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M. Chenost and R. Sansoucy


Numerous studies and reviews have already been completed on this topic, e.g. Osbourn (1976), Minson (1976), Stobbs (1976), Balch (1977), Chenost and Meyer (1977), Jarrige (1979), Gohl (1981), Lane (1981), Preston (1982) and Devendra (1988). In the context of the present consultation, we will therefore restrict ourselves to reviewing the main characteristics of the tropical feed resources which should be taken into consideration when defining diets and feeding systems in accordance with the new principles of ruminant digestive physiology and nutrition.


The individual cow's daily production depends not only on its genetic characteristics and its stage of lactation but also a great deal on the quantity and quality of nutrients to its intermediary metabolism. This supply is the result of the voluntary intake and the nutrient density of feed intake.

Voluntary intake depends both on:

The voluntary intake of feed depends essentially on the rate of degradation of its digestible matter into particles of a size small enough to enable their passage from the reticulo-rumen to the lower gut. This degradation is achieved by means of the chewing process (eating and rumination) and the microbial fermentation which takes place in the reticulo-rumen. The cell wall content and the magnitude and nature of lignification of these cell walls are amongst the most important factors which govern the degradability and the rate of passage of a forage.

Good microbial activity will require:

Nutrients required at the tissue level for both maintenance and milk synthesis are supplied by the end products of rumen fermentation (amongst which are the volatile fatty acids (VFAs) and microbial cell proteins) and by the dietary nutrients which have escaped rumen degradation and are digested in the intestine. Depending on the level of production of the host animal, it may be necessary to provide, in addition to the forage, dietary supplements in order to meet its nutritional requirements. These supplements should be administered in a certain amount and should possess characteristics, such that the rumen ecosystem is not impaired and generates the proper amount and relative proportions of microbial protein, VFA energy and glucogenic energy.

In order to define an optimum diet it will therefore be necessary to choose the feeds according to the quality and quantity of energy and nitrogen available. These characteristics cannot be determined by the classical routine analysis. In addition to Crude Protein content and Organic Matter (or energy) digestibility it is important to know:

All these considerations are undoubtedly more important in tropical than in temperate regions even if the levels of animal production are lower. In fact, shortages of nitrogen (tropical feeds also contain less by-pass protein) and of digestible cell-wall energy may occur quite often in tropical countries. It is therefore important to be able to choose the proper missing components among the other locally available resources. As already discussed in several instances, supplements of the tropical basic diets have often more than an additive effect on both intake and animal performances (Preston, 1982; Van Es and Taminga, 1987).

We will briefly distinguish between the basic resources which compose the main parts of the diet and the other various resources which can supplement them.

The former are pastures and green fodders which are of course the principal natural ruminant feed. They are also crop residues, including the fibrous agricultural residues (FAR) which can be used as a substitute (partly or entirely) for herbage in those populated regions like South East Asia where land must firstly be devoted to production of food for man. Another group is the perennial food crops (e.g. sugarcane, bananas), and also roots and tubers which were formerly grown for man and are now more and more considered as feed, either for the dry season or even as the basis of the diet for new feeding systems.

Table 1. Main nutritional characteristics of the principal categories of tropical feed resources.
FeedsRumen fermentationObservations
Pasturesslowly fermentablefair CP content-fair intake
Green fodders  -lignified
ForagesVFA - C2fermentable-rumen function and rumination
   Basis of diet
Crop residues:slowly fermentablevery low CP content-increased OMD and intake with treatment
straws  -need fermentable N + PDIA 
canetops  Basis of diet
Feed crops:  -need fermentable
sugarcaneslowly) fermentable low CP content+ PDIA nitrogen
(whole)quickly) Basis/supplement
Foliages - treefermentablevery high CP-rumen function
crops including  -aa sources (PDIA)
Leucaena,C2 - C3unfermentable*-good intake
Glyricidia, etc. (by-pass)Supplement/Basis
Agro-industrial byproducts:
molasseseasily fermentablelow N 
 (“sugar” type) Supplement/Basis
pulps (citrus)easily fermentablelow N 
 (“cell wall” type)  
energy + N
bran/polishingsfermentable +unfermentableSupplement
 Lipids LCFAN source-PDIA + bypass
   energy, glucogenic
oil cakes + seeds high CP 
animal/fishLipidsPDIA a.a.Supplement
NH3 - urea fermentableindustrial NPN

* Further research is needed regarding the tannin effect on digestion(enzymatic) in the intestine

The other category is roughly made-up of the agro-industrial and various by-products which can be utilized only as part of the diets.

We will now consider the way these feed resources can be utilized in the appropriate combination so that:

Although pastures and green fodders are the principal natural feeds for ruminants, there are also other feed resources which can be used as substitutes during the dry season (e.g., crop presidues), supplements (agro-industrial by-products: cereal brans, molasses, oilcakes, etc.) or as the basis of the diet (sugar cane, roots and tubers, bananas).


It is well recognized that the tropical herbaceous and shrub plants become high in lignified carbohydrates and low in total nitrogen when they mature. In addition their mineral content is low and unbalanced; phosphorous is amongst the most frequent deficient macro-elements.

The digestibility of tropical forages decreases at a lower rate than that of temperate ones but this decrease starts earlier and from a lower value at the young vegetation stage (Chenost, 1975; Evans, 1977). As a result, tropical grasses, and to a lesser extent legumes, always have a lower digestibility than the temperate ones (Minson, 1976) as shown in Table 2. In fact, the high content and the type of encrustation of lignin in the plant tissues and cell walls and the low N supply to rumen microbes are reasons which lead to a slow rate of breakdown and passage of particles to the lower gut and reduced intake of tropical grasses.

However, except in the case of natural pastures in dry tropical areas, tropical pastures have a tremendously high dry matter productivity. This productivity enables maintenance of high stocking rates (carrying capacity) as shown in Table 3.

The yields of tropical C4 grasses (e.g. Digitaria decumbens) responds linearly to annual rainfall (or water supply when irrigated) when fertilized with nitrogen up to 400 kg N/ha (Salette, 1970). Nitrogen fertilization however does not increase the animal's daily production since it has very little or no effect on digestibility and voluntary intake.

Milk production per area unit can thus reach high levels, thanks to the stocking rate, but with low individual production per animal, as shown by numerous authors quoted by Evans (1977) (Table 4). Such low daily milk production levels (seldom higher than 12 kg) may hamper the duration of the lactation curve.

Table 2. Examples of digestibility, intake and rumen fermentation by sheep of some tropical grass hays. (from O. Kawamura et al., 1985)
Nature of grasses(*) Regrowth number Vegetative stage Dry matter intake g/kg LW 0.75 Dry matter digestibility (per cent) Crude Protein content (per cent DM) ADF VFA mm/100ml C2 percent C3 VFA
Green panic 1 flowering 47.7 58.9 10.2        
  2 heading 45.9 53.4 13.5        
  3 heading 52.9 54.4 11.9        
Sudan grass 1 flowering 42.0 53.7 11.9        
  2 vegetative 50.7 55.0 15.6 37.5 5.62 75.1 13.0
  3 vegetative 48.0 58.0 14.9 to to to to
            49.1 8.96 82.6 18.2
Rhodes grass 1 vegetative 44.2 56.3 10.0        
  2 vegetative 42.4 60.7 13.2        
  3 flowering 47.4 59.6 12.2        
African millet 1 vegetative 49.5 60.7 14.0   10.24    
Italian Ryegrass 1 heading 70.1 72.5 13.6 28.1 12.24 70.1 22.3
  2 heading 59.1 58.6 11.1 41.8 10.20 71.6 21.0

(*)wilted and air driedADF = Acid Detergent Fibre; VFA = Volatile Fatty Acids; C2 = Acetic Acid; C3 = Propionic Acid.

Many studies have shown that trying to increase individual production by exploiting the grass cover at an earlier stage of growth is wasteful and uneconomic. A recent study on buffaloes has also lead to the same conclusions (Wanapat and Topark-Ngarm, 1985). In fact, the loss in DM production/ha is not compensated for by the very small benefit, in terms of DOM intake, which could be expected from a faster turn-over.

Whereas a 4,500 kg milk lactation needs, in addition to a typical temperate forage-based diet, an average of 150 g concentrate for each kg of milk produced, the same level of milk production requires an average of 300 to 350 g concentrate in the case of a typical tropical forage-based diet. Such amounts are uneconomic (except when concentrates are subsidized) and illogical (substitution effect of concentrate depresses the DM intake of forage). It is therefore necessary to resort either to an improvement of the basic diet or to design a strategy of supplementation taking into account other local feed resources (see below).

In areas where rainfall is higher than 750 mm per year, it is possible to oversow natural pastures with legumes which, most of the time, are not present in the primary grass cover. The effect of legumes is two-fold: firstly fixation of substantial amounts of N and therefore increase of the production of the associated grasses, and secondly an increase in the feeding value of the grass cover resulting from the higher N content and by-pass protein, the higher OMD and intake of legumes. A lot of research work has shown the importance of fodder legumes (Table 4) on the individual cow's daily production. The strategy to be finally adopted regarding the type of pasture (pure grasses versus grass/legume association) is however not only a nutritional problem but also an agronomical and managerial one.

But tropical green fodder, which represents the cheapest sources of forage, cannot in general ensure high individual milk secretion levels. The main limiting factors are intake and N content and quality.


The fibrous agricultural residues (FAR) represent a considerable potential forage resource in the populated countries where land must be devoted to human food production as a priority. A comprehensive review of their potential in the developing countries and of the strategies for expanding their utilization has been achieved respectively by FAO (1985) and IDRC and ICAR (1988).

Table 3. Carrying capacity and milk production per hectare from various pasture systems (from Stobbs, 1976, quoted by Jarrige, 1979).
Pasture system Stocking rate (cows/ha) Milk production (kg/ha/year)
- unfertilized grass 0.8 – 1.5 1,000 – 2,500
- grass-legume 1.3 – 2.5 3,000 – 8,000
- nitrogen fertilized grass (+P, S, K) 2.5 – 5.0 4,500 – 9,500
- nitrogen fertilized grass, irrigated (+P, S, K) 6.9 – 9.9 15,000 – 22,000

Table 4. Milk production from tropical pastures without supplementary feed (extracted from Evans, 1977)
Pasture Stocking rate (cow/ha) Breed Milk yield
kg/cow/day kg/ha/yr
Unfertilized pastures
P. maximum 1.1 Jersey 6.8 2,667
M. miniflora
P. maximum 1.0 Jersey/Criollo 6.9 2,667
D. decumbens 1.5 Friesian/Zebu 6.9 3,760
Grass-legume fertilized pasture
P. maximum/Glycine 1.3–2.5 Friesian 12.4–13.7 4,954–8,221
D. decumbens/Centro 1.7 Friesian/Zebu 7.3 4,530
Nitrogen fertilized pure grass
D. decumbens 2.5 Jersey 6.8 6,014
D. decumbens 6.9 Friesian/Zebu 10.9 17,408
D. decumbens 8.0 Jersey 6.5 22,466

Amongst the world's total crop residues maize yields the largest amount and wheat, rice and paddy and pulses each yield about half the amount of maize. The remainder consists of sorghum stovers, barley straws, sugarcane tops and leaves, roots and tubers, oil plants stovers and foliage (Kossila, 1985). They are still underutilized as feed resources, except in Asia where they form the first component of the ruminants' diet.

Their feeding value is limited by their poor voluntary intake, low digestibility and low nitrogen, mineral and vitamin content. In addition they are very slowly fermented in the rumen. In fact, they consist essentially of lignified structural carbohydrates, since they represent the dead aerial part of the mature plant after harvest.

The use of FAR as cattle feed has generated considerable research work in the last 20 years but unfortunately much less development application. However, they can represent the basic part of ruminants' diet provided:

Their better digestive utilization can be achieved either through an appropriate supplementation (legumes, molasses, fruit pulps, poultry manure, urea, etc.) or chemical pre-treatments (urea/ammonia treatments) which both facilitate the microbial breakdown of the cell-walls. Appropriate supplements which enable a good cellulolysis can be chosen among the local feed sources on the ground of the characteristics listed in Table 1: the appropriate fermentable N supply can be of natural (poultry manure) or industrial (urea) origin; the fermentable energy (of the “digestible cell-wall” type) is typically fresh grass or good quality foliage and, of course, all the easy digestible agro-industrial pulps, e.g. citrus, pineapple, etc. The breakdown of FAR can also be improved by chemical treatments (sundstø1 and Owen, 1984) among which urea-generated NH3 is probably the technique which best fits in with the socio-economical conditions found in tropical developing countries where inputs must be kept at the lowest level possible.

Treated or not, FAR must be combined with feed supplements which provide adequate nutrients to the rumen microorganisms. The former, however, still require to be known with a better accuracy than at present. Recent research works (Silva and ørskov, 1988; Ramihone, 1987) have shown the importance - in addition to NH3-N - of true protein sources on the cellulolytic microbial growth. This is another reason for supplying any cheap protected protein source (e.g. legume trees and foliages, ricebran), which, as seen above, are necessary for the production requirements of the host animal.


Various perennial food crops which were formerly grown only for human consumption are now more and more considered as feed sources for the dry season and even as the basis of feeding systems. The main ones are sugarcane, cassava, banana, etc.

Amongst the various reasons for such their increasing use, two are probably most important. The first is the tremendous dry matter productivity/ha of these crops. The second one lies in the fact that, as opposed to conventional fodder crops, their nutritive value is not affected by the age of the plant which has already reached its stage of maturity. Their exploitation is therefore very flexible and easier than that of herbage.

1) The most typical example is probably the sugarcane, exploited as a whole plant. Sugarcane could play the same role in tropical animal production as the forage maize - whole plant - in the temperate countries. First considered in experiments by Preston in the 1970's, the sugarcane (whole plant) is typically the addition of two opposite types of forage components: structural carbohydrates of low and slowly digestible energy and soluble carbohydrates (sucrose) rapidly fermentable. In addition its N content is very low.

Whole sugarcane-based systems have proven to be technically and economically a very attractive solution for small to average dairy or dual purpose units in sugarcane producing countries where areas for fodder pastures are limited. As described by Preston and Leng (1978) the deficient nutrients may essentially be provided by locally grown (or available) feed resources:

2) Cassava (Manihot esculenta) is another fodder crop of great interest as a feed resource (Devendra, 1977). Its tubers are a valuable energy source which can also provide glucose at the intestine level as its starch can partially escape the rumen fermentation. Its leaves, exploited either as a green fodder (several cuts before harvesting tubers have proved to be still compatible with a satisfactory tuber yield) or, at the time of harvesting, the tubers are valuable sources of both PDIN and, to a reasonable extent, PDIA.

3) Another interesting plant is the banana, either considered as a whole plant (when blown down by tropical winds and hurricanes) or as fruit (starch source) when considering the discarded bananas which remain available on the premises of the conditioning exportation units (Le Dividich et al., 1976). The banana as a basis of the diet is more adapted to beef production in view of its high starch and poor N content. As seen above it may however remarkably complement sugarcane. The whole plant can also be envisaged as the basis of the diet for milk producing animals.


They can be classified into 4 groups:

All these by-products have been reviewed by various authors (Chenost and Meyer 1977, IDRC and ICAR 1988). We will therefore restrict to a brief account of the more important ones taken as examples.


This is a feed which is rapidly and entirely fermented in the rumen. Between 10 and 30% of the diet, as is traditionally the case there is no particular problems with molasses for all types of livestock. However when the diet is based on molasses (eg: >70%) the behavior of cattle is different and the management of the herd must be more careful (Preston and willis 1974). A small amount of fibre is vital for ensuring the normal physical function of the rumen. Non-protein nitrogen is essential for the development of the micro-organisms of the rumen. Furthermore the animal responds dramatically to small amount of protein like fish meal, which can escape the rumen fermentation (Preston 1985).

However it has never been possible to incorporate as high levels of molasses in the diet of lactating cows as in the case of fattening cattle. The reason is that diets high in molasses lead to insufficient amount of glucose and glucose precursors (low propionate and high butyrate) in the end products of digestion (Leng and Preston 1976).

Molasses which is an excellent carrier for urea as a source of non protein nitrogen for ruminants can be more easily used as a supplement and distributed to small farmers when is part of solid multinutrient blocks (Leng 1984, Sansoucy 1986, Sansoucy et al 1988).

Citrus and sugarbeet pulps

Due to the high digestibility of their non lignified cell walls they favor, as opposed to molasses, the cellulolytic activity of the rumen. Due to the relatively moderate rate of fermentation (as opposed to sugars) they also represent good carriers of NPN and ensure an efficient microbial synthesis (synchronization of both ATP and NH3 releases). They can constitute the major part of the diet as well as an excellent energy supplement for diets based on fibrous crop residues.

Oil cakes and seeds and by-products of animal origin

They have been comprehensively reviewed in the 1988 IDRC and ICAR's publication. They constitute the largest source of supplementary protein. As mentioned earlier in this paper, the assessment of their potential use as protein supplement will be based on the degree of degradability of their nitrogen in the rumen.

As they represent a source of foreign exchange and of high quality protein for human and non ruminant animals, their use as protein supplement for ruminants should be considered against the local availability of legumes and or legume trees.

Cereal milling by-products

They are very well known and their use is expanding. Their major asset is the fact that they supply at a time, moderately fermentable energy, dietary protein and neoglucogenic nutrients. As an example, rice polishings can play a remarkable role in balancing sugarcane based diets.


As a main concluding remark and as clearly observed by Preston and Leng (1987), the tropical basic feed resources have in common the fact that they are poor in nitrogen (namely in protected dietary PDIA) and rich in carbohydrates. These carbohydrates are however either structural and slowly fermentable or too easily fermentable compared to those of the temperate fodder plants, rich both in cell wall type and in less fermentable type of energy. As a result, taken alone or in combination with each other, they will be fermented in the rumen at very different rates. In addition there is another drawback in that fermentable N (predominant in the main tropical feedstuffs) may also be released too quickly and not in time with the energy. Supplementing tropical feeds with crop residues, feed crops and agro-industrial by-products, will therefore have to take into consideration not only the above described characteristics but also the kinetics of release of the various nutrients. Attention to the rationing aspects will be of major importance.


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