3. L. Moil and H. Kuypers, Exp. Brain Res. 23 (Suppl.), 142 (1975).

21 OCTOBER 1977


Thr Thr


Gly Gly

to amino acids whose codons terminate only with a pyrimidine. The total number 319

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4. J. F. Fulton, Arch. Neurol. Psychiatry 31, 221 and in which commissurotomy was not and J. P. Schade, Eds. (Elsevier, Amsterdam, (1934); M. A. Kennard, H. R. Viets, J. F. Ful1964), p. 168. performed, the abnormal behavior, even ton, Brain 57, 69 (1934); A. M. Travis, ibid. 78, 10. H. G. J. M. Kuypers and D. G. Lawrence, Brain 174 (1955). Res. 4, 151 (1967); C. E. Catsman-Berrevoets when tested soon after the operation, 5. I. Stepidn,Acta Neurobiol. Exp. 34,145(1974). and H. G. J. M. Kuypers, Neurosci. Lett. 3, 245 was either mild or did not occur. 6. D. R. Meyer, C. Hughes, D. J. Buchholz, A. D. (1976). L. Dalhouse, J. Enloe, P. M. Brain M. Kemp and T. P. S. Powell, Brain 93, 525 Res. J. Meyer, 11. The monkeys' behavior with the af108, 397 (1976); N. Butters and D. Pandya, SciW. J. Nauta and W. P. Mehler, Brain (1970); fected arm may represent the visual ence 165, 1271 (1969). Res. 1, 3 (1966); D. A. Hopkins and L. W. Nies7. N. C. in Woolsey, Biological and Biochemical Neurosci. Lett. 2, 253 (1976). sen, counterpart of the somatosensory guided Bases of Behavior, H. F. Harlow and C. N. 12. We thank D. Hopkins for many helpful remarks, Woolsey, Eds. (Univ. of Wisconsin Press, grasping and groping behavior (4) that E. Dalm for his assistance during surgery, W. Madison, 1958), p. 502; I. H. Wagman, in The van den Oudenalder for help with the photogracan occur in monkey and man with conOculomotor System, M. B. Bender, Ed. (Harper phy, and J. Verhoeven for the invaluable help in & Row, New York, 1964), p. 18. tralateral premotor lesions. The behavior handling the animals. This investigation was 8. P. H. Schilier and M. Stryker, J. Neurophysiol. supported in part by grant 134615 of FUNGO in some respects resembles the defect in 35, 915 (1972). (Dutch Organization for Fundamental Research 9. G. D. Lawrence and H. J. M. G. in Brain Kuypers, Medicine). delayed alternation response in which 91, 1 (1968); ibid., p. 15; H. G. J. M. Kuypers, in animals show an overriding tendency to Organization of the Spinal Cord, J. C. Eccles I February 1977; revised I June 1977 reach to where food had previously been obtained. It also resembles Stepien's (5) "magnet reaction," in which animals presented with a conditioned stimulus at How Many Anticodons? one place and a reward at another tend to go and to remain at the place where the Abstract. Much new information on codon composition is becoming available conditioned stimulus is delivered. How- from the sequencing of molecules of DNA and RNA. The "wobble rules" for codonever, these behavioral defects are the re- anticodon pairing are applicable to this information. These rules provide for only 54 sult of bilateral lesions located more ros- anticodons to pair with 61 codons, because the base A is not found in the first positrally than our premotor ablation (5, 6). tion of anticodons. The behavior of our animals might be due to the fact that the ablation, which Recent advances in molecular biology sition of the codons of OX174. Such inpresumably interferes with the cortical have enabled long sequences of nucle- formation draws attention to the steering of axial and proximal move- otides to be identified in DNA and RNA "wobble rules" of pairing (4) between ments (7), also results in a disinhibition molecules (1). This new information en- the first bases of anticodons and the third of a subcortical region that directs the larges the scope of molecular evolution, bases of codons. Some amino acids have contralateral arm and hand straight to a which formerly depended mainly on only pyrimidine-terminated codons. The visual target, in the same way as the su- comparisons of the amino acid se- only known anticodons for such amino perior colliculus, for example, may di- quences of proteins. Such sequences re- acids start with G, for example, GAA for rect the gaze (8). veal very little about the third bases of phenylalanine. The wobble rules do not This postulated subcortical region codons, but, with the new information exclude the possibility of AAA; the abprobably contributes fibers to the ventro- on nucleotide sequences, these are now sence of A from the first position of all medial group of descending brainstem identified. As a result, Sanger et al. (2) known anticodons is thought to result pathways (9), since accurate reaching found that there is a disproportionately from the action of anticodon deaminase movements of the arm to a visible object high use of T (U) (3) in the third base po- (5). This changes adenine to hypoxancan be readily elicited in bilaterally pyrthine, so that anticodons starting with I amidotomized animals (9), but are large(inosine) occur in the case of amino acids ly abolished when in these animals the Table 1. Amino acid anticodons, possible and with more than two codons, such as valventromedial brainstem pathways (9) are identified (italic). ine. The absence of an IAA anticodon also transected. The cells of origin of GAA for phenylalanine is explainable by conAla Phe IGC these brainstem pathways receive many UAA Ala cluding that this anticodon is eliminated Leu GGC UGC Ala by lethality, for it could "mis-pair" with Leu cortical connections directly from the CAA Ala UUA Leu CGC ablated premotor areas and the rostral IAG (leucine), and similarly pari passu Leu GUA Tyr part of the precentral gyrus (10), as well GAG for the other amino acids whose codons His Leu UAG GUG as indirectly by way of the striatum and CAG UUG Leu Gln terminate only with a pyrimidine. It is the substantia nigra (11). Elimination of IAU Ile Gln therefore anticodon deaminase rather CUG Ile GUU Asn than [as stated elsewhere (6)] the wobble some of these connections may be re- GAU UAU Ile UUU Lys rules that sponsible for the disinhibition of the pos- CAU explain "why there are not 61 Met CUU Lys tulated subcortical region steering the IAC Val GUC Asp transfer RNA's (tRNA's)." Amino acids arm directly to a visual target. GAC Val Glu with two purine-terminated codons can UUC Val Glu have two anticodons, such as lysine with CUC L. MOLL UAC Val GCA Cys H. G. J. M. KUYPERS CAC IGA Ser CCA Trp anticodons UUU and CUU. However, Department ofAnatomy, GGA ICG Ser Arg in the case of, for example, valine, with Medical School, Erasmus University, UGA Ser GCG Arg four codons, there are three known anRotterdam, Netherlands CGA Ser UCG Arg ticodons (IAC, GAC, and UAC) and IGG Pro CCG Arg there is no reason why CAC may not exGGG Pro Ser GCU d Notes Reerenes UGG Pro UCU Arg ist. The wobble rules provide for redun1. R. Haaxma and H. G. J. M. Kuypers, Brain 98, *CGG Pro GCU Arg dancy in coding rather than for pos239 (1975). ICC Thr Gly tulating a number of anticodons less than 2. D. N. Pandya and H. G. J. M. Kuypers, Brain 'ICU Thr GCC Gly 61, but the redundancy does not extend Res. 13, 13 (1969); E. G. Jones and T. P. S. Pow- GlGU Brain 793

of anticodons, counting them as transcribed, before modification of bases, should therefore be 54: one apiece for Phe, Met, Tyr, His, Asn, Asp, Cys, and Trp; two apiece for Gln, Lys, and Glu; three for Ile; four apiece for Val, Pro, Thr, Ala, and Gly; five for Ser; and six apiece for Leu and Arg. Thirty-seven of these have so far been identified in tRNA molecules (7). Some of the anticodons in tRNA molecules contain modified first bases (other than hypoxanthine). Such modifications may either restrict (8) or extend (9) wobble pairing without, of course, engendering ambiguity in amino acid incorporation during peptide synthesis.

RDerencs and Notes 1. W. Fiers et al., Nature (London) 260, 500 (1976); A. M. Maxam and W. Gilbert, Proc. Natl. Acad. Sci. U.S.A. 74, 560 (1977). 2. F. Sanger, G. M. Air, B. G. Barrell, N. L. Brown, A. R. Coulson, J. C. Fiddes, C. A. Hutchison III, P. M. Slocombe, M. Smith, Nature (London) 265, 687 (1977). 3. The abbreviations used in this report are: A, adenine; T, thymine; U, uracil; G, guanine; C, cytosine; I, inosine; Ala, alanine; Arg, arginine; Asn, asparagine; Asp, aspartic acid; Cys, cysteine; Gln, glutamine; Glu, glutamic acid; Gly, glycine; His, histidine; Ile, isoleucine; Leu, leucine; Lys, lysine; Met, methionine; Phe, phenylalanine; Pro, proline; Ser, serine; Thr, threonine; Trp, tryptophan; Tyr, tyrosine; and Val, valine. 4. F. H. C. Crick, J. Mol. Biol. 19, 548 (1966). 5. H. 0. Kammen and S. J. Spengler, Biochim. Biophys. Acta 213, 352 (1970). 6. W. M. Fitch, Science 194, 1173 (1976). 7. T. H. Jukes, Nature (London) 246, 22 (1973); Adv. Enzymol., in press. 8. S. Nishimura, Prog. Nucleic Acid Res. Mol. Biol. 12, 49 (1972). 9. H. Ishikura, Y. Yamada, S. Nishimura, FEBS Lett. 16, 68 (1971).

when the deer population was adequate to sustain wolf numbers (2). However, the deer herd declined rapidly during the next few years as a result of a combination of adverse factors including a high wolf population (10). Wolf pups then starved (9, 11), wolf productivity dropped (9), and wolves became desperate for food (2). Only then did packs begin trespassing widely into neighboring territories, and only then did the Harris Lake pack begin killing deer in its buffer zone (2). Other packs did likewise. Meanwhile, the few remaining deerwintering areas lay in wolf-pack buffer zones (3, 12). Deer migrated from them through one or two pack territories and THOMAS H. JUKES summered in the buffer zones of other packs (3). Furthermore, those deer were Space Science Laboratory, University 21 July 1977 of California, Berkeley 94720 generally older and had survived longer than deer that had lived throughout the area when the population was higher (3). There is little evidence that the deer sought out the buffer zones. Rather, it Wolf-Pack Buffer Zones as Prey Reservoirs appears that these animals just happened Abstract. In a declining herd, surviving deer inhabited overlapping edges ofwolf- to live there and that they survived longpack territories. There, wolves hunted little until desperate, in order to avoid fatal er because such areas were less used by encounters with neighbors. Such encounters reduce wolf numbers and predation wolves. Although such interactions bepressure and apparently allow surviving deer along territory edges to repopulate the came apparent only when the deer to area through dispersal of their prime, less vulnerable offspring into territory cores. wolf ratio decreased drastically, similar but less extreme interactions probably Predator-prey systems tend to survive are accumulating (4, 5) to warrant the ex- take place when deer to wolf ratios are for long periods despite the negative ef- position of a theory of the role of wolf- more usual. fect of the predators. However, the me- pack territory edges in the survival of I propose that these relationships are chanics of the survival process have deer populations. important in helping to perpetuate the been little discussed. The process must Wolf packs in northeastern Minnesota prey population, thereby also helping to involve evolutionary strategies of preda- inhabit a mosaic of adjoining territories perpetuate the predator and, thus, the tor, prey, or both, that (i) benefit one or of 125 to 310 km2 each (6). Around each entire predator-prey system. My theory both, (ii) allow the predator to obtain suf- territory lies a strip about 2 km wide, the is that because wolf packs tend to avoid ficient prey, and (iii) allow enough prey "buffer zone," in which the pack on ei- intensive use of buffer zones, deer into survive. ther side can be found, but in which nei- habiting those areas tend to survive longOne such evolutionary strategy is the ther probably spends much time (7). er and form a reservoir for maintaining tendency of wolves (Canis lupus) to prey Deer usually live throughout wolf terri- and recovering deer populations in the disproportionately on older animals. Al- tories. Individual deer inhabit areas of wolf territory cores. though those predators attempt to catch 0.48 to 4.10 km2 in summer and tend durThis theory implies that when deer any prey they can, their physical abilities ing winter to congregate in "yards" as populations are high, summer deer denrestrict them to capturing primarily, if far as 38 km away from summer ranges sities may be higher in buffer zones than not exclusively, disadvantaged or debili- (3). in territory cores, but the disparity will tated prey (1). Thus wolves can eat, yet The precise size and nature of the be less than when populations are low, their prey populations can themselves wolf-pack buffer zone, as well as the be- when it may not even be measurable. If survive and produce a crop that wolves havior of adjacent wolf packs when with- deer numbers decline, they will become can continue to harvest. This strategy re- in it, are unknown. However, evidence lowest in wolf-pack territory cores first, quires a precise adjustment of the abili- indicates that wolves may feel insecure and dispersing deer from the buffer zone ties of both predator and prey. in this peripheral strip and thus may min- reservoirs can help replenish the cores. A second strategy, which relies on the imize the time they spend there. Wolves If the decline is too great and wolves are spatial organizations of both predator will try to kill members of neighboring forced increasingly into the buffer zones, and prey, is the subject of this report. I packs when they meet (8, 9), and the the probability of mortal strife (9) among recently discovered this strategy in maximum chance of an encounter is in wolf packs increases. This tends to disstudying drastically declining popu- the buffer zone. The rate of scent-mark- courage pack use of buffer zones and relations of white-tailed deer (Odocoileus ing by each pack in the buffer zone is duces the wolf population (9), thus minivirginianus) and wolves in northeastern about twice that in the territory center mizing predation in that area. Because Minnesota. Deer surviving the decline (7), which suggests higher anxiety near buffer zones would constitute 25 to 40 were distributed almost exclusively the territory edge. percent of a region, enough deer would along the edges of wolf-pack territories. One pack of wolves (Harris Lake survive there to help repopulate the rest Although the published evidence is only pack) studied intensively for seven win- of the area. suggestive (2, 3), enough supporting data ters killed few deer in its buffer zone Because deer in the buffer zones 320


How many anticodons?

eHl, 93, (1970). 3. L. Moil and H. Kuypers, Exp. Brain Res. 23 (Suppl.), 142 (1975). 21 OCTOBER 1977 UGU CGU Thr Thr UCC CCC Gly Gly to amino...
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