Venom Yield and Toxicity
One of the most important yet undeterminable factors of the gravity of snakebite is the amount of venom injected into the victim. Since this volume varies considerably in every bite, attempts have been made to determine the amount and toxicity of venom produced by each species of poisonous snake. Individual yield is so variable that a large number of snakes must be milked in order to determine the average yield. Even then there remains an uncertainty as to how this amount may compare with that injected by a biting snake.
Wolff and Githens (1939b:234) made 16 venom extractions from a group of cottonmouths in a two-year period. The average yield per snake fluctuated between 80 and 237 milligrams (actual weight), and toxicity measured as the minimum lethal dose for pigeons varied from 0.05 to 0.16 milligrams (dry weight). No decrease in yield or toxicity was evident during this period. Another group of cottonmouths from which venom was extracted over a period of five years also showed no decrease in yield or toxicity. Of 315 individual extractions the average amount obtained from each individual was 0.55 cubic centimeters of liquid or 0.158 grams of dried venom (28.0 per cent solids). The minimum lethal dosage (M. L. D.) which was determined by injecting intravenously into 350-gram pigeons was found to be 0.09 milligrams (dry weight). Each snake carried approximately 1755 M. L. D.'s of venom.
The record venom extraction for the cottonmouth was 4.0 cubic centimeters (1.094 grams dried venom) taken from a five-foot snake which had been in captivity for 11 weeks and milked five weeks earlier (Wolff and Githens, 1939a:52). The average yield of venom of cottonmouths is about three times the average yield reported for copperheads by Fitch (1960:256), a difference correlated with the greater bulk and relatively large head of the cottonmouth.
Allen and Swindell (1948:13) stated that cottonmouth venom rates third in potency, compared drop for drop to that of Micrurus fulvius and Crotalus adamanteus. Freshly dried cottonmouth venom tested on young white rats showed the lethal dose to be from 23 to 29 milligrams per kilogram of body weight. The venom of 11 one-week-old cottonmouths was found to be more potent than that of adult males. Githens (1935:171) rated C. adamanteus venom as being weaker than that of the copperhead (A. contortrix), which he rated only slightly lower than cottonmouth venom. The crotalids which he ranked more toxic than cottonmouths are: the Pacific rattlesnake (C. viridis oreganus) and the massasauga (S. catenatus). He found A. bilineatus, C. durissus, and C. v. lutosus to have the same toxicity as cottonmouths. Minton (1953:214) found that the intraperitoneal "lethal dose 50" (the dose capable of killing half the experimental mice receiving injections of it) was 6.36 milligrams per kilogram for copperheads. However, in later publications Minton (1954:1079; 1956:146) reported that the "lethal dose 50" for copperheads was 25.65 milligrams. Approximately the same potency was determined for cottonmouths. Several rattlesnakes that he tested showed a higher toxicity than copperheads or cottonmouths.
Criley (1956:378) found the venom of copperheads to be 6.95, nearer Minton's earlier estimate, and rated cottonmouth venom as being twice as toxic as that of copperheads. The relative toxicities of other crotalids tested, considering the cottonmouth to be one unit, were: C. basiliscus, 0.3; A. contortrix, 0.5; C. viridis oreganus, 1.4; A. bilineatus, 2.2; C. adamanteus, 2.3; C. v. viridis, 3.2; C. durissus terrificus, 27.5.
It can be seen from the above examples that toxicity of venoms and the resistance of the animal receiving an injection of venom is highly variable. Possibly the venom of each species of snake has greatest effect on animals of the particular group relied on for food by the snake. If that is so, the venom of cottonmouths would be expected to be more toxic when tested on fish, reptiles, and amphibians than on birds and mammals. Likewise, the venom of most species of rattlesnakes would be expected to be more virulent when injected into mammals than when injected into lower vertebrates. But, according to Netting (1929:108), species of rattlesnakes that prey on cold-blooded animals, which are less susceptible to venoms than warm-blooded animals, are thought to have highly toxic venoms. This explanation accounts for the powerful venom of Sistrurus catenatus; and, in this respect, venom of cottonmouths should be highly toxic also. However, no clear-cut trends have been shown in most cases. Allen (1937) injected 250-gram guinea pigs with 4 milligrams of venom of various poisonous snakes. Survival time was recorded in order to indicate the relative potency of the venoms. Of 16 such tests C. adamanteus held places 1, 2, 3, 12, and 16; Bothrops atrox held places 4, 9, 10, and 13; and A. piscivorus held places 5, 7, 8, and 15. Places 6, 11, and 14 were held by three individuals of different species. No relationship to size or sex was indicated by the results of this experiment.