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Plants causing poisoning outbreaks of livestock in South America: A review

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Official websites use. Share sensitive information only on official, secure websites. This paper reviews toxic plants for livestock in South America. We included plants in this review, considering only the plants that caused at least one demonstrated outbreak of poisoning. Plants reported as toxic in other regions and present in South America, but with no confirmed outbreaks of poisoning in this region, are not included. We report the main aspects of the epidemiology, clinical signs, pathology and economical importance of the poisonings. This review highlights the great diversity of toxic plants capable of affecting livestock in South America, and for this reason studies in this regard should be intensified. Some important toxic plants in the region are Senecio spp. Pteridium spp. Mimosa tenuiflora and Cesnotigma pyramidalis are important cause of malformations in livestock in Northeastern Brazil. Plants containing fluoroacetate cause severe losses in livestock in Brazil and Solanum glaucophyllum in Argentina. Some good forages like Trifolium spp. According to Tokarnia et al. In most of these plants, the toxicity was demonstrated by observing outbreaks and by the experimental reproduction of the disease. Also, we include as toxic plants for livestock those that contain a known toxic compound causing outbreaks of disease typical of this compound e. In this manuscript, we did not consider as toxic plants many species that occur in South America and have been reported as toxic in other regions, sometimes with a known toxic compound, but outbreaks in South America have not been demonstrated and published e. We also did not include those plants whose toxicity has been experimentally demonstrated in South America, but no spontaneous outbreaks of intoxication have been recorded. Toxic plants cause severe economic losses, including direct losses due to the death of animals, reproductive losses, reduced production, subclinical diseases, and decreased immunity, and indirect losses due to the cost of control measures, loss of forage, the replacement of dead animals, reduced value of the farms, and costs of diagnosis and treatment. There is little data about South America's economic losses caused by toxic plants. In Brazil, based on data obtained from diagnostic laboratories from different Brazilian regions, annual losses due to animal deaths caused by plants are estimated as approximately 1,3 million cattle, , sheep, 58, goats, and 38, horses from a population of nearly million cattle, 17,6 million sheep, 9,3 million goats, and 5,5 million horses Pessoa et al. It has also been estimated that in the Brazilian semiarid region, from a population of 7,6 million goats and 7,7 million sheep, , kids and , lambs died annually due to malformations caused by Mimosa tenuiflora and Cesnotigma pyramidalis Pessoa et al. Losses due to toxic plants had not been estimated in other South American countries. In the last few years, mainly in Brazil, Argentina and Uruguay, research groups and diagnostic laboratories reported many plant poisonings. At present, there are known toxic plants in Brazil, 68 in Argentina, and 45 in Uruguay. At all we include toxic plants from different South American countries. These studies, most of them performed by diagnostic laboratories from the universities, refer mainly to the description of the epidemiology, clinical signs, and pathology of intoxications. Fewer studies include the identification of the toxic compounds of plants. In this review, poisonous plants to livestock in South America will be presented, with emphasis on aspects related to the epidemiology of the intoxications, economic impact, toxic compounds, pathogenesis, characterization of clinicopathological features, and control and prevention techniques. The plants are grouped initially according to their toxic compound. After describing the poisoning by plants with a specific toxic compound, we include the plants with a similar effect but with an unknown toxic compound. In cases where the toxic compound is not known and the toxicity of the plant is not similar to that of known toxic compounds, the plants were classified by the affected organ system. In Brazil, 12 species from three plant families contain monofluoroacetate MFA and cause sudden death associated with exercise. This group comprises four species of the Rubiaceae family Palicourea marcgravii Fig. Monofluoroacetate has been detected in other species such as Palicourea amapaensis , Palicourea longiflora , Palicourea barraensis , Palicourea macarthurorum , Palicourea nigricans , Palicourea vacillans , Palicourea aff. However, these species have not been associated with outbreaks Lee et al. Palicourea marcgravii is probably the most important Brazilian toxic plant found in forests in all of Brazil except the southern region. After deforestation, large areas of pasture created a favorable environment for P. Plants containing monofluoracetate. Poisoning by MFA-containing plants mainly affects cattle, but sheep and goats can also be affected, mainly by A. Buffalos are nearly six times more resistant than cattle to P. Clinical manifestations are peracute and characterized by collapse, tremors, difficulty breathing, limb paddling, and finally death due to acute heart failure within a few minutes Cunha et al. Gross findings are nonspecific or absent. Hydropic vacuolar degeneration of distal convoluted tubule cells with nuclear pyknosis in the kidneys is observed in most cases Tokarnia et al. Abortions, embryonic deaths, and neonatal mortality are observed in goats experimentally intoxicated by A. In Colombia, sudden death following exercise has been associated with Amorimia concinna Gonzalez, and Tanaecium exitiosum Mora, , but no MFA has been determined in these species. Tanaecium bilabiatum causes sudden death in cattle in the region of the Orinoco River and its tributaries in Venezuela Cortes, Fencing off areas to prevent animals from accessing the plant is the most effective preventive measure, and animals that have consumed the plant should not be subjected to exercise. The use of bacteria that hydrolyze fluoroacetate to prevent poisoning has been experimentally proven but is not being used under field conditions Silva et al. Nerium oleander Apocynaceae and Kalanchoe blossfeldiana Crassulaceae containing cardiotoxic glycosides have been reported in Brazil. Both plants are ingested when plants or their branches are cut and placed within reach of animals. Outbreaks of poisoning by N. Persea americana Lauraceae poisoning was recently described in horses in Southeast Brazil. The disease presents a chronic clinical course associated with severe cardiac fibrosis. Poisonings by Ateleia glazioveana Leguminosae in southern Brazil, Niedenzuella acutifolia Malpighiaceae in southeastern Brazil, and Niedenzuella multiglandulosa Malpighiaceae in southeastern and midwestern Brazil cause three clinically different diseases in domestic ruminants, which can occur singly or together: 1 cardiac form with fibrosis of the heart causing sudden death or congestive heart failure; 2 a perinatal form with abortion or neonatal death; and 3 a nervous form with spongiosis status spongiosus of the nervous system Riet-Correa et al. However, considerable overlap exists between the different forms of the disease. The observation of different clinical manifestations depends on the amount and duration of plant consumption. Aborted fetuses, and calves and lambs that died after parturition had cardiac and brain lesions similar to those observed in adult animals, indicating that the unknown toxin passed through the placental barrier Riet-Correa et al. In South America, five plant genera from three families contain pyrrolizidine alkaloids PA and may cause three syndromes: 1 chronic liver fibrosis cirrhosis ; 2 hepatogenous photosensitization; and 3 centrilobular liver necrosis with high doses of PA. Senecio Asteraceae is the genus most often associated with outbreaks in cattle. In Brazil, it is the most important poisonous plant for cattle in the state of Rio Grande do Sul Southern Brazil ; species related to outbreaks include Senecio brasiliensis , Senecio selloi , Senecio cisplatinus , Senecio heterotrichius , Senecio oxyphyllus , Senecio madagascariensis, and Senecio tweedei Panziera et al. In Uruguay, there are cases of poisoning by S. In Chile, several species of Senecio occurred, but only Senecio erraticus has been associated with poisoning Araya, The disease mainly affects cattle Panziera et al. Poisoning has also been described in sheep in Brazil Giaretta et al. However, sheep are more resistant to intoxication and are used to control the plant Stigger et al. In the state of Rio Grande do Sul, southern Brazil, it has been demonstrated that a stocking rate of 0. However, some species, such as S. Crotalaria retusa Leguminosae is the most important poisonous plant for horses in the northeast region of Brazil causing hepatic fibrosis and hepatic encephalopathy Nobre et al. In this same region, when seeds are consumed, C. Experiments have shown that single doses of However, sheep can ingest daily doses of These resistant animals through daily administration of non-toxic doses can control the plant, as they ingest it without showing signs. The only problem is that this resistance is lost in a short period 7—15 days Anjos et al. As outbreaks in animals ingesting growing C. Sheep ingesting low, non-toxic amounts of Senecio brasiliensis also develop resistance, while high doses can cause acute poisoning Grecco et al. In the last years, poisoning by Crotalaria spp. Intoxication by Crotalaria spectabilis causes fibrosis and centrilobular necrosis in horses that ingest oats contaminated by seeds Lacerda et al. Poisoning by seeds of Crotalaria mucronata in cattle and Crotalaria juncea in horses have been reported as the cause of respiratory signs and lung lesions in animals ingesting contaminated grains. Main microscopic lesions included thickening of the alveolar septa, interstitial fibrosis, alveolar, bronchiolar and perivascular edema, proliferation of type II pneumocytes, and proliferation of Clara cells Pessoa et al. A single outbreak of intoxication by Erechtites hieracifolia Asteraceae , containing 0. An outbreak of hepatogenous photosensitization in cattle associated with the consumption of Heliotropium elongatum Boraginaceae was reported in Uruguay Dutra, a. Tephrosia cinerea Leguminosae is a weed that causes marked ascites due to liver fibrosis in sheep in Northeast Brazil. Intoxication occurs due to prolonged consumption of T. Sheep of all ages are affected, and the leaves are the most toxic part of the plant. The clinical course is chronic and clinical and pathological features include ascites, periacinar hepatic fibrosis, portosystemic shunts, and hepatic encephalopathy Riet-Correa et al. Plants that contain diterpene glycosides include Xanthium strumarium Fig. These substances with close structural similarity with carboxyatractyloside cause centrilobular or massive hepatic necrosis, with clinical signs of acute liver failure, including hepatic encephalopathy Pearce et al. Plants containing carboxyatractyloside or substances with similar action. Xanthium strumarium is a toxic plant, mainly for cattle in Uruguay Rivero et al. However, it was diagnosed as causing high mortality in cattle in the State of Minas Gerais in Southeast Brazil. The plant has adapted to the areas flooded by the Araguari River due to several hydroelectric plants that create transient flood conditions similar to what happens in the southern region of Brazil Pampa biome Machado et al. Intoxication occurs by ingestion of cotyledons or burs mixed with silage or cereals Driemeier et al. In Argentina, there have been cases of mortality due to the direct consumption of Xanthium spinosum fruits in the field Costa et al. Pascalia glauca is the main plant causing acute poisoning in cattle in Argentina, but it also affects sheep and goats in this country Medina et al. The plant rarely causes poisoning in cattle in Uruguay Rivero et al. Poisoning occurs by directly consuming the plant in areas invaded by it or by ingesting contaminated hay. Although P. Outbreaks of Cestrum parqui poisoning occur in Uruguay Rivero et al. Most outbreaks are related to forage shortage conditions and mainly affect cattle and occasionally sheep Dutra et al. Cestrum axillare is one of the most important toxic plants in Southeastern Brazil and also occurs in the Northeastern region of this country. It affects mainly cattle, and occasionally buffalo and goats. Most outbreaks occur from April to September, during the dry season Ubiali et al. Cycas revoluta Cycadaceae is an ornamental plant that causes hepatic necrosis in cattle. Poisoning occurs when animals consume it from gardens or when the plant is mixed with forage. Clinicopathological features of C. Evidence of plant consumption is essential for diagnosis Rivero et al. Other plants causing hepatic necrosis in South America, but with unknown toxic compounds are shown in Table 1. Lantadenes A and B are hepatotoxic pentacyclic triterpenoids identified in several Lantana species Verbenaceae. Much of the available information deals with the toxicity of Lantana camara. In Brazil, natural cases have been identified with L. Poisoning by L. Affected animals commonly present photosensitization, with weight loss, jaundice, and photophobia due to obstructive liver damage. The main gross lesions are severe jaundice and enlarged yellow to orange liver. The consumption of high doses of L. Myoporum laetum Myoporaceae contains furanosesquiterpenes, the best known of which is ngaione. Poisoning mainly affects cattle and rarely sheep in Uruguay, Argentina, and southern Brazil Odriozola et al. Cases of poisoning occur from the consumption of the leaves directly from the trees or from their branches that fall after storms. The clinical course is acute, and the animals show apathy, serous eye discharge, abdominal pain, constipation, jaundice, and photosensitization. In South America, three plant genera have been described as the cause of hepatogenous photosensitization caused by the presence of lithogenic saponins: Urochloa syn. Brachiaria spp, Panicum spp. Poaceae and Tribulus terrestris Zygophyllaceae. Poisoning by Urochloa spp. In Argentina, isolated cases of intoxication with a commercial hybrid of U. All Urochloa species are considered toxic, but the concentration of saponins differs between species. Urochloa decumbens contains high levels of lithogenic saponins protodioscin , followed by Urochloa brizantha and Urochloa ruziziensis which contain lower amounts. The least toxic is Urochloa humidicola. Cattle, sheep, and goats are affected, but sheep are more susceptible than cattle, and young animals are more susceptible than adults. However, there is a difference in susceptibility between animals of the same breed. In Brazil, this natural selection has been associated with a decrease in the frequency of poisoning in sheep and cattle in regions where the plant is most used. It is also suggested that there is some degree of adaptation of the animals by gradual consumption or by the transfaunation of rumen microbiota Castro et al. Another factor that has led to a lower frequency of intoxication is the replacement of U. Horses are susceptible to poisoning, but only U. In Brazil, Urochloa spp. Knowing the epidemiological characteristics mentioned above is essential for preventing and controlling intoxication. However, both the use of resistant animals and the utilization of Urocloa spp. Other plants containing lithogenic saponins are Panicum dicotomiflorum , Panicum miliaceum and Tribulus terrestris. Poisoning by Panicum dicotomiflorum has been reported in sheep in northeastern Brazil Riet-Correa et al. Similarly, in the central zone of Argentina, sporadic outbreaks of cattle intoxicated with Panicum miliaceum have been recorded Odriozola, Tribulus terrestris is a perennial, flat-growing creeper with an underground root system adapted to dry environments. In Argentina, this species is found in the dry central region of the country and few outbreaks have been detected in sheep Tapia et al. Although outbreaks are rare, when they occur the incidence is usually high and many animals are affected. It has a variety of toxic and potentially toxic compounds in the plant. However, saponins and some alkaloids seem to be the toxic compounds involved in liver damage and photosensitization Odriozola, Poisonings by pods of Enterolobium contortisiliquum , Enterolobium gummiferum , Stryphnodendron fissuratum , and Stryphnodendron obovatum Fabaceae affect ruminants in the Midwest and Northeast of Brazil Tokarnia et al. Poisoning by Enterolobium cyclocarpum also occurs in Colombia Lozano et al. This group of plants causes three syndromes: 1 ruminal acidosis; 2 hepatogenous photosensitization; and 3 abortion. Animals consume the pods during drought season when they fall to the ground. Triterpene saponins isolated from E. It has been demonstrated that goats consuming non-toxic doses of pods of E. Horses poisoned by E. The plant is most toxic during seed production, when photodynamic compounds are concentrated. Plants causing primary photosensitization. Ricardo Lucena. Heterophyllaea pustulata Rubiaceae Fig. Other plants causing primary photosensitization but with unknown toxic compounds include Froelichia humboldtiana Amaranthaceae Fig. Poisoning by M. In South America, many plants from three families contain swainsonine and, in some cases, calystegines. Six toxic Ipomoea Convolvulaceae species are recognized in Brazil: Ipomoea carnea subsp. Fistulosa Fig. All poisonings by the different species occur in goats in the Northeast region of Brazil, while intoxication by I. The latter was also diagnosed in cattle in the Brazilian Pantanal central western Brazil , and I. Plants that contain indolizidine alkaloids. A Ipomoea carnea var. All these plants contain swainsonine, and some of them also contain calystegines. It has been proven that in I. In Brazil, in the southern and southeastern regions, poisoning by Sida carpinifolia Malvaceae Fig. In the State of Santa Catarina, S. Reproductive failure mainly in goats with chronic clinical signs Oliveira et al. In Argentina, swainsonine poisoning is caused by Ipomoea carnea var. A characteristic of the intoxication by plants that contain swainsonine is that animals usually do not ingest these plants; however, when some animals, due to lack of forage availability, start to consume the plant, by social facilitation, the other animals of the herd acquire the habit to ingest it. Control of poisoning by plants that contain swainsonine in small ruminants by the technique of conditioned food aversion has been successfully used, using lithium chloride LiCl as an aversive. In the case of poisoning in goats by I. Another efficient control measure, especially in areas severely invaded by Ipomoea spp. Animals can be placed in areas invaded by the plant for 15—30 days or until the first clinical signs appear. After this time, the animals must be transferred to areas free of Ipomoea spp. After this period, they can return to ingest Ipomoea spp. Cook et al. Plants containing indole diterpene alkaloids are infected by endophytic fungi that produce this toxic substance. They include: Cynodon dactylon Poaceae Fig. Gardner et al. Indole diterpenoid alkaloids have also been identified in Ipomoea pes caprae Convolvulaceae Stephen T. Lee, personal communication , which causes a disease similar to that caused by I. Plants that contain diterpene alkaloids. The intoxications mentioned above affect goats, sheep, and cattle. The toxic compound can be eliminated in milk affecting offspring Lopes et al. Typical signs of intoxication include generalized muscle tremors that are initially mild but increase when animals are moved, leading to falls. If the animals are left to rest, they recover and return to standing. The most characteristic clinical signs are intention tremors cerebellar with lateral movements of the head and neck that increase if the animals are exercised, leading to loss of balance and fall. Affected animals fully recover within 7—15 days after being removed from infested areas. Only chronic cases can show permanent signs caused by degeneration and subsequent disappearance of Purkinje cells Riet-Correa et al. Other plants show very similar clinical signs. In northern Argentina, several outbreaks of a tremorgenic disease were observed in cattle grazing Megathyrsus maximus Poaceae. However, indole diterpenoids were not found in the analyzed samples of M. These substances have demonstrated neurotoxic potential in vitro and are suggested to be the toxic compound of Prosopis juliflora Fig. The main lesions responsible for the clinical signs is the degeneration and disappearance of motor neurons in the nuclei of cranial nerves, mainly in the trigeminal nerve. Animals poisoned by pods of Prosopis spp. A Prosopis julifllora ; B pods of P. Prosopis spp. In Brazil, outbreaks have been described in cattle, goats, and sheep consuming P. In Peru, P. In Argentina, natural cases of intoxication have been described in cattle and goats consuming Prosopis nigra Fig. Sporadic cases of Prosopis caldenia intoxication have been diagnosed in the central region of Argentina Odriozola, In the semiarid region of Northeastern Brazil, P. In contrast, P. Tryptamine alkaloids present in Phalaris spp. Their toxicity is higher when the plants are young and growing. Phalaris spp. The sudden death syndrome does not occur in South America, and its pathogenesis is unclear. The acute form is the most commonly diagnosed in cattle and sheep Riet-Correa et al. Intoxicated animals initially show hyperexcitability, tremors, marked incoordination, recumbency, inability to stand, convulsions and death within 3—5 days. The chronic form occurs several weeks up to 2 months after the removal of cattle from Phalaris pastures. Affected animals have poor nutritional status, dullness, different degrees of incoordination, and paralysis of the tongue and lips. Because animals frequently fall on the chest they develop edema, bruising and skin lesions on the chest. A hallmark of Phalaris poisoning is the greenish discoloration in the thalamus, midbrain, spinal cord, and the intracytoplasmic brown pigment deposited in neurons, Recently, it was demonstrated the accumulation of neuromelanin and alpha-synuclein in the neurons Tayebi et al. Indigofera lespedezioides Leguminosae which contains indospicine causes chronic neurological disease in horses in the state of Roraima, Brazil Lima et al. Most poisoning cases occur at the end of the drought when I. Histologically, a PAS positive pigment with the characteristics of lipofuscins is observed in neurons and Wallerian-like degeneration is present in some mesencephalic tracts Lima et al. Other toxic compounds found in South America poisonous plants include tetrahydrocannabinol in Cannabis sativa , which caused an outbreak in cattle Driemeier, , caffeine in Coffea arabica and Coffea canephora husks affecting horses Delfiol et al. Other plants affecting the nervous system, in which toxic compounds are unknown are mentioned in Table 2. Oxalate is a common component of plants and it occurs in two forms: soluble and insoluble. Soluble oxalates are one of a number of anti-nutrients in forage plants and block calcium Ca , magnesium Mg and some trace minerals such as iron Fe absorption. High oxalate blood concentration may result in severe hypocalcaemia and hypomagnesaemia. High oxalate salts concentrations in the blood are filtered by the kidney, and form insoluble oxalate crystals that may cause kidney failure. In cattle, nephrosis was reported to be associated with the consumption of Setaria anceps Schenk et al. In contrast to soluble oxalates that cause nephrosis, insoluble oxalates are not absorbed causing gastrointestinal tract irritation. In Brazil, cases of intoxication have been described in cattle and goats that consumed Dieffenbachia picta and Dieffenbachia seguine Dantas et al. After chewing the leaves of these plants, animals develop swelling edema and protrusion of the tongue, and edema of the lips, face and glottis. There is salivation, difficulty apprehending food, hyperthermia, tachycardia, tachypnea and diarrhea. The high concentration of oxalates in some tropical pastures, including Urochloa humidicola and Megathyrsus Panicum maximum is also responsible for fibrous osteodystrophy in horses. The high concentrations of oxalates in these grasses form complexes with Ca preventing its absorption. The bones of the face are enlarged and there is a generalized bone fragility Tokarnia et al. Tannins are naturally occurring plant polyphenols. Their main characteristic is that they bind and precipitate proteins. They can have a large influence on the nutritive value of many foods eaten by humans and animals. In cases of intoxication, the animals present gastrointestinal lesions and severe nephrosis. In South America, tannin containing plants toxic to cattle include Quercus robur and Combretum Thiloa glaucocarpa. Spontaneous poisoning by Q. The poisoning usually occurs in late summer or autumn when forage is scarce and the availability of green acorns is very high, or in early spring when shoots and young oak leaves are widely available. The first signs are observed several weeks after the animals enter the grazing area with Q. Intoxicated animals show weakness, progressive weight loss, poor body condition, pale mucous membranes, tachycardia, ruminal impaction, and constipation with dry feces covered with mucus. Combretum glaucocarpa is the most important nephrotoxic plant in Brazil and as Q. The leaves are ingested by cattle, causing a subacute disease, with evolution between 5 and 20 days. The affected animals show subcutaneous edema, especially in the thigh, perineum, and scrotum, extending to the ventral abdomen, thorax, and submandibular region. Usually, the lethality is high. The outbreaks occur annually, always at the start of the rainy season when C. Different species of Amaranthus have been reported as nephrotoxic in Brazil, including Amaranthus hybridus, Amaranthus quitensis, Amaranthus blitum and Amaranthus retroflexus for cattle Ferreira et al. The intoxication occurs when the animals are introduced into paddocks severely invaded by the plant, mainly in harvested fields of corn, soybean, sorghum and watermelon, where large amounts of the plants remain in the paddock. In Argentina and Uruguay, the only species associated with kidney failure in cattle was Amaranthus quitensis Rodriguez et al. The poisoning occurs in autumn when Amaranthus spp. Clinical signs and pathology of these nephrotoxic plants are similar to other plants causing nephrosis. Leucaena leucocephala Leg. Mimosoideae is a forage plant characterized by high leaf concentrations of mimosine, and toxicity associated with its consumption in livestock is relatively frequent. Mimosine is a toxic non-protein amino acid chemically similar to tyrosine. Leucaena toxicity occurs due to the incorporation of mimosine into biologically vital proteins in place of tyrosine. Spontaneous poisoning cases have been described in Brazil and Argentina in cattle, sheep, goats, and horses. The main clinical signs are loss of hair or wool alopecia and, in horses, hair loss of the tail and mane. Lethargy and decreased appetite, and excessive salivation is also observed. The animals recover after being removed from access to this toxic plant Riet-Correa et al. The ergot alkaloids are mycotoxins that have a significant impact on livestock health and productivity around the world. They are secondary metabolites of the Claviceps spp. In cattle, tall fescue poisoning includes three clinical syndromes: 1 dysthermic syndrome; 2 gangrenous syndrome; and 3 bovine fat necrosis. It is an important forage species in some South American countries, particularly Argentina's central region. In this country dysthermic syndrome is a common problem observed during summer, causing economic losses. Gangrenous ergotism or fescue foot syndrome appears sporadically in cattle during the autumn-winter when temperatures are low and in flooded fields in Argentina Odriozola et al. Bovine fat necrosis syndrome, characterized by abnormal accumulation of masses of hard fat in the abdominal cavities of cattle, has not been reported in the region. In horses, the consumption of ergot alkaloid in S. Tephrosia noctiflora Leguminosae causes dermatopathy and, in severe cases, emaciation and death of cattle in Northeastern Brazil. The plant is found invading pasture areas of Urochloa decumbens. Clinical and pathological features include alopecia, exudative dermatitis, lichenification, limb and dewlap edema, and weight loss. The animals recover within two weeks of being removed from the areas invaded by the plant Barbosa et al. Quinolizidine and piperidine alkaloids contained in several plants induce birth defects. The mechanism of action of quinolizidine and piperidine alkaloid-induced teratogenesis is similar, however, there are differences in incidence, susceptible gestational periods, and severity between livestock species. The toxicity of this species is attributed to the presence of N-methyl and N,N-dimethyltryptamine Gardner et al. Mimosa tenuiflora poisoning causes malformations in goats, sheep, and cattle that naturally consume the plant, mainly during drought Pimentel et al. Newborn kids and lambs have various types of malformations, including congenital contractures affecting the legs arthrogryposis , spine scoliosis and lordosis , neck kyphosis , and rib cage. Malformations of the bones of the head and face, including micrognathia, primary cleft lip due to unilateral or bilateral hypoplasia or aplasia of the incisive bone, and secondary cleft palate palatoschisis , are also observed. Some animals are born blind, with varying degrees of opacity of the cornea and microphthalmia, and others with ocular dermoids. Plants causing malformation abortion and embryonic deaths. Similar malformations have been described in goats and sheep consuming Cenostigma pyramidale Fabaceae Fig. Poincianella pyramidalis, Caesalpinia pyramidalis in the same region of Brazil. In this case it is estimated that the toxic principles are the same or similar to M. Both plants also produce embryonic mortality and abortion Lopes et al. Conium maculatum hemlock is a common annual or biennial herb commonly distributed in temperate regions of South America. Although cases of acute intoxication with this plant have been described, they are very infrequent. The toxicity of C. An abortion outbreak in cattle was associated with consumption of Hesperocyparis Cupressus macrocarpa and Cupressus arizonica needles. There were no significant macroscopic or histological findings in the fetuses. Abortifacient labdane acids were detected in the needles of H. Buroni et al. In Argentina, abortion in cattle was associated with the consumption of Pinus ponderosa Pinaceae needles with concentrations of isocupressic acid greater than 0. Trifolium subterraneum Leguminosae contains phytoestrogens that cause reduced fertility rates in cattle and sheep. Formononetin levels greater than 0. Phytoestrogen poisoning is uncommon in South America. Only one outbreak affecting Holstein heifers grazing on pastures predominantly of Trifolium subterraneum var. Yarloop for four months has been reported from the state of Rio Grande do Sul, Brazil. The heifers showed development of the mammary gland and vulva, dilatation of the cervical canal, and production of vaginal secretions Pimentel et al. Aspidosperma pyrifolium Apocynaceae Fig. Poisoning occurs when the plant is consumed during the dry period or when it re-sprouts shortly after the rains. In goats, embryonic death occurs mainly in the first 34 days of pregnancy, and abortion can occur at any stage of pregnancy Riet-Correa et al. Ricin and ricinine are the toxins of Ricinus communis Euphorbiaceae. Ricinine is an alkaloid present in the leaves and pericarps, that causes neurotoxic effects probably due to increased release of glutamate and inhibition of the postsynaptic receptors, but without gross and microscopic lesions in nervous tissue. Ricin is a lectin in seeds, primarily affecting the alimentary tract causing necrotic gastroenteritis, hypersalivation, diarrhea, dehydration, teeth grinding, hypothermia, colic, ruminal stasis, and recumbency. Simultaneous manifestations neurological and gastroenteritis occur in cases of consumption of the whole plant. Ricinine can be detected in the liver and rumen fluids Machado et al. Poisoning occurs in conditions of forage shortage or when the plant is cut and offered to animals. In Brazil, the poisoning has been reported in cattle Albuquerque et al. Macrocyclic trichothecenes cause severe digestive signs in ruminants and horses. In South America, the first known toxic plant poisoning reported since the 17th century, was Baccharis coridifolia Asteraceae. Roridins and verrucarins were also produced by endophytic fungus isolated from B. The poisoning caused acute clinical digestive signs; most affected animals died in 12—96 h. Other plants containing macrocyclic trichothecenes and causing similar clinical signs to B. Baccharis vulneraria Costa et al. Phorbol esters are the main toxic compound of Jatropha spp. Poisoning by Jatropha curcas in livestock occurs mainly by the ingestion of by-products of oil extraction press cake , seeds, or seed hulls. However, such poisonings have only been reported experimentally in Brazil. In contrast, spontaneous outbreaks of goat poisoning by the ingestion of Jatropha ribifolia are reported in the semiarid region of Northeastern Brazil. The first cases of poisoning occurred when the goats started to consume the plant due to the severe shortage of forage during the dry season. Later, social facilitation influenced other goats to start eating the plant, which caused outbreaks in successive years. Affected goats presented apathy, anorexia, soft feces, weight loss, severe dehydration, and a reddish pigment in the skin, lips, horns, and teeth. This pigment is present on J. Emaciation was the main lesion observed on necropsies Pimentel et al. Plants that affect the digestive system in which toxic compounds are unknown are presented in Table 3. Of the plants mentioned in Table 3 it is important to highlight Megathyrsus Panicum maximum. Presently there are nearly 20 million hectares of M. However, since the introduction of M. The disease was reproduced experimentally in penned horses ingesting growing P. Colic in Equidae is due to bloat caused probably by rapid hydrolysis and fermentation of carbohydrates, which are in high content during the growing stage of the plant Cerqueira et al. Pastures that induce colic contain significantly higher starch concentration than pastures that do not cause the disease Souza et al. Tanzania was also identified in Northeastern Brazil as a cause of severe cecal dilatation, tenesmus and abdominal bilateral distention in cattle. The disease did not cause deaths and occurred in a rotational system where the paddocks were grazed with high stocking rate for one day and the animals returned to the same paddock after 28 days. After each grazing period the pastures were fertilized with urea Riet-Correa et al. In Northern Brazil a similar disease has been associate with the ingestion of M. The only Brazilian toxic plant that contains n-propyl-disulfide is Allium cepa Amaryllidaceae onion which causes hemolytic anemia and hemoglobinuria in carnivores, horses, and ruminants. In Brazil, poisoning has been reported in buffalo consuming the onion bulb in the state of Santa Catarina Borelli et al. Other plants causing hemolytic anemia in Brazil are Urochloa arrecta syn. Brachiaria radicans Poaceae tanner-grass , Ditaxis desertorum Euphorbiaceae , and Indigofera suffruticosa Fabaceae. The poisoning by U. Ditaxis desertorum rarely causes hemolytic anemia in cattle in northeastern Brazil Tokarnia et al. The poisoning by Indigofera suffruticosa also affects cattle in northeastern Brazil Barbosa-Neto et al. These plants cause anemia, hemoglobinuria, and progressive weight loss. Additionally, I. The pigment is probably the toxic compound of I. Several outbreaks of osteolathyrism caused by Lathyrus hirsutus Fabaceae were diagnosed in cattle in Uruguay from to Affected cattle, mainly calves, were recumbent or had difficulties for moving and showed progressive weight losses. They recovered within days after being removed from the pastures. The poisoning was reproduced experimentally by the administration of L. Melia azedarach Meliaceae is a plant native to Asia that can be found in some regions of South America. Tetranortriterpenes such as meliatoxin A1, A2, B1, and B2 are the most relevant toxic compounds identified in the fruits of this plant Oelrichs et al. Intoxication occurs by ingesting ripe fruits when they fall on the ground or when branches are within reach of animals. Affected animals virtually show two syndromes: 1 characterized by nausea, vomiting, constipation, or diarrhea, often bloody; 2 characterized by excitement or depression and dyspnea, followed by death. Cyanide poisoning is frequently reported in Brazil, Uruguay, and Argentina Tokarnia et al. In Argentina, it is relatively frequent in the country's central region during dry years. Poisonings by other Sorghum species are rare Juffo et al. Additionally, atypical cases of ataxia and urinary incontinence have been reported in horses and cows grazing forage Sorghum Odriozola, More recently, abortions and calf births with arthrogryposis have been observed in cows grazing grain and forage Sorghum in Brazil and Argentina F. Castro, personal communication. Several native species of Manihot Euphorbiaceae found in the northeastern region of Brazil cause cyanide poisoning when accessed by animals for direct consumption or when eaten after being cut Amorim et al. Manihot esculenta Euphorbiaceae is also a frequent cause of cyanide poisoning when animals are fed with the tubers or their by-products cassava wastewater or cassava husks Silva et al. In Brazil, other cyanogenic plants such as Anadenanthera macrocarpa and Piptadenia viridiflora Fabaceae Tokarnia et al. Poaceae , and Sorghum sudanense cause poisoning in the southern region Gava et al. Nitrate and nitrite poisoning occur in Uruguay, Argentina and Brazil in grass pastures, mainly of Lolium multiflorum or Avena sativa Poaceae , fertilized with nitrogen or after severe drought followed by abundant rainfall Rivero et al. In Argentina, it has been caused by the consumption of sorghum and maize forage during dry years Odriozola, In southern Brazil, poisoning was reported in a stubble reed invaded by Amaranthus spp. In Chile, poisoning by nitrates and nitrites has been described in cattle by the ingestion of Silybum marianum Asteraceae Freire and Araya, In South America, four native calcinogenic plants of the Solanaceae family cause enzootic calcinosis characterized by systemic calcification, hypoparathyroidism, hypercalcitoninism, hypercalcemia, hyperphosphatemia, thyroid C-cell hyperplasia, parathyroid atrophy, osteonecrosis, and osteopetrosis Machado et al. The pathogenesis of enzootic calcinosis is complex but studies demonstrated a similarity with the bone-forming process. Recently, it was demonstrated that extracellular vesicles produced by vascular smooth muscle cells are crucial for developing vascular mineralization Machado et al. Poisoning by Solanum glaucophyllum Fig. The disease mainly affects cattle and rarely sheep, buffaloes, horses and pigs by consumption of the fallen leaves mixed with the pasture Machado et al. Additionally, poisoning caused by contaminated hay was reported Micheloud et al. Calcinogenic plants. Carlos E. Two species of Nierembergia cause enzootic calcinosis in sheep and less frequent in cattle Riet-Correa et al. Nierembergia veitchii Fig. In the case of Nierembergia veitchii , an action similar to 1,25 OH 2 D 3 was demonstrated in chickens Riet-Correa et al. Solanum stuckertii Fig. The ptaquiloside is a norsesquiterpenoid glycoside present in several fern species. The ferns most frequently linked to toxicity in South America are species of the Pteridium complex, mainly Pteridium arachnoideum and Pteridium caudatum Furlan et al. However, recent reports demonstrated that other fern species, such as Pteris deflexa , Pteris plumula and Adiantopsis chlorophylla induce toxicity similar to that of Pteridium spp. The toxic effects of bracken ingestion are diverse, depending on the animal involved and the consumed doses. In bovine, ferns poisoning includes three clinical syndromes: 1 the most common syndrome is bovine enzootic haematuria, which results from neoplastic urinary bladder damage and is induced by chronic ingestion of bracken. It is reported in several South American countries such as: Peru Herencia et al. This presentation is relatively common in Southern Brazil and has been associated with chronic bracken fern consumption Faccin et al. However, bovine papillomavirus type 4 BPV-4 is considered a cofactor in developing these neoplasms Medeiros-Fonseca et al. Such cases occur when a large amount of bracken is ingested by cattle and results in acute depression of hematopoietic activity of the bone marrow inducing anemia, leucopenia, thrombocytopenia and haematuria. Outbreaks of this syndrome have been described in Brazil Tokarnia et al. Melilotus alba and Melilotus officinalis are common legumes used in Argentina. When these plants are used for hay making and silage with high levels of humidity, some species of fungi colonize the material and the coumarins in sweet clover are converted to dicumarol. Dicumarol is a potent vitamin K antagonist and anticoagulant. When hay or toxic silage is consumed for several weeks, changes in the proenzymes necessary for prothrombin synthesis result in a bleeding syndrome. Furthermore, dicumarol can cross the placenta in pregnant animals and the newborn can be affected at birth. Outbreaks of dicumarol poisoning have been sporadically recorded in cattle consuming sweet clover hay in Argentina Colque-Caro et al. The genus Vicia comprises more than species, including Vicia villosa and Vicia sativa Barros et al. The toxic compound that causes hairy vetch poisoning is unknown. Intoxication causes a type IV hypersensitivity reaction, suggesting that some plant constituents act as antigens that sensitize lymphocytes causing multisystem granulomatous inflammation through a mechanism not yet described Sonne et al. The poisoning is characterized by dermatitis, conjunctivitis, fever, diarrhea, and lesions of extensive granulomatous infiltration of various organs. The disease mainly affects adult cattle of different breeds Sonne et al. Although the incidence is usually low, the lethality is high, and many animals die even after being removed from the problem paddock. Spontaneous cases of poisoning by Senna occidentalis have been reported in cattle in Brazil Carmo et al. The toxic compound of S. However, dianthrone, an anthraquinone in S. Poisoning by Senna obtusifolia has been reported in cattle Furlan et al. Usually, poisoning by S. However, intoxication by S. Seeds can also be eaten mixed with chopped green forage Mussart et al. Petiveria alliacea is a perennial herb that can reach 1 m in height, native to the Amazon and other tropical areas of South America. The toxicity of P. Affected animals have incoordination and progressive cachexia with persistent flexion of one or both hocks. Main gross lesions are characterized by diffuse yellow flaccid muscles and yellow discoloration of the kidneys. Main histologic lesions are degeneration, necrosis and atrophy with proliferation of fibrous tissue of skeletal and cardiac muscles, and chronic nephrosis Ruiz, Many compounds have been identified in the plant, but it is not clear which is the cause of the clinical signs. Two toxic plants were reported to cause lymphatic tissue necrosis as the main alteration: Polygala klotzschii Polygalaceae Tokarnia et al. Clinical signs are acute and death occurs after 10—38 h of clinical manifestation. The most characteristic histological lesion is necrosis of lymphatic tissue affecting the spleen, lymph nodes, Peyer's patches, and peribronchial lymphoid tissue. There is diffuse necrosis in the germinal center of lymphoid follicles. Enteritis is observed in both small and large intestines Tokarnia et al. The toxic compound of P. The toxic compound of R. Other plants causing necrosis of the lymphatic tissue, including Baccharis spp. In La Hulla and Tolima departments of Colombia, the consumption of Bunchosia pseudonitida and Bunchosia armeniaca causes pigmentation of different tissues in cattle, sheep, and goats. The poisoning is characterized by bluish-purple or pink pigment in mucous membranes, teeth, elastic tissues, subcutaneous tissue, muscles, bones, and hair. Most of the animals remain without other clinical signs, but some show gait disturbances, emaciation, prostration, recumbency, and death. A similar pigmentation of the viscera and carcasses have been reported in sheep in the Brazilian northeastern region, but the plant responsible for the disease was not identified Pimentel et al. First cases are often seen after at least three months grazing in such pastures. The intoxication is observed throughout the year but is more frequent in spring. The poisoning is characterized by a haemolytic crisis, with jaundice and hemoglobinuria. The death occurs in 24—96 h. The main gross lesions are jaundice, subcutaneous yellow edema, serous liquid in cavities, swollen friable ochre-coloured liver with distended and edematous gallbladder, dark kidneys and dark urine haemoglobinuria. Microscopically the main lesions are degenerative and necrotic lesions of the liver and nephrosis. Chronic phytogen copper intoxication occurs in pastures with low molybdenum, less than 0. The diagnosis is based on the epidemiological data, clinical signs, macroscopic and histologic lesions, and the determination of Cu concentrations in the liver over ppm and kidneys over 80 ppm. For the prevention of intoxication, grazing periods of no more than three months in pastures with a predominance of T. In this review, we report species of toxic plants for livestock. Most of these poisoning have been reported in Argentina, Brazil, and Uruguay, where many diagnostic laboratories and Universities are researching toxic plants for livestock. This research expands the number of known toxic plants reported in the last 40 years; for example, in Brazil, there were 90 known toxic species and 52 genera in , species and 64 genera in , species and 72 genera in , species and 80 genera in , and species and 96 genera in This significant increase in the knowledge of poisonous plants in Brazil in the last 40 years was due to the increase in the number of federal and state universities and diagnostic laboratories in different states with veterinary and animal science education, improving the recognition and diagnosis of livestock diseases. In Northeastern e. As research continues, and as livestock producers become better informed, it is expected that the list of toxic species will continue to increase in the coming years. In general, the study of toxic plants has been related to their epidemiology, clinical signs, and pathology. In contrast, the determination of toxic compounds i. Despite this, several toxic compounds have been identified with the collaboration of researchers from the USA, Brazil, Denmark, and Germany. At this point, it is worth highlighting the collaboration of the USDA Poisonous Plant Research Laboratory at Logan, Utah through which the toxic compounds of various plants were determined, including fluoroacetate, pyrrolizidine alkaloids, diterpene glycosides, lithogenic saponins, indolizidine alkaloids, indole diterpene alkaloids, quinolizidine and piperidine alkaloids, and labdane acids. However, to advance on this issue, it is necessary to create more research groups in Latin America to identify toxic compounds with government funding. Some plants, such as Xanthium spp. Mimosa spp. Research and investments in the creation of biological and chemical control methods are necessary to mitigate these losses. Some of the most important toxic plants reported in South America include: Pteridium spp. Bloat caused mainly by Trifolium spp. Swainsonine-containing plants Ipomoea spp. An important toxic plant for horses mainly in northeastern Brazil is Crotalaria retusa as a cause of liver fibrosis and hepatic encephalopathy; this plant also causes liver necrosis in sheep and goats Anjos et al. Also, in horses and mules, the use of Megathyrsus maximus as the main forage for these species in northern Brazil resulted in great economic losses due to the high number of deaths and the difficulties of using other grasses for Equidae in the region Cerqueira et al. Despite recent advances in the research of toxic plants for livestock in South America, it is important to increase the number of research groups working on plant poisonings and emphasizing the determination of toxic compounds. Micheloud prepared the review and reviewed the manuscript. The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper. We acknowledge all the researchers and institutions that collaborate with us in numerous research projects on poisoning plants, especially to the researchers of the Poisonous Plant Research Laboratory, ARS, USDA, at Logan, UT for their continuous collaboration with South American researchers in the last 20 years. Queiroz for reviewing the manuscript. This section collects any data citations, data availability statements, or supplementary materials included in this article. As a library, NLM provides access to scientific literature. Toxicon X. Find articles by Franklin Riet-Correa. Find articles by Mizael Machado. Find articles by Juan F Micheloud. Open in a new tab. Toxic plants without known toxic compounds that cause hepatic necrosis. Toxic plants without known toxic compounds that affect the nervous system. Toxic plants without known toxic compounds that affect the digestive system. Similar articles. Add to Collections. Create a new collection. Add to an existing collection. Choose a collection Unable to load your collection due to an error Please try again. Add Cancel. Gava et al. Odriozola et al. Andrade et al. Colodel et al. Cruz et al. Canella et al. Rivero et al. Traverso et al. Brum et al. Dutra et al. Riet-Correa et al. Barbosa et al. Bezerra et al. Perdomo and Freitas ; Giannitti and Margineda Bedotti et al. Alvim Barros et al. Borelli et al. Carvalho et al. Araya et al. Pessoa et al. Nascimento et al. Guarana et al. Lima et al. Medeiros et al. Mello et al. Medicago sativa , Trifolium repens , and Trifolium pratense. Cerqueira et al. Brito et al. Odini et al. Torino et al. Peixoto et al. Galiza et al.

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