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Origins of life                                                                                              boundary layer in Stevns Klint, Denmark,
                                                                                                             were not terrestrial contaminants. The same
A left-handed Solar System?                                                                                  criterion was recently applied by Cronin and
                                                                                                             Pizzarello10 to the Murchison meteorite.
Christopher F. Chyba
                                                                                                             These authors discovered enantiomeric
                                                                                                             excesses of up to 9% in apparently non-bio-
         ow and why did life on Earth come to           This controversy led Pillinger6 to empha-            logical amino acids. So it appears that two

H        use almost exclusively laevorotatory,
         or left-handed amino acids (L-enan-
tiomers), rather than their mirror-image
                                                    size the need for a criterion independent of
                                                    enantiomeric excess to judge the authenticity
                                                    of meteoritic amino acids. He suggested
                                                                                                             very different approaches in two laboratories
                                                                                                             confirm enantiomeric excesses in the
                                                                                                             Murchison meteorite.
dextrorotatory, or right-handed forms (D-           determining the isotopic composition of indi-                The same isotope ratios that distinguish
enantiomers)? The paper by Engel and                vidual enantiomers, because terrestrial stable           Murchison amino acids from their terrestrial
Macko on page 265 of this issue1 re-examines        isotope ratios differ from those in meteorites.          counterparts imply that these molecules or
the Murchison meteorite, and reinforces the             Engel and Macko1 appear to have                      their chemical precursors originated in
surprising result that some process favour-         achieved this. These authors had previously7             interstellar clouds1,8. Why should chemistry
ing L-enantiomers seems to have operated            used the carbon isotope ratio 13C/12C of ala-            in such a cloud have a bias towards one-
before the origin of life on Earth, and proba-      nine in Murchison to argue for an excess of              handedness? One possibility10 is that an
bly before the formation of the Solar System.       L-alanine over D-alanine, but reservations               enantiomeric preference may have been
If this is correct, our Solar System may have       about these measurements persisted8. In                  imposed on the cloud out of which our Solar
formed with a built-in bias for left-handed         their present letter1, they instead examine the          System formed by circularly polarized light,
                                                    15
amino acids.                                          N/14N ratio of individual Murchison                    perhaps synchrotron radiation from a
    An excess of L-enantiomers is an intrigu-       amino-acid enantiomers. They find L-enan-                neutron star11. If a substantial fraction of the
ing result, as it suggests that the handedness,     tiomer excesses in the amino acids alanine               organic inventory of early Earth was then
or chirality, of terrestrial biology might have     and glutamic acid — both protein amino                   derived from comets and asteroids12, the
an extraterrestrial cause. But an L-excess          acids — of over 30% and 50%, respectively,               synchrotron-radiation hypothesis would
indigenous to the meteorite is difficult to         with 15N/14N ratios that are too high to                 connect terrestrial biochemistry with the
prove, as it may instead be due to contamina-       include much terrestrial material. The con-              extreme physics of collapsed stars.
tion by the terrestrial biosphere of an origi-      tradiction between this and an earlier                       If the excess was indeed established by
nally racemic mixture. (In a racemic mix-           report2 that alanine and glutamic acid have              some process specific to the molecular cloud
ture, L- and D-enantiomers are present in           L-enantiomer excesses of about 0 and 10%,                out of which our Solar System formed, simi-
equal abundance, as in typical laboratory           respectively, suggests that the Murchison                lar enantiomeric excesses should be found in
chemical syntheses of amino acids.)                 meteorite is very heterogeneous.                         other organic-rich meteorites and in comets.
    When amino acids were first discovered              Another way to establish an extraterres-             Examination of other carbonaceous chon-
in the Murchison meteorite2, apparent               trial origin is to use amino acids that are              drites could therefore test this hypothesis.
excesses of L-enantiomers ranging from 0 to         extremely rare in the terrestrial biosphere.             Unless there is a more widespread process
20% were found. However, the conservative           This was used by Zhao and Bada9 to argue                 operating, other solar systems could have
interpretation of these results was that cont-      that two amino acids found above and below               been formed with a preference for D-amino
amination was to blame, especially as non-          the 65-million-year-old Cretaceous/Tertiary              acids, or with no preference at all — with
protein amino acids were reported to show                             H                           H
no enantiomeric excess3. (‘Protein’ amino                                                                                  Figure 1 Left- and right-handed
acids are of the type used by terrestrial life in    H2N                      CH3       H3C                       NH2      versions of the amino acid alanine.
proteins, and are therefore the most likely to                    C                                   C                    A meteoritic excess of L-alanine
suffer from biological contamination.) A                                                                                   has now been shown not to be
later study4, claiming indigenous enan-                                                                                    from terrestrial contamination, so
                                                                      COOH                    HOOC
tiomeric excesses in Murchison of up to 70%,                                                                               life’s chemical left-handedness
was criticized on similar grounds5.                           L-alanine                          D-alanine                 may be extraterrestrial.



    Avoided objects

    Freudian abstracts                      Dust and fibres from Freud’s couch




234                                                          Nature © Macmillan Publishers Ltd 1997                        NATURE | VOL 389 | 18 SEPTEMBER 1997
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possible consequences for life in those sys-        trial life in the Solar System might be to                             detectors respond to more distant objects.
tems. Alternatively, if the enantiomeric            search for enantiomeric excesses. Mounting                             This is a position-disparity mechanism. Dis-
excess somehow arose during chemical evo-           evidence for (evidently non-biological)                                parities may also be detected by differences
lution within Murchison itself, no correla-         enantiomeric excesses in the Murchison                                 in the distribution of excitatory and
tion between meteorites is demanded.                meteorite means that this criterion alone                              inhibitory regions within two receptive
    Although these enantiomeric excesses            may be less useful than we had hoped —                                 fields that do not differ in position. This is a
suggest a link between extraterrestrial organ-      whether within martian meteorites, on the                              phase-disparity mechanism.
ic molecules and the origin of terrestrial life,    martian surface, or elsewhere.                                             A simple disparity detector that receives
the connection is by no means certain. A            Christopher F. Chyba is in the Department of                           direct inputs from the two eyes does not pro-
preference may have arisen during prebiotic         Planetary Sciences, University of Arizona, Tucson,                     duce a pure disparity signal — one that is
or biotic evolution as well. For example, con-      Arizona 85721, USA.                                                    not influenced by incidental changes in the
sider peptide nucleic acid (PNA), a candidate       1. Engel, M. H. & Macko, S. A. Nature 389, 265–268                     stimulus. For example, the signal for black
DNA precursor molecule. It has been shown               (1997).                                                            bars is not the same as that for white bars, and
                                                    2. Kvenvolden, K. et al. Nature 228, 923–926 (1970).
that an otherwise achiral PNA strand can            3. Kvenvolden, K. A., Lawless, J. G. & Ponamperuma, C. Proc. Natl
                                                                                                                           simple detectors are sensitive to slight
have its chirality fixed by the presence of an L-       Acad. Sci. USA 68, 486–490 (1971).                                 changes in the location of the object7. In the
or D-lysine residue attached to its end13. One      4. Engel, M. H. & Nagy, B. Nature 296, 837–840 (1982).                 cat, more robust disparity signals are pro-
can imagine a picture in which this random          5. Bada, J. L. et al. Nature 301, 494–497 (1983).                      duced by so-called complex cells at a later
                                                    6. Pillinger, C. T. Nature 296, 802 (1982).
‘seeding’ of chirality led to a chiral preference   7. Engel, M. H., Macko, S. A. & Silfer, J. A. Nature 348, 47–49
                                                                                                                           stage of processing. But, even for some com-
in either prebiotic chemistry or early life14. So       (1990).                                                            plex detectors, the sign of the disparity signal
even if the Solar System was born with a pref-      8. Cronin, J. R. & Chang, S. in The Chemistry of Life’s Origins (eds   is inverted when the contrast sign is reversed
erence for L-amino acids, there may have                Greenberg, J. et al.) 209–258 (Kluwer, Amsterdam, 1993).           such that white regions in one eye are super-
                                                    9. Zhao, M. & Bada, J. L. Nature 339, 463–465 (1989).
been other opportunities for chiral choices         10. Cronin, J. R. & Pizzarello, S. Science 275, 951–965 (1997).
                                                                                                                           imposed on black regions in the other8.
to be made, and our understanding is far too        11. Bonner, W. A. Origins of Life 21, 59–111 (1991).                       Cumming and Parker3 now report that
limited to know whether these subsequent            12. Chyba, C. F. & Sagan, C. in Comets and the Origin and Evolution    most binocular cells in V1 of the monkey
choices might have dominated any initial                of Life (eds Thomas, P. J., Chyba, C. F. & McKay, C. P.) 147–173   produce a normal disparity signal to images
                                                        (Springer, New York, 1997).
enantiomeric bias.                                  13. Wittung, P. et al. Nature 368, 561–563 (1994).
                                                                                                                           with the same sign of contrast, but an invert-
    Finally, Lederberg15 suggested in 1965          14. Cohen, J. Science 267, 1265–1266 (1995).                           ed signal to reversed-contrast images. Visu-
that one criterion for detecting extraterres-       15. Lederberg, J. Nature 207, 9–13 (1965).                             ally, images with reversed contrast evoke
                                                                                                                           rivalry, but they do not create an impression
Binocular vision                                                                                                           of depth (see Fig. 1 on page 281). These
                                                                                                                           results are important because they show that
Seeing in reverse                                                                                                          disparity detectors in V1 of the primate
                                                                                                                           respond to a type of disparity that is not used
Ian P. Howard
                                                                                                                           directly for depth perception. Furthermore,
                                                                                                                           they indicate that (contrary to the generally
   t is remarkable that we perceive a three-        rather than fuse. When similar images fall on                          held view), cells in V1 cannot detect when

I  dimensional world using two-dimensional
   images. Artists discovered that depth per-
ception arises from perspective, shading and
                                                    slightly different retinal regions, we see one
                                                    object standing out in depth (Fig. 1). This is
                                                    the magic of stereoscopic vision. The sim-
                                                                                                                           images are in register — this function pre-
                                                                                                                           sumably occurs in a higher visual centre.
                                                                                                                               What are these inverted-disparity signals
occlusion of distant objects by closer ones. Yet    plest idea is that optic-nerve fibres serving                          used for? Two or more simple detectors
it was not until Wheatstone invented the            slightly different regions (receptive fields) in                       could feed into a complex detector which
stereoscope, in 1838, that it was realized that     each of the two retinas converge onto the                              accepts only same-contrast signals. Other
differences in the positions of images in our       same binocular cell. Such a cell is a ‘disparity                       complex cells could use reversed-contrast
two eyes —binocular disparities — also create       detector’, because it responds best when the                           signals to evoke rivalry and help to indicate
impressions of depth1. (One of Wheatstone’s         images in the two eyes have a disparity that                           whether complex images in the two eyes are
stereograms is shown in Fig. 1.)                    matches the difference in the positions of the                         in proper register (that is, whether they have
    Recently, people have become fascinated         two receptive fields6. Different disparity                             a maximum proportion of same-contrast
by autostereograms, in which a three-               detectors respond to different signs and                               edges). But periodic stimuli, even when in
dimensional scene emerges when the eyes             magnitudes of disparity. Zero-disparity                                register, can contain regions of opposite con-
misconverge on an apparently random array           detectors respond to objects in the plane of                           trast. Under these circumstances, complex
of dots. We now have stereoscopic virtual           fixation, crossed-disparity detectors react to                         cells that accept opposite-contrast stimuli
reality and stereoscopic ‘vision’ in robots.        nearer objects, and uncrossed-disparity                                could use them to improve their response8.
But what are the brain mechanisms that are                                                                                     When we look at a nearby object our
involved in stereopsis (binocular vision)2?                                                                                eyes converge, and when we look at a far
An advance in our understanding of these                                                                                   object our eyes diverge. These eye move-
mechanisms comes from results by Cum-                                                                                      ments bring the images of the object of
ming and Parker3 and by Masson et al.4, on                                                                                 interest, as far as possible, onto corre-
pages 280 and 283, respectively, of this issue.                                                                            sponding points, and simplify the task of
    When we look at a small object, the                                                                                    detecting disparities due to relative depth.
images of it fall on corresponding points in        Figure 1 One of the original stereograms                               Given that reversed-contrast images pro-
the two retinas. Neural signals from these          constructed by Wheatstone in 1838. When the                            duce inverted disparity signals, do they also
corresponding points then converge on the           displays are fused by convergence, the inner ring                      evoke reversed vergence?
same ‘binocular cell’ in the primary visual         appears nearer than the outer ring, because its                            To answer this question, Masson et al.4
cortex (V1) of the brain5. As a result, the         images have a crossed disparity. When the                              measured the vergence responses of monkeys
images fuse and we see one object. But if the       displays are fused by divergence, the inner ring                       and humans to arrays of overlapping
images differ widely in shape, orientation or       appears more distant, because its images have an                       reversed-contrast disks. Vergence occurred in
luminance contrast, they rival one another          uncrossed disparity.                                                   the opposite direction to that evoked by same-
NATURE | VOL 389 | 18 SEPTEMBER 1997                            Nature © Macmillan Publishers Ltd 1997                                                                235

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389234a0

  • 1. news and views Origins of life boundary layer in Stevns Klint, Denmark, were not terrestrial contaminants. The same A left-handed Solar System? criterion was recently applied by Cronin and Pizzarello10 to the Murchison meteorite. Christopher F. Chyba These authors discovered enantiomeric excesses of up to 9% in apparently non-bio- ow and why did life on Earth come to This controversy led Pillinger6 to empha- logical amino acids. So it appears that two H use almost exclusively laevorotatory, or left-handed amino acids (L-enan- tiomers), rather than their mirror-image size the need for a criterion independent of enantiomeric excess to judge the authenticity of meteoritic amino acids. He suggested very different approaches in two laboratories confirm enantiomeric excesses in the Murchison meteorite. dextrorotatory, or right-handed forms (D- determining the isotopic composition of indi- The same isotope ratios that distinguish enantiomers)? The paper by Engel and vidual enantiomers, because terrestrial stable Murchison amino acids from their terrestrial Macko on page 265 of this issue1 re-examines isotope ratios differ from those in meteorites. counterparts imply that these molecules or the Murchison meteorite, and reinforces the Engel and Macko1 appear to have their chemical precursors originated in surprising result that some process favour- achieved this. These authors had previously7 interstellar clouds1,8. Why should chemistry ing L-enantiomers seems to have operated used the carbon isotope ratio 13C/12C of ala- in such a cloud have a bias towards one- before the origin of life on Earth, and proba- nine in Murchison to argue for an excess of handedness? One possibility10 is that an bly before the formation of the Solar System. L-alanine over D-alanine, but reservations enantiomeric preference may have been If this is correct, our Solar System may have about these measurements persisted8. In imposed on the cloud out of which our Solar formed with a built-in bias for left-handed their present letter1, they instead examine the System formed by circularly polarized light, 15 amino acids. N/14N ratio of individual Murchison perhaps synchrotron radiation from a An excess of L-enantiomers is an intrigu- amino-acid enantiomers. They find L-enan- neutron star11. If a substantial fraction of the ing result, as it suggests that the handedness, tiomer excesses in the amino acids alanine organic inventory of early Earth was then or chirality, of terrestrial biology might have and glutamic acid — both protein amino derived from comets and asteroids12, the an extraterrestrial cause. But an L-excess acids — of over 30% and 50%, respectively, synchrotron-radiation hypothesis would indigenous to the meteorite is difficult to with 15N/14N ratios that are too high to connect terrestrial biochemistry with the prove, as it may instead be due to contamina- include much terrestrial material. The con- extreme physics of collapsed stars. tion by the terrestrial biosphere of an origi- tradiction between this and an earlier If the excess was indeed established by nally racemic mixture. (In a racemic mix- report2 that alanine and glutamic acid have some process specific to the molecular cloud ture, L- and D-enantiomers are present in L-enantiomer excesses of about 0 and 10%, out of which our Solar System formed, simi- equal abundance, as in typical laboratory respectively, suggests that the Murchison lar enantiomeric excesses should be found in chemical syntheses of amino acids.) meteorite is very heterogeneous. other organic-rich meteorites and in comets. When amino acids were first discovered Another way to establish an extraterres- Examination of other carbonaceous chon- in the Murchison meteorite2, apparent trial origin is to use amino acids that are drites could therefore test this hypothesis. excesses of L-enantiomers ranging from 0 to extremely rare in the terrestrial biosphere. Unless there is a more widespread process 20% were found. However, the conservative This was used by Zhao and Bada9 to argue operating, other solar systems could have interpretation of these results was that cont- that two amino acids found above and below been formed with a preference for D-amino amination was to blame, especially as non- the 65-million-year-old Cretaceous/Tertiary acids, or with no preference at all — with protein amino acids were reported to show H H no enantiomeric excess3. (‘Protein’ amino Figure 1 Left- and right-handed acids are of the type used by terrestrial life in H2N CH3 H3C NH2 versions of the amino acid alanine. proteins, and are therefore the most likely to C C A meteoritic excess of L-alanine suffer from biological contamination.) A has now been shown not to be later study4, claiming indigenous enan- from terrestrial contamination, so COOH HOOC tiomeric excesses in Murchison of up to 70%, life’s chemical left-handedness was criticized on similar grounds5. L-alanine D-alanine may be extraterrestrial. Avoided objects Freudian abstracts Dust and fibres from Freud’s couch 234 Nature © Macmillan Publishers Ltd 1997 NATURE | VOL 389 | 18 SEPTEMBER 1997
  • 2. news and views possible consequences for life in those sys- trial life in the Solar System might be to detectors respond to more distant objects. tems. Alternatively, if the enantiomeric search for enantiomeric excesses. Mounting This is a position-disparity mechanism. Dis- excess somehow arose during chemical evo- evidence for (evidently non-biological) parities may also be detected by differences lution within Murchison itself, no correla- enantiomeric excesses in the Murchison in the distribution of excitatory and tion between meteorites is demanded. meteorite means that this criterion alone inhibitory regions within two receptive Although these enantiomeric excesses may be less useful than we had hoped — fields that do not differ in position. This is a suggest a link between extraterrestrial organ- whether within martian meteorites, on the phase-disparity mechanism. ic molecules and the origin of terrestrial life, martian surface, or elsewhere. A simple disparity detector that receives the connection is by no means certain. A Christopher F. Chyba is in the Department of direct inputs from the two eyes does not pro- preference may have arisen during prebiotic Planetary Sciences, University of Arizona, Tucson, duce a pure disparity signal — one that is or biotic evolution as well. For example, con- Arizona 85721, USA. not influenced by incidental changes in the sider peptide nucleic acid (PNA), a candidate 1. Engel, M. H. & Macko, S. A. Nature 389, 265–268 stimulus. For example, the signal for black DNA precursor molecule. It has been shown (1997). bars is not the same as that for white bars, and 2. Kvenvolden, K. et al. Nature 228, 923–926 (1970). that an otherwise achiral PNA strand can 3. Kvenvolden, K. A., Lawless, J. G. & Ponamperuma, C. Proc. Natl simple detectors are sensitive to slight have its chirality fixed by the presence of an L- Acad. Sci. USA 68, 486–490 (1971). changes in the location of the object7. In the or D-lysine residue attached to its end13. One 4. Engel, M. H. & Nagy, B. Nature 296, 837–840 (1982). cat, more robust disparity signals are pro- can imagine a picture in which this random 5. Bada, J. L. et al. Nature 301, 494–497 (1983). duced by so-called complex cells at a later 6. Pillinger, C. T. Nature 296, 802 (1982). ‘seeding’ of chirality led to a chiral preference 7. Engel, M. H., Macko, S. A. & Silfer, J. A. Nature 348, 47–49 stage of processing. But, even for some com- in either prebiotic chemistry or early life14. So (1990). plex detectors, the sign of the disparity signal even if the Solar System was born with a pref- 8. Cronin, J. R. & Chang, S. in The Chemistry of Life’s Origins (eds is inverted when the contrast sign is reversed erence for L-amino acids, there may have Greenberg, J. et al.) 209–258 (Kluwer, Amsterdam, 1993). such that white regions in one eye are super- 9. Zhao, M. & Bada, J. L. Nature 339, 463–465 (1989). been other opportunities for chiral choices 10. Cronin, J. R. & Pizzarello, S. Science 275, 951–965 (1997). imposed on black regions in the other8. to be made, and our understanding is far too 11. Bonner, W. A. Origins of Life 21, 59–111 (1991). Cumming and Parker3 now report that limited to know whether these subsequent 12. Chyba, C. F. & Sagan, C. in Comets and the Origin and Evolution most binocular cells in V1 of the monkey choices might have dominated any initial of Life (eds Thomas, P. J., Chyba, C. F. & McKay, C. P.) 147–173 produce a normal disparity signal to images (Springer, New York, 1997). enantiomeric bias. 13. Wittung, P. et al. Nature 368, 561–563 (1994). with the same sign of contrast, but an invert- Finally, Lederberg15 suggested in 1965 14. Cohen, J. Science 267, 1265–1266 (1995). ed signal to reversed-contrast images. Visu- that one criterion for detecting extraterres- 15. Lederberg, J. Nature 207, 9–13 (1965). ally, images with reversed contrast evoke rivalry, but they do not create an impression Binocular vision of depth (see Fig. 1 on page 281). These results are important because they show that Seeing in reverse disparity detectors in V1 of the primate respond to a type of disparity that is not used Ian P. Howard directly for depth perception. Furthermore, they indicate that (contrary to the generally t is remarkable that we perceive a three- rather than fuse. When similar images fall on held view), cells in V1 cannot detect when I dimensional world using two-dimensional images. Artists discovered that depth per- ception arises from perspective, shading and slightly different retinal regions, we see one object standing out in depth (Fig. 1). This is the magic of stereoscopic vision. The sim- images are in register — this function pre- sumably occurs in a higher visual centre. What are these inverted-disparity signals occlusion of distant objects by closer ones. Yet plest idea is that optic-nerve fibres serving used for? Two or more simple detectors it was not until Wheatstone invented the slightly different regions (receptive fields) in could feed into a complex detector which stereoscope, in 1838, that it was realized that each of the two retinas converge onto the accepts only same-contrast signals. Other differences in the positions of images in our same binocular cell. Such a cell is a ‘disparity complex cells could use reversed-contrast two eyes —binocular disparities — also create detector’, because it responds best when the signals to evoke rivalry and help to indicate impressions of depth1. (One of Wheatstone’s images in the two eyes have a disparity that whether complex images in the two eyes are stereograms is shown in Fig. 1.) matches the difference in the positions of the in proper register (that is, whether they have Recently, people have become fascinated two receptive fields6. Different disparity a maximum proportion of same-contrast by autostereograms, in which a three- detectors respond to different signs and edges). But periodic stimuli, even when in dimensional scene emerges when the eyes magnitudes of disparity. Zero-disparity register, can contain regions of opposite con- misconverge on an apparently random array detectors respond to objects in the plane of trast. Under these circumstances, complex of dots. We now have stereoscopic virtual fixation, crossed-disparity detectors react to cells that accept opposite-contrast stimuli reality and stereoscopic ‘vision’ in robots. nearer objects, and uncrossed-disparity could use them to improve their response8. But what are the brain mechanisms that are When we look at a nearby object our involved in stereopsis (binocular vision)2? eyes converge, and when we look at a far An advance in our understanding of these object our eyes diverge. These eye move- mechanisms comes from results by Cum- ments bring the images of the object of ming and Parker3 and by Masson et al.4, on interest, as far as possible, onto corre- pages 280 and 283, respectively, of this issue. sponding points, and simplify the task of When we look at a small object, the detecting disparities due to relative depth. images of it fall on corresponding points in Figure 1 One of the original stereograms Given that reversed-contrast images pro- the two retinas. Neural signals from these constructed by Wheatstone in 1838. When the duce inverted disparity signals, do they also corresponding points then converge on the displays are fused by convergence, the inner ring evoke reversed vergence? same ‘binocular cell’ in the primary visual appears nearer than the outer ring, because its To answer this question, Masson et al.4 cortex (V1) of the brain5. As a result, the images have a crossed disparity. When the measured the vergence responses of monkeys images fuse and we see one object. But if the displays are fused by divergence, the inner ring and humans to arrays of overlapping images differ widely in shape, orientation or appears more distant, because its images have an reversed-contrast disks. Vergence occurred in luminance contrast, they rival one another uncrossed disparity. the opposite direction to that evoked by same- NATURE | VOL 389 | 18 SEPTEMBER 1997 Nature © Macmillan Publishers Ltd 1997 235