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4.3 Anti-nutritive and Toxic Factors in Forage Tree Legumes

B.W. Norton

Screening Techniques for Plant Toxins
Secondary Plant Compounds in Forage Tree Legumes


Plants have co-evolved with predator populations of bacteria, insects, fungi and grazing animals, and have developed defence mechanisms which assist their survival. Leguminous trees and shrubs often have thorns, fibrous foliage and growth habits which protect the crown of the tree from defoliation. Many plants also produce chemicals which are not directly involved in the process of plant growth (secondary compounds), but act as deterrents to insect and fungal attack. These compounds also affect animals (including humans) and the nutritive value of forages. Mycotoxins (fungal metabolites) produced by saprophytic and endophytic fungi are also a potential source of toxins in forages.

The effects of both secondary metabolites and mycotoxins vary with animal species. Non-ruminants (e.g. pigs, poultry and horses) are usually more susceptible to toxicity than ruminants which have the capacity to denature potential toxins in the rumen. The nature and action of toxins in plants have been the subject of several reviews (Duke 1977, Rosenthal and Janzen 1979, Hegarty 1982, Seawright et al. 1985, Barry and Blaney 1987), in which attention was focused on pasture plants of commercial importance.

This section reviews the information available on the anti-nutritive and toxic compounds which have been found in forage tree legumes.

Screening Techniques for Plant Toxins

In plant introduction and range evaluation programmes, there is a need to rapidly screen large numbers of plants for nutritive value, palatability and potential toxicity. Since it is usually not feasible to collect sufficient herbage to feed ruminants, alternative techniques have been developed using laboratory animals (e.g. mice, rats, rabbits and guinea pigs) as test animals. Rats have been used to detect a wide range of plant toxins potentially harmful to man and other monogastric animals. Since the rat is more sensitive to plant toxins than ruminants, several workers have used a rat bioassay to test the presence of toxins in tropical pasture legumes (Bindon and Lamond 1966, Strickland et al. 1987). Although toxicity to rats is not necessarily indicative of similar problems in ruminants, the process allows rapid identification of potentially toxic species which can then be investigated further in feeding trials with ruminants. The results of these bioassays will be discussed in this section. With the increasing interest in tree legume species, there is a need to extend these trials to a wider range of species.

Secondary Plant Compounds in Forage Tree Legumes

Secondary plant compounds may produce toxic effects in ruminant animals (e.g. cyanide, nitrate and fluoroacetate), may depress intake and/or utilisation of feed components (mycotoxins, high tannins), or may enhance feed nutritive value (low tannins, anti-protozoa! activity). Table 4.3.1 provides a summary of the compounds which have been found in tree legume species which may affect animal productivity. The list of species and compounds is not exhaustive, and some compounds listed may not be toxic. Although mycotoxins in forages are known to cause some commercially important toxicoses in grazing stock (e.g. ryegrass staggers, lupinosis and facial eczema), no comparable reports can be found for stock consuming tree species.

The mode of action of some of the compounds (tannins, cyanogenic glycosides and saponins) has been described in detail by other reviewers and will not be attempted here. In the following section, the significance of these compounds to the nutritive value of the major tree legume species will be discussed.


Mulga (Acacia aneura) is the most common Acacia species used for stock feeding in Australia. As mentioned in Section 4.2, mulga is of low nutritive value. Many acacias have high concentrations of phenolic compounds in their leaves, the major compounds being lignin and tannins. The tannins may be further categorised into hydrolysable tannins (polyesters of garlic acid and hexahydroxyphenic acid derivatives) and condensed tannins (proanthocyanidins). In ruminants, the ingestion of hydrolysable tannins can cause death, but these animals have a higher tolerance of condensed tannins. The condensed tannins in mulga inhibit plant protein degradation in the rumen and decrease rumen availability of sulphur, which then depresses the digestibility of plant cell walls. It is also possible that these tannins inhibit microbial enzymes in the rumen and decrease the availability of plant proteins for digestion in the intestines.

Table 4.3.1. A list of secondary compounds found in some forage tree legume species.


Plant part



Acacia aneura


condensed tannins, oxalate


Acacia cambagei


cyanogenic glycoside (CG)


CG hydrolase



Acacia cana

leaf, stem



Acacia doratoxylon

leaf, stem



Acacia georgina

leaf, stem

CG hydrolase (no CG)




Acacia salicina

leaf, bark






Albizia chinensis


echinocystic acid


glycosides, oleanolic acid



condensed tannins


Albizia lebbeck


various sterols



pipecolic add derivatives



echinocystic acid


Calliandra calothyrsus


condensed tannins


Calliandra haematocephala


pipecolic acid derivative


Calliandra portoricensis


tannins, saponins, flavonoids,



Gliricidia sepium





condensed tannins



coumarins, melilotic add



CG, nitrate



canavanine, heat stable toxin


Leucaena leucocephala





condensed tannins



flavanol glycosides


Sesbania grandiflora

leaf, seed

condensed tannins, glycosides



methyl oleanolate


Sesbania sesban


(glucuronide-oleanolic acid)



saponin, heat labile toxin



(stigmasta galactopyranoside)


* References: 1. Gartner and Hurwood (1976); 2. Cunningham et al. (1981); 3. Everist (1969); 4. Hall et al. (1972); 5. Rawat et al. (1989); 6. Ahn et al. (1989); 7. Asif et al. (1986); 8. Romeo (1984); 9. Sotelo et al. (1986); 10. Marlier et al. (1979); 11. Aguwa and Lawal (1987); 12. Calle et al. (1987); 13. Griffiths (1962); 14. Manidool (1985); 15. Hegarty et al. (1964); 16. Lowry et al. (1984); 17. Andal and Sulochana (1986); 18. Kalyanaguranathan et al. (1985); 19. Dorsaz et al. (1988); 20. Shqueir et al. (1989); 21. Kholi (1988)

Tannins do not appear to be present in all Acacia species, and perhaps low tannin species could be selected for further study. Tannins may also be associated with the poor acceptability of young mulga leaves, although volatile oil content is also highest in young leaf (Melville 1947). There is recent evidence that some ruminal microorganisms are able to metabolise tannins, or able to remain active in a high tannin environment, and may be used as inoculants to overcome the detrimental effects of tannins in ruminants (Section 4.6).

Mulga also contains sufficient insoluble oxalate to affect the availability of calcium (Gartner and Hurwood 1976). Where oxalate concentrations are high, calcium oxalate crystals may be formed in the kidney leading to urolithiasis. However, given time to adapt, the microorganisms in the rumen can metabolise moderate amounts of oxalate, and there is little reason to suspect that oxalate poisoning is a serious problem when feeding mulga.

Cyanogenic glycosides (CG) occur in many Acacia species and, when ingested and hydrolysed to free hydrogen cynanide (HCN), cause cyanide toxicity. Cynanide combines with haemoglobin in blood and inhibits respiratory enzymes, ultimately causing death. The response of ruminants to CG ingestion varies. In the rumen, HCN is converted to thiocyanate using available sulphur and thiocyanate is absorbed and excreted. Thiocyanate is a goitrogen, inhibiting the activity of the thyroid gland, and often the effect of CG ingestion is seen as the development of goitre (thyroxine or iodine deficiency). Iodine supplementation overcomes this effect. The ruminal trapping of sulphur may also induce a sulphur deficiency which can be corrected by supplemental sulphur (Wheeler et al. 1975). The formation of HCN from this glycoside requires the presence of a specific hydrolytic enzyme in the plant tissue. Table 4.3.1 shows that both enzymes are not always present and, in the absence of the hydrolytic enzyme, CG are not toxic.

The seeds and pods of Georgina gidyea (Acacia georgina) contain fluoroacetic acid (FA), an organic acid found in a range of other plants (e.g. Dichapetalum, Gastrolobium and Spondanthius). This compound inhibits the Krebs cycle by formation of fluorocitrate and is used as a poison for rats and rabbits. FA and its derivatives are also used as insecticides. Fluorine is a cumulative poison, and its effects are often observed only after stock have been grazing plants containing these compounds for a significant time. When compared with other FA containing plants, G. gidyea has only low concentrations, and FA poisoning is only a problem when the plant is the sole source of food during a drought. It is of some interest that native Australian mammals (e.g. kangaroos and possums) normally consume the seeds of G. gidyea with impunity, suggesting that these species have developed a mechanism for tolerance of toxicity (Twigg et al. 1986).


Albizia lebbeck or Indian siris has been more intensively studied than the faster growing A. chinensis. A major difference between the species is in tannin content. Albizia chinensis contains significant levels of condensed tannins and proanthocyanidins while A. lebbeck contains no extractable tannins (Ahn et al. 1989). Green leaf, fallen leaf and flowers of A. lebbeck have all been shown to be highly palatable and of high nutritive value for sheep (Lowry 1989). Less is known about A. chinensis although it is readily accepted (either fresh or dried) by young goats as a supplement to low quality straws (Robertson 1988, Ash 1990) and is eagerly browsed by does and their kids.

A variety of secondary compounds have been isolated from Albizia species, some having biological activity. A range of sterols (taxerol, cycloartemol, lupeol, campesterol and sitosterol) have been found in the flowers of A. lebbeck (Asif et al. 1986) and a saponin (echinocystic acid) was reported in root extracts (Shrivastava and Saxena 1988). Saponins are glycosides of steroid or triterpenoid compounds (e.g. ursane, oleanane and lupane) and, by their detergent action, have been implicated in the formation of bloat in cattle grazing white clover pastures. Triterpenic substances and glycosides of echinocystic acid (saponin) have been isolated from the bark of A. chinensis, and these bark extracts have been found to have molluscicidal (Ayoub and Yankov 1986), spermacidal (Rawat et al. 1989) and insecticidal (Tripathi and Rizvi 1985) properties. Rahman et al. (1986) also reported that alkaloids from the seeds of A. lebbeck are fungicidal and cytotoxic to selected lines of cancer cells growing in vitro. As the name suggests, the neutral non-protein amino acid albizzine was first isolated from Albizia lebbeck, but no toxic activity has been reported.

Whilst these compounds may provide some protection against plant predators, they do not appear to affect the palatability and intake of forage trees by ruminants. It has been observed that whilst goats will eat the bark of some browse trees, little bark damage is found when goats browse A. chinensis. The high content of saponins in bark may be deterring consumption. There appear to be no reports of saponins in Albizia leaf and the dried leaf is non-toxic when fed to rats (Ahn 1990). Although there is a paucity of information on the effects of A. chinensis fed to ruminants, it appears that Albizia species may prove to be a valuable new source of forage for ruminants.


All Calliandra species appear to contain condensed tannins, with high levels (>10%) in C. calothyrsus. When fed to rats (20% of diet), feed intake (palatability) was high but all rats lost weight (Ahn 1990). Tannins are known to have a direct effect on metabolism. Chickens fed high tannin sorghums develop leg abnormalities (Elkins et al. 1978). Barry et al. (1986) found that plasma growth hormone levels increased with increased intake of condensed tannins by sheep. Tannins react not only with dietary protein but also with enzymes of the gut wall and protein in saliva

Ruminants appear to be more tolerant of tannins than non-ruminants, although few studies have been conducted with forage tree legumes. Palmer and Schlink (1992) have reported that wilting (25°C for 24 h) Calliandra calothyrsus (calliandra) depressed feed intake in sheep when given a sole diet over an 8 day period. Ahn et al. (1989) have shown that drying decreases extractable tannin content of tree legumes, including Calliandra. Table 4.3.2 shows the results from an experiment where frozen (fresh) and dried calliandra were fed as supplements to sheep given a low quality (barley straw) diet. The removal of tannins by polyethylene glycol (PEG) infusion resulted in an increased consumption of straw when frozen calliandra was fed. Drying alone increased straw intake and the digestibility of the cell walls (NDF). PEG infusion increased the digestibility of N. particularly in the rumen, which resulted in higher urinary N losses as ammonia. It was concluded from this study that the presence of both tannins and a heat labile compound in fresh calliandra depresses feed utilisation, and that drying removes a factor (not tannin) which is depressing the digestibility (and intake) of barley straw by these sheep. The nature of this factor is not known, and clearly deserves further study. At the level of feeding used in this study, drying effectively removed the detrimental effect of both tannins and the unknown heat labile factor. There is a need to confirm these effects at higher levels of forage intake, and to reconcile these positive effects of drying with the negative effects found by Palmer and Schlink (1992).

Although there are no reports of secondary compounds in C. calothyrsus, the leaves of C. portoricensis were found to contain saponins, flavonoids and glycosides (Aguwa and Lawal 1987). These extracts have bactericidal (Adensina and Akinwusi 1984, Aguwa and Lawal 1987) and helminthicidal properties in dogs (Adewuni and Akubue 1981). Pipecolic acid, a non-protein amino acid, and its derivatives have been isolated from the leaves of C. haematocephala (Marker et al. 1979) and these compounds were shown to have insecticidal properties (Romeo 1984). The effects of these compounds on sheep is not known, but it does seem possible that some may be useful as stock and human medicines.


There is varying opinion about the nutritive value of Gliricidia sepium (gliricidia). It is generally agreed that it is a high quality forage, but of low palatability when first introduced to animals. Carew (1983) found that gliricidia in the diet of sheep and goats induced diarrhoea and depressed consumption of dried leaves during the first 3 weeks of feeding. Similar observations were made by Robertson (1988) where goats took 5 days to adapt to prescribed intakes of fresh and dried gliricidia leaves. The odour of the leaves has been implicated in this initial reluctance of animals to eat gliricidia (Brewbaker 1986). However, once adapted, there appear to be no long-term detrimental effects on sheep and cattle (Chadhokar 1982). The composition and quality of milk of dairy cows given 30% of their diet as fresh gliricidia leaves was not affected (Chadhokar 1982).

Table 4.3.2. The effects of drying Calliandra calothyrsus and infusion of polyethylene glycol (PEG) on dry matter and N utilisation in sheep given barley straw diets supplemented with Calliandra (adapted from Ahn 1990).








Intake (g/day)






Barley straw















Digestibility (%)

Dry matter










Nitrogen (N)





N utilisation (g/day)

N intake





N in faeces





N in urine





N balance





* Values within a line with different subscripts differ significantly (P < 0.06)

The tannin content of gliricidia leaves does not appear to interfere with plant protein availability but may be one of the factors affecting palatability (Table 4.3.3). Ahn (1990) found that drying removed all extractable tannins from gliricidia increased straw intake, dry matter and N digestibility and N balance in sheep. However, it is not possible to decide whether these effects were due to the tannins or some other factor removed or inactivated during drying. Even though drying removed tannins and improved nutritive value, sheep still consumed gliricidia with reluctance, suggesting that the factor(s) associated with poor palatability were not removed by drying.

The factors affecting palatability of gliricidia in ruminants are probably the same as those that depress digestibility and growth in rabbits and chickens given gliricidia leaf meal diets (Raharjo and Cheeke 1985). Gliricidia and calliandra were the least palatable of the forages offered in this study. Ahn (1990) also found depressed intakes, weight loss and foetal deaths in rats offered a diet containing 20% dried gliricidia leaf. Gliricidia bark and seeds are reported to be used as a rat poison in some countries (Sotelo et al. 1986) suggesting that a toxic principle is present. Coumarins have been found in gliricidia leaf (Table 4.3.1); these compounds are precursors of phyto-oestrogens which have caused infertility and abortion in sheep grazing clover in Australia (Cox and Braden 1974). However, Chadhokar (1982) fed diets containing 75% gliricidia to pregnant sheep and found only beneficial effects of supplementation. Sotelo et al. (1986) have reported a thermostable toxin in gliricidia seeds which killed mice within l week of feeding. These authors isolated a non-protein amino acid, canavanine (2-amino-4-guanidooxy-butyric acid) from gliricidia seeds, and this compound may be associated with the toxicity of gliricidia in non-ruminants. However, many of these reports require confirmation by further experimentation. Nevertheless, despite the problem with palatability in ruminants, the undoubted value of gliricidia as a source of forage makes continued study of this species still worthwhile.


The value of Leucaena leucocephala (leucaena) as a feed for stock has been documented by many workers even though all parts of the plant contain the non-protein amino acid mimosine (b -[N-(3-hydroxy-4-oxopyridyl)]-a -aminopropionic acid) which is highly toxic to non-ruminants. Mimosine acts by interfering in cellular mitosis, and the symptoms of toxicity are alopecia, reduced appetite, reduced weight gain and often death. It is recommended that diets for pigs and poultry should contain less than 10% leucaena.

It is now known that in areas where leucaena is indigenous (Central America), and in parts of Asia, ruminants consuming leucaena appear able to degrade the ruminal metabolite of mimosine, 3-hydroxy-4(1H)-pyridone (DHP), to harmless end-products (Jones and Lowry 1984). This capacity is associated with a specific bacterial population in the rumen of these adapted animals. However, where leucaena has been introduced to ruminant populations without this adaptation, symptoms of toxicity such as alopecia, excessive salivation, lack of coordination of gait, enlarged thyroid glands (low serum thyroxine) and reduced fertility are often observed (Jones 1979). These symptoms have been reported in Papua New Guinea (Holmes et al. 1981) Australia (Hegarty et al. 1976) and can be expected in other areas of the tropics where leucaena has been newly introduced.

Toxicity in ruminants is caused by DHP, which is a potent goitrogen. The severity of toxicity is related to the level of leucaena consumed, and diets containing less than 30% are generally considered safe for ruminants. Alternatively, ruminal organisms capable of degrading DHP may be introduced into the rumen of stock grazing leucaena (Jones and Megarrity 1986), thereby removing this restraint to increased use of leucaena. The management strategies needed to maximise animal productivity from leucaena are reported in Section 4.5.

Table 4.3.3. The effects of drying supplemental Gliricidia and Calliandra leaf on the intake and utilisation of barley straw by sheep (adapted from Ahn 1990).








(a) Intake (g/day)






Barley straw





Tree forage










(b) Digestibility (%)

Dry matter










(c) N utilisation (g/day)

N intake





N in faeces





N in urine





N balance





* Values with a line with different subscripts differ significantly (P < 0.06)


Two species of Sesbania are potentially useful forage sources - the slower growing tree S. grandiflora and the rapidly growing short-lived species S. sesban. Sesbania grandiflora leaves and pods are reported to be palatable and non-toxic for cattle (NAS 1979). Some other reports suggest that the white flowering variety is non-toxic, while the purple flowering type is toxic (Hutagalung 1981). Dried leaves of S. grandiflora have been fed (20% of diet) to milking cows (Tendulkar et al. 1984) and goats (15% of diet) without detrimental effects (van Eys et al. 1985, Ash 1990). Sesbania sesban has also been successfully fed as a sole diet to goats (Singh et al. 1980) and as a supplement to low quality forage for young sheep (Reed and Soller 1987). In this latter study, S. sesban (50% dry matter) was fed for 91 days, during which time the sheep grew at a rate of 48 g/day.

A major difference between the species is that S. grandiflora contains condensed tannin precursors (cyanidins) in leaves, whilst no tannin can be detected in S. sesban. However, both species contain compounds potentially toxic to non-ruminants. The methyl ester of oleanolic acid has been isolated from the flowers of S. grandiflora and shown to have haemolytic effects on sheep and human erythrocytes (Kalyanaguranathan et al. 1985). Olvera et al. (1988) found poor growth and high mortality in Tilapia fingerlings at all levels of inclusion (10-35%) of S. grandiflora leaf meal in a fish meal diet. Similar results were obtained when leaf meal of S. grandiflora was substituted in starter rations for chickens (A. Ash, personal communication).

Oil from the seeds of S. sesban is accorded specie properties in Ayurvedic medicine, and is reported to have bactericidal, cardiac depressant and hypoglycaemic actions. The saponin, stigmasta-galactopyranoside has recently been isolated from S. sesban seeds (Kholi 1988). Dorsaz et al. (1988) have isolated glucuronide derivatives of oleanolic acid which has molluscicidal activity against Biophalaria glabrata, one of the known snail vectors of schistosomiasis. This saponin also showed spermicidal and haemolytic activity. Tripathi and Rizvi (1985) also found that S. sesban extracts had anti-feeding activity against moth larvae. Shqueir et al. (1989) found that the inclusion of S. sesban leaf meal in poultry diets (10% of diet) proved fate. to young chicks, and that the provision of either cholesterol or sitosterol with the diet significantly improved survival. The authors reported that the leaves contain a saponin-like toxin and a heat labile factor. It is clear from these studies that both Sesbania species contain a number of toxins with specific activity against a variety of organisms.

Although no reports of acute toxicity in ruminants were found, many of these trials were of limited duration. In grazing studies with goats in Australia, goats showed a high preference for the bark of S. sesban, even when sufficient leaf was available, killing many trees. No toxic effects were found in goats consuming this bark. It has been observed that the bark of S. sesban accessions may be either green or red, and goats readily consumed green bark. It is not known whether goats have the same preference for the red-barked variety, or whether the colour is indicative of compounds detrimental to ruminants. This aspect of grazing behaviour needs further study. In a year-long field grazing study at the University of Queensland, heifer cattle were initially reluctant to browse S. sesban. However, after 3 months they began to consume Sesbania selectively and their weight gains dramatically improved with no indication of toxicity (Gutteridge and Shelton 1991).


Although many different, and some potentially dangerous, compounds have been isolated from many of the potentially useful forage legume trees, little is known about the specific effects of these compounds on ruminant metabolism. The intense interest in leucaena has generated research which firstly identified the regional problem of mimosine and DHP toxicity, and then proceeded to provide a practical solution to the management of this problem. A newer set of forage trees is now available and is being agronomically evaluated. Their nutritive value needs to be intensively studied before they can be released for widespread use.


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