of requirements.

    Urea content in blood and milk is affected by the NSC/CP ratio. Where there is a larger quantityof fermentable energy available for ruminal microflora, the ammonia which forms may be moreeffectively transformed into microbial proteins. This leads to a lower production of urea (Journetet al., 1975). In the trial described, a low dietary protein content and high NSC content shouldhave produced even lower urea values in biological liquids. This did not occur; clearly, in buffaloa mechanism is triggered which allows them to make full use of protein sources during periodsof deficiency and enables them to survive the dry seasons in difficult tropical environments.

    Where there has been long-term intake at low protein concentrations, by reducing theaminoacid tissue turnover the buffalo is able to maximise use of nitrogen sources, which areemployed for bacterial production through an excellent use of fermentable organic material(Bertoni et al., 1993). Moreover, the buffalo is able to transform nitrogen sources into proteins,even under energy deficiency, and more effectively than bovine are able to do (Langer, 1969)

    in that it has a ruminal environment which is more suitable to the growth of microorganismswhich use non-protein nitrogen.

    Lastly, compared with the dairy cow, the buffalo can adapt better to protein deficiency (Bertoniet al., 1993) even when the energy/protein ratio is not optimal.

    Indirect evidence is provided by the paucity or absence of negative effects of hyperprotein dietson integrity of hooves and on buffalo welfare, which occur under current practices withberseem-rich diets in tropical countries North of the equator or when, in springtime, the freshforage is used in some southern Italian plains.

    A lowering of urea blood levels, following the reduction of protein concentration, occurred withdiets characterised by high levels of fermentable energy. The diets in question may have led toa rise in insulinemia and thus reduced or blocked the normal breakdown of aminoacids, therebycontributing to reduce the level of circulating urea.

    We subsequently conducted other studies though the data were not examined in such detail.

    At the beginning of lactation, in low-producing animals (5.5 kg of milk) a 25% protein deficitand a 2.4% theoretical energy deficit produced an altered cryoscopic index and low titratableacidity (SH). Also in this case, correction of the diet normalised cryoscopy, titratable acidity,cheese-making capacity and determined a not significant production increase.

    On another farm in which low titratable acidity was recorded, the buffaloes were at thebeginning of lactation and received a ration which appeared to satisfy production requirements,in that there was a 20% deficit in only protein requirements. The increase in dietary protein(14.12 vs 9.73%) and energy content (0.9 vs 0.82 MUF/ kg DM) enabled titratable acidity toreturn to regular levels. Also in this case, milk quantity increased only slightly.

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    Although fat content oscillates broadly, fatty acid composition undergoes only slight changesduring lactation insofar as it must first satisfy calf requirements. Hence, with a deficit of one ofthe three components (ruminal, intestinal or hematic) there is little change in fatty acidcomposition and the milk fat quantity decreases.

    For the above reasons, at the beginning of lactation the use of protected fats has a favourableeffect on milk production in that it covers any energy deficits while subsequently when drymatter intake is regularised and thus when there are enough ruminal and intestinal precursors it causes an increase in milk fat content.

    The above effect depends not only on the lactation phase but also on production level: moreproductive animals usually increase or persist in production for longer while less productiveanimals increase their milk fat content.

    In a species with poor catabolic habits their use allows us to maximise the use of forage,promotes its persistence, reduces weight loss and, given the same energy density, enabling tolimit the content of starches that cause fattening, shorten the length of lactation and in thebuffalo are not strictly indispensable for bacterial synthesis.

    In contrast with the above comments on the high tolerance of buffalo towards protein surplus,

    the species is unable to maintain homeostasis with an excess of soluble sugars+starch in itsdiet. In summer months in Italy, on farms where excessive amounts (30 kg) of fresh corn orsorghum are administered, milk titratable acidity (SH) increases up to 12 SH with detrimentalconsequences for cheese-making aptitude. The effects of rationing in buffalo are often lessevident than those observed in dairy cows.

    The above difference is much more evident when the herds genetic level is not high.

    When the results of the experimental trial in buffalo are examined, interpretation is notstraightforward in that the effect of treatment may be reflected in production, in milk fatcontent, in both or in one only to the detriment of the other due to a dilution effect. Theresult is can be more easily understood if they are evaluated in terms of ECM or, better still, ofkg of mozzarella produced. If each farmer were to process his/her own milk, more attentionwould be paid to rationing!

    Rationing which best meets the needs of animals allows to higher mozzarella production. Thisshould drive livestock farmers and processors to formulate commercial regulations which bettermeet the requirements of both. This is a difficult goal to pursue insofar as their interests do notalways coincide. It should be easier to take such ideas on board when farmer and cheese-maker work for a single producer. However, even in this case the capacity to evaluate economicreturns and apply extended time horizons cannot always be taken for granted.

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