Pumping Up Intelligence
         
        Abrupt Climate Jumps and the Evolution of
        Higher Intellectual Functions during the Ice Ages
        
        
        
        
        William H. Calvin
        University of Washington
        Seattle WA 98195-1800 USA
        faculty.washington.edu/wcalvin
        WCalvin@U.Washington.edu
        
        
        The title is not a metaphor, though past tense
        might be better as this chapter is about how each of the many hundred
        abrupt coolings of the last several million years could have served as a
        pump stroke, each elevating intelligence a small increment - even though
        what natural selection was operating on was not intelligence per se.
        While we often use the term 'intelligence' to encompass both a broad
        range of abilities and the efficiency with which they're enacted, it
        also implies flexibility and creativity, an "ability to slip the
        bonds of instinct and generate novel solutions to problems" (Gould
        and Gould 1994, p. 70). Those three pillars of animal intelligence -
        association, imitation, and insight - are also impressive (Byrne 1994),
        as are the occasional symbolic (Deacon 1997) and reasoning (Gould &
        Gould, 1998) abilities. But Piaget (1929; 1952) said that intelligence
        is what you use when you don't know what to do, when neither innateness
        nor learning has prepared you for the particular situation.
        Intelligence is improvisational. Still, most of the time, not much
        improvisation is necessary; the individual has encountered somewhat
        similar situations before, has a repertoire of actions, and simply
        starts one - and gropes, using feedback's progress reports to guide to
        the goal. No major planning is needed in most cases, and thus not much
        in the way of intellectual wherewithal. This suggests a focus on those
        few behaviors that require an elaborate multistage plan prepared during
        "get set." An example would be the ballistic movements
        (hammering, clubbing, throwing, kicking, spitting) where the speed of
        feedback is so inadequate (they're often over-and-done by the time that
        progress reports can start modifying the movement), where only
        near-perfect plans will succeed.
        What the mind is often seeking during "get set", I suspect,
        is coherence - finding a conceptual combination of sensory
        input, memories, and movement plans that fit together particularly well
        - though, because of novelty, most will rate less than the "perfect
        ten" of exact, unambiguous fits. Similar to this is Barlow's (1987)
        suggestion that intelligence is all about making a guess that discovers
        some new underlying order. "Guessing well" neatly covers a lot
        of ground relevant to higher intellectual functions: finding the
        solution of a problem or the logic of an argument, happening upon an
        appropriate analogy, creating a pleasing harmony or witty reply, or
        guessing what's likely to happen next.
        
        
        Higher Intellectual Functions
        
 
        
        
        Because they all involve pattern-finding and all emphasize human
        abilities not widely shared with the other apes, I want to restrict
        myself here to the higher intellectual functions. I will use
        "intelligence" as a term denoting the speed-and-scale of
        individual performance of them, not unlike the manner in which much of
        the variance of the general factor g can be accounted for
        (Jensen 1992) by the subtests that emphasize speed of performance and
        the number of items that must be borne in mind simultaneously - as in
        those multiple choice analogies: A is to B as C
        is to [D, E, or F] which require six concepts to be
        simultaneously managed. Closely related is our ability to remember phone
        numbers long enough to dial them. Many people can retain a seven-digit
        number for 5-10 seconds, but will resort to writing it down if faced
        with an out-of-area number or an international one of even greater
        length. When getting close to your limit, you try to collapse several
        items into one chunk, so as to make more room (Simon, 1983).
        Definitions of "higher intellectual function" vary; I tend
        to use the phrase to refer to the structured mental abilities
        such as
        
           syntax, the structuring schemes of phrases and clauses
          used to disambiguate sentences longer than a few words, e.g., "I
          think I saw him leave to go home" has nested embedding involving
          four verbs. (Syntax is an evolutionary puzzle because there aren't
          obvious intermediate forms in development or aphasia between short
          structureless protolanguage sentences and recursive embedding.)
           planning, those speculative structured arrangements,
          e.g., "Maybe we can go to the country this weekend if I get my
          work finished, but if I have to work Saturday, then maybe we can go to
          a movie on Sunday instead." (Squirrels hoarding nuts isn't
          planning but an innate behavior triggered by longer nights releasing
          more melatonin from the pineal. Holding an intention for a few hours
          isn't planning either, and I'd also exclude foraging behaviors that
          could be explained more simply by choosing between familiar migration
          routes. Perhaps we should reserve the term for something that requires
          multiple stages of the move to be assembled in advance of action,
          rather than organizing the later stages after getting the initial
          moves in motion, which goal-plus-feedback can accomplish.)
           chains of logic, our prized rationality. But the
          emphasis here is on novel chains, not routine ones. The most mindless
          of behaviors may be segued, the completion of one calling forth the
          next: courtship behavior may be followed by intricate nest building,
          then a segue into egg laying, then incubation, then the stereotyped
          parental behaviors. (Kφhler's chimps, that piled up boxes to reach
          the hanging banana, might qualify under the novel chaining
          requirement, if simpler explanations can be eliminated.)
           games with arbitrary rules, such as hopscotch. (Both Pan
          species, chimpanzees and bonobos, have a version of "blind man's
          bluff" - but it isn't structured.)
           music of a structured sort, such as harmony and
          counterpunctual themes. (Perhaps not rhythm per se but certainly
          rhythms within rhythms.)
        
        Obviously, if one relaxes the nested-or-chained structural
        requirement, there are various primate behaviors that are possible
        evolutionary precursors.
        I am focusing here on structuring because humans exhibit such a large
        increment in ability over our Pan cousins across these five
        areas - and because I am interested in whether there is a "common
        core" of neural machinery that is shared by all such structured
        behaviors, one where improvements in any one of the five might improve
        the other four "for free."
        Evolutionary arguments often commit the reification fallacy (a
        "gene for intelligence"). Indeed, we often assume that an
        abstraction like language implies a real concrete entity such as a
        "language module." (Separate, of course, from any planning
        module! And so on, to the balkanization of the mind.) Yes, there is
        localization of function - and we certainly tend to name a cortical area
        according to the first of its functions we discover - but multiple
        functions are commonplace (see chapter six of my Cerebral Code
        for a discussion of how a neocortical area could alternate between being
        a narrow specialist and performing as a general-purpose scratch board).
        
        
        Language in the Multiple Use Context
        
 
        
        
        Multiple uses of a structural entity are common, and a familiar
        example is the "curb cut" where the steplike curb is locally
        smoothed into a gentle ramp. What paid for curb cuts was, of course,
        wheelchair requirements. But as soon as a curb cut is in place, 99
        percent of the traffic is for secondary uses such as bicycles,
        suitcases, baby carriages, grocery carts - none of which would have
        "paid for it."
        A secondary "free" use may, of course, later pay for
        further improvements (just imagine the skateboarders holding a bake sale
        to pay for widening!), suggesting that the evolutionary history of
        higher intellectual function might first emphasize one structured use
        and later others. If the notion of a "free lunch" offends,
        note that it is commonly assumed that music is a spare-time use of the
        language-related parts of the brain, that there was likely little
        natural selection for four-part harmony via barbershop quartets.
        Language is the most defining feature of human intelligence: without
        the orderly arrangement of verbal ideas permitted by syntax, we might be
        little more clever than Pan. For a glimpse of life without
        syntax, consider the Sacks (1989) description of Joseph, an 11-year-old
        deaf boy. Because he could not hear spoken language and had never been
        exposed to fluent sign language, Joseph did not have the opportunity to
        learn syntax during the critical years of early childhood:
         
        
          "Joseph saw, distinguished, categorized, used; he
          had no problems with perceptual categorization or generalization, but
          he could not, it seemed, go much beyond this, hold abstract ideas in
          mind, reflect, play, plan. He seemed completely literal -- unable to
          juggle images or hypotheses or possibilities, unable to enter an
          imaginative or figurative realm....He seemed, like an animal, or an
          infant, to be stuck in the present, to be confined to literal and
          immediate perception, though made aware of this by a consciousness
          that no infant could have."
        
         
        "Language cortex" isn't just the traditional Broca and
        Wernicke areas but much of the lateral aspects of the temporal and
        frontal lobes, plus the parietal lobe areas near the left sylvian
        fissure (see, for example, Ojemann 1991). Language localizations have a
        strong overlap with nonlanguage sequential functions such as sound
        strings and hand-arm sequencing (most aphasics have some form of
        hand-arm apraxia; see Kimura 1993).
        And this overlap brings me to an important point: the use that
        initially "paid" for the structuring abilities seen in the
        higher intellectual functions need not be any one of them. The original
        "wheelchair" analog could, for example, be the structured
        planning needed for some nonintellectual function, such as the
        ballistic hand-arm movements used for hammering, clubbing, and throwing.
        Throwing is a particularly interesting possibility because targets
        are located at many different distances and elevations, making each
        hunting throw a novel situation, quite unlike the more stereotyped dart
        throws and basketball free throws where long practice can "find the
        right groove." There is also a premium on being right the first
        time, as dinner is likely to flee if you miss.
        Apes have elementary forms of the rapid arm movements that we're
        experts with - hammering, clubbing, and throwing - and one can imagine
        hunting and toolmaking scenarios that, in some settings (more in a
        moment), were important additions to the basic hominid gathering and
        scavenging strategies. The evolutionary rewards for individuals having
        better-than-average projectile hunting skills could thus set the stage
        for free secondary uses, such as planning on longer time scales, such as
        logical trains of thought, and perhaps even music and syntax. Some of
        these, once they had a chance to show their stuff, are exposed enough to
        natural selection to "pay" for further improvements - and thus
        improve throwing accuracy, pari passu (Calvin, 1983, 1993,
        1996b).
        
        
        Finding the Right Level
        
 
        
        
        Although it seems to have played little role so far in our modern
        concepts of intelligence, the concept of levels of organization
        is a common one in the sciences. Much of guessing well involves finding
        the right level at which to address a problem, neither too literal nor
        too abstract - or, sometimes, inventing a new level on the fly.
        Levels are best defined by certain functional properties (Calvin
        & Bickerton, 1999), not anatomy. As an example of four levels, fleece
        is organized into yarn, which is woven into cloth,
        which can be arranged into clothing. Each of these levels of
        organization is transiently stable, with ratchet-like mechanisms that
        prevent backsliding: fabrics are woven, to prevent their disorganization
        into so much yarn; yarn is spun, to keep it from backsliding into
        fleece.
        A proper level is also characterized by "causal decoupling"
        from adjacent levels (Pagels, 1988); it's a "study unto
        itself." For example, you can weave without understanding how to
        spin yarn (or make clothing). Chemical bonds illustrate a proper level:
        Mendeleyev discovered the patterns of the table of elements, and thereby
        predicted the weights and binding properties of undiscovered elements,
        without knowing anything about the underlying patterns of electron
        shells (or the overlying patterns of stereochemistry).
        Mental life can pyramid a number of levels, thereby creating
        structure. Some of the major tasks of early childhood involve
        discovering four levels of organization in the apparent chaos of the
        surrounding environment:
        
           Infants discover phonemes and create standard categories for
          them; six-month-old Japanese infants can still tell the difference
          between the English /L/ and /R/ sounds but after another six months of
          regular exposure to the Japanese phoneme that lies in between them in
          sound space, the baby will treat the occasional English sounds as mere
          imperfect versions of the Japanese phoneme (Kuhl et al 1992) and so be
          set up for later confusing the English words 'rice' and 'lice'.
           With a set of basic speech sounds, babies start discovering
          longer-duration patterns amid strings of phonemes, averaging nine new
          words every day during the preschool years.
           Between 18-36 months of age, they start to discover
          still-longer patterns of words called phrases and clauses, rules such
          as add -s for plural, add -ed for past tense.
           After syntax, they go on to discover Aristotle's rule about
          narratives having a beginning, middle, and end (and they then demand
          bedtime stories with a proper ending).
        
        Indeed, we find it very rewarding to discover half-hidden patterns
        all through life: that's the basis for the popularity of crossword and
        jigsaw puzzles. It's why science is so much fun.
        Pyramiding four levels in a mere four years is impressive. But levels
        can also be created on the fly, as we seek an analogy or make a novel
        abstraction. To spend more time at the more abstract levels in this
        house of cards, the prior ones have to be sufficiently shored up to
        prevent backsliding over your concentration span.
        
        
        From Stratified Stability to Darwinism
        
 
        
        
        But there's no child inside the head to stack up those higher stories
        of the house of cards, so what self-organizes a higher level? In the
        simpler physical systems, noise (as in diffusion) can provide the raw
        material for self-organizing structures (such as crystals). As Bronowski
        (1973) observed: "The stable units that compose one level or
        stratum are the raw material for random encounters which produce higher
        configurations, some of which will chance to be stable..
" If
        there is an organizational principle in the universe that is even more
        elementary than Darwin's, it is Bronowski's.
        But, as Darwin first realized, competitions between stable
        alternatives can improve the results, providing a quality bootstrap
        under certain conditions. Not all of what is loosely called
        "Darwinian" qualifies, however, as many pruning processes do
        not have a result that copies and competes. As I have discussed
        elsewhere (Calvin 1996, 1997), there appear to be six essential features
        of a recursive Darwinian process:
        
           It involves a pattern. Classically, this is a string
          of DNA bases called a gene. But the pattern could be a melody or the
          brain activity associated with a thought.
           Copies are somehow made of this pattern, as when cells
          divide or you whistle an overheard tune. Indeed, the unit pattern is
          defined by what's semi-reliably copied, e.g., the gene's DNA sequence
          is semi-reliably copied while whole chromosomes or organisms usually
          are not.
           Patterns occasionally change. Point mutations from
          cosmic rays may be the best known alterations, but far more common are
          copying errors and (as in meiosis) shuffling the deck.
           Copying competitions occur for occupation of a limited
          environmental space. For example, several variant patterns called
          bluegrass and crabgrass compete for my back yard.
           The relative success of the variants is influenced by a multifaceted
          environment. For grass, it's the nutrients, water, sunshine, how
          often it's cut, etc. We sometimes say that the environment
          "selects" or that there is selective reproduction or
          selective survival. Darwin called this biasing by the term 'natural
          selection.'
           The next generation is based on which variants survive to
          reproductive age and successfully find mates. The high mortality among
          juveniles makes their environment much more important than that of
          adults. This means that the surviving variants place their own
          reproductive bets from a shifted base, rather than the original center
          of variants at conception (this is what Darwin called an inheritance
          principle). In this next generation, a spread around the
          currently successful is again created. Many new variants will be worse
          than the parental average but some may be even better
          "fitted" to the environment's collection of features.
        
        From all this, one gets that surprising darwinian drift toward
        patterns that almost seem designed for their environment. In the
        cardboard version of darwinian, particular parts such as "natural
        selection" are often confused with the entire darwinian process,
        but no one "essential" by itself will suffice. Without all six
        essentials, the process will shortly grind to a halt.
        For example, neural patterning in development (all of that culling of
        cells and synapses) is an example of a sparse case: just a pattern
        carved by a multifaceted environment. There is no replication
        of the pattern, no variation, no population of the pattern to compete
        with a variant's population, and there's nothing recursive about
        achieving quality because there's no inheritance principle. It's very
        useful but it's not a quality bootstrap.
        
        
        Making Darwinism Fast Enough
        
 
        
        
        Speed is of the essence in behavior, however, and one might
        reasonably worry about whether a neocortical version of the darwinian
        process can operate quickly enough to provide an answer within the
        windows of opportunity afforded by either hunting or social repartee.
        There are at least four "catalysts" which can greatly speed up
        evolutionary processes:
        
           Systematic recombination (crossing over, sex)
          generates many more variants than do copying errors and the far-rarer
          point mutations. There's also nonsystematic recombination, such as
          bacterial conjugation or the conflation of ideas.
           Fluctuating environments (seasons, climate changes,
          diseases) change the name of the game, shaping up more complex
          patterns capable of doing well in several environments. For such
          jack-of-all-trades selection to occur, the climate must change much
          faster than efficiency adaptations can track it (more in a minute).
           Parcellation (as when rising sea level converts the
          hilltops of one large island into an archipelago of small islands)
          typically speeds evolution. It raises the surface-to-volume ratio (or
          perimeter-to-area ratio) and exposes a higher percentage of the
          population to the marginal conditions on the margins.
           Local extinctions (as when an island population becomes too
          small to sustain itself) speed evolution because they create empty
          niches. The pioneers that rediscover the niche get a series of
          generations with no competition, enough resources even for the odder
          variants that would never grow up to reproduce under any competition.
          For a novel pattern, that could represent the chance to
          "establish itself" before the next climate change, for which
          it might prove better suited than the others.
        
        There are also catalysts acting at several removes, as in Darwin's
        example of how the introduction of cats to an English village could
        improve the clover in the surrounding countryside: The (i) cats would
        (ii) eat the mice that (iii) attack the bumble bee nests and, thereby,
        (iv) allow more flowers to be cross pollinated. Although a Darwinian
        process will run without these catalysts, using Darwinian creativity
        often requires some optimization for speed.
        Explaining how neocortical circuitry can implement the six essentials
        and the four catalysts on a milliseconds-to-minutes time scale, thereby
        facilitating intelligent "get set" groping, lies beyond the
        scope of this article (see Calvin 1996, 1998b). However, these ten
        facets of a rapid evolutionary process will be useful in considering how
        we got our big brains so quickly (2.5 million years is quick). What sped
        up the slow biological evolution of the rapid neural evolutionary
        machinery underlying the higher intellectual functions?
        
        
        Hominid Evolution and the Ice Ages
        
 
        
        
        The earliest known changes in hominids, seen soon after the
        australopithecines diverged from the other Pan cousins about
        five million years ago, were rearrangements of hips and knees for
        upright posture. Brain size (an admittedly inadequate indicator of
        functional capacities) didn't change very much, remaining in the great
        ape ballpark. But a number of interesting things all started to happen
        between three and two million years ago.
        
           The archaeologists have traced stone tools back that far: the
          simplest types (the split pebbles which make such good cutting edges
          for getting through animal hides) go back to about 2.5 million years,
          with much more elaborate ones developing by 1.5 million years ago.
          While various mammals use found objects to open shells and
          the like, simple toolmaking (shatter a rock and select the sharp
          edges) seems to have been on the rise by 2.5 million years ago.
           The onset of the ice ages has been moved back to about the same
          time by the paleoclimatologists. Since then, ice sheets have slowly
          built up. They melt off somewhat more quickly (the rise in sea level
          takes about 8,000 years) and remain at a minimum for another similar
          period. The major meltoffs occurred about every 40,000 years - until
          about 0.7 million years ago, when a 100,000-year cycle became more
          prominent. It isn't clear what this has to do with African-based
          hominid evolution, as the average temperatures there only drop about 5°C
          during the colder periods, enough to create some glaciers on the
          equatorial volcanos but hardly enough to create a wintertime for
          animals living in the Rift Valley. All of the ice sheets that formed
          at higher latitudes were surely unseen by our African ancestors.
           And brain size finally starts to change back about 2.4 million
          years ago as the australopithecine lineage split off a distinctive Homo
          lineage with increased cranial capacity (and it kept increasing in
          size; nothing of the kind is known for other animals). Many animal
          lineages also split between 3 and 2 million years ago: chimp-bonobo,
          gibbon-siamang, mastodon-elephant (accelerated speciation is best
          studied, however, in the antelope and pig lineages).
        
        What do these three things have to do with one another, or with
        intelligence? Cause and effect? Or merely three independent trains set
        in motion by the major rearrangement of ocean currents and climate that
        followed the damming up of the "Old Panama Canal" about 3
        million years ago, when North and South American finally joined up and
        forced the equatorial currents that equilibrated the Atlantic and
        Pacific Oceans into a long detour around the southern continents?
        
        
        The Mark Twain Transition Time Principle
        
 
        
        
        The writer Mark Twain once observed that "A round man cannot be
        expected to fit into a square hole right away. He must have time to
        modify his shape." Evolution can often track climate changes,
        selecting for variants with more or less body insulation. Indeed, up
        until a decade ago, we thought that the ice ages were "glacially
        slow," that slow changes in the earth's orbit caused gradual
        cooling, which caused more ice to gradually form, and sea level to
        gradually lower. No animal lived long enough to realize that climate
        changes were happening because the change during the lifetime of any one
        generation was so minuscule.
        What happens if the climate changes abruptly, so that adaptations
        over the generations cannot track it? So that the habitat is largely
        disrupted (no more customary plants or prey, a different setting for
        reproduction, etc.), all within one generation's time on earth?
        There is a general answer to this (Calvin, 1996b) and a much more
        specific one. The general answer is that the circumstance provides a
        selective pressure for versatility, one that counters the usual
        lean-mean-machine tendencies that reduce unneeded anatomy and behavior
        in the name of efficiency. Evolutionary theory suggests a tendency
        towards the latter if the environment remains the same for long
        enough. But when the habitat changes so drastically in so short a
        time, only reserve capacity in behavior can solve the problems. Lean
        mean machines don't survive the downsizings very well. The more
        versatile may have what it takes.
        The more specific answer, however, is grass.
        
        
        Abrupt Climate Change
        
 
        
        
        Inferring ancient climates can be done from layers of sediments that
        accumulate in lake and ocean floors. From cores, one can study such
        proxy climate indicators as pollen types and oxygen isotope ratios, and
        how they change over time. But there's a problem: worms and
        bottom-scavenging fish stir the bottom, mixing together hundreds (if not
        thousands) of years of sediments. Like a moving average of a
        stock-market index, a smoothed record of paleoclimate can miss some
        dramatic fluctuations that made and lost fortunes.
        When year-by-year high-resolution records became available from
        Greenland ice cores, where tree-ring-like records can be seen, we became
        aware that climate could - and frequently did - change quite rapidly. We
        now know that this is not merely a peculiarity of Greenland, that these
        were worldwide events in many cases. Unlike the high latitude ice
        sheets, these abrupt climate shifts affect the tropics as well.
        Atop those glacially slow rhythms, abrupt coolings and warmings have
        often occurred. By 'abrupt' is meant decade transition times; 'often'
        means every few thousand years; these flips may last for centuries (or
        even 1500 years) before flipping back, just as abruptly. Obviously, this
        is not the abrupt climate change which volcanos and Antarctic ice sheet
        collapse can cause; it's bistable. Ocean circulation seems to have
        several modes, and when it switches between modes, the abruptness of the
        transition profoundly affects ecosystems. In the tropics, average annual
        temperatures fall by 3-5°C. In Europe, it's more like 5-9°C and some
        high latitude locations cool more than twice as much.
        The magnitude of the temperature change isn't the big
        problem. It's how fast it occurs, Mark Twain's problem. If
        these coolings were to occur slowly, with temperature ramping down over
        500 years, one might expect high altitude plants and animals to slowly
        move down the hillsides into the valleys below. Each generation of
        hominids could have continued to make a living in much the way their
        parents taught them, though their diet mix would shift over the
        centuries. But with major changes within a decade, other events
        supervene.
        First comes drought (there is much less evaporation from tropical
        oceans, which also reduces a major greenhouse gas, water vapor). Then,
        similar to what we saw in the 1997-98 El Niρo, vast fires occur even in
        tropical forests. After that comes a succession of plants leading, over
        a few centuries, from grass to forests (of species better suited to the
        new annual temperature).
        For many terrestrial species, this is a trial - and, eventually, an
        opportunity. Because of vanishing resources, the habitat becomes patchy.
        There are refugia, where a more-or-less traditional way of life can be
        maintained, but they don't support very many. These subpopulations
        inbreed because the resources are too scarce to support a journey to
        find another subpopulation. But they tend to mix during the downsizing
        itself, as stray individuals locate remaining subpopulations. Some
        subpopulations may form up entirely from strays.
        Thus an abrupt cooling is likely to provide three (recombination,
        climate stress, parcellation) of the catalysts that speed up
        evolutionary processes. The fourth, re-expanding into empty niches, may
        occur during an abrupt warming (which is accompanied by increased
        rainfall), where pioneers discover new territories of untapped
        resources, and have many offspring survive, even the odder ones.
        
        
        Why Us?
        
 
        
        
        The foregoing is likely true for many mammalian species, not just our
        ancestors. The remaining great apes likely went through this cycle many
        times. What was special about hominids?
        Eating grass indirectly. In the first few years after the great
        forest fires, it is a boom time for the remaining grazing animals, with
        all that grass and all of those succulent shoots. But the waterholes
        would be scarce, and so whole herds would bunch together to visit the
        remaining waterholes. They would lose a few peripheral individuals to
        the predators that lay in wait there.
        What was so special about hominid predators lingering around the
        waterholes? For one thing, upright posture. The grazing animals have
        innate "search images" for four-legged predators; they keep
        their distance. But bipeds can get much closer (as my colleague Arnold
        Towe notes, if you drop down on all fours, grazing animals move away
        promptly). Upright posture may, if efficient enough, allow hominids to
        run animals to exhaustion. But clearly, at some stage, projectile
        predation became a part of the picture (Calvin 1993). Even something so
        simple as flinging a tree branch can be effective in the context of a
        tightly-packed herd at a waterhole: the herd immediately starts to wheel
        around and flee, but the branch lands somewhere in their midst and trips
        an animal or two. Attempts to get back up are delayed by other animals
        stampeding past, and injuries will often occur. In any event, the
        hunters will often have time to run up to a downed animal before it can
        escape. Chimpanzees love to fling branches and they also covet fresh
        meat -- but don't seem to have made the connection, perhaps because of
        lacking savannah waterholes frequented by herds. Following a
        cool-crash-and-burn downsizing, there are lots of temporary savannahs.
        
        
        Growth Curves
        
 
        
        
        Chattering between two climate states thus has the potential to speed
        up evolution, and cool, crash, and burn cycles provide some
        opportunities that our upright australopithecine ancestors might have
        exploited. What, however, makes this an important driver for hominid
        evolution and intelligence, as compared to a bookshelf full of varied
        suggestions on what might have been behind it all?
        Compared, say, to the invention of the carrying bag? The basic idea
        of a carrying bag must have been very important for both gathering and
        for small-game hunting. But one cannot reinvent the carrying bag for
        extra credit. Some inventions can be repeated; for example, the aquatic
        mammals have all discovered that a small reduction in body hair buys
        them greater swimming efficiency. Another reduction buys them even more.
        No matter where along the "growth curve" they are, another
        increment has additional rewards. (There is, however, a limit: you can
        only become so naked.) Some growth curves are also steeper than others,
        faster at driving evolution than slower candidates (which might, like
        many on that bookshelf of plausible candidates, have done the job in the
        long run). So the steepness and extent of growth curves are important
        considerations in sorting out where our intelligence came from.
        There are two aspects of the "eat grass indirectly"
        scenario which have long growth curves, and they involve things where we
        humans have considerably enhanced abilities over the great apes. They
        can also be "pumped" in one percent increments to produce
        many-fold improvements. Neither is that abstraction called
        'intelligence' but either could be the wheelchair-like curb cut that
        gave higher intellectual functions their entry-level jobs.
        
        
        Cooperation and Group Selection
        
 
        
        
        The "good of the group gene" possibility was dismissed a
        few decades ago (see Sober and Wilson 1998) on the theoretical grounds
        that it would, like a leaking tire, backslide. Even if somehow
        concentrated into a subpopulation yielding a majority of cooperators,
        you'd still expect that tendencies to share could be swamped by all the
        non-reciprocating freeloaders, who would out-reproduce the sharers.
        If this were the prime consideration, of course, we would also have
        to conclude that car tires would never work because they all, sooner or
        later, go flat. We just pump them back up occasionally, and the
        cool-crash-and-burn cycle suggests both a concentration mechanism and a
        pump that might allow widespread cooperation to become established for
        long enough to invent other solutions to the freeloader problem.
        There is a certain amount of random recombination of populations
        during the downsizing, as noted earlier. It is not unlike randomly
        selecting a jury of 12 from a jury pool of many hundreds. Even if the
        jury pool is representative (half male, 90 percent right-handed,
        appropriate racial mixtures), some individual juries are dramatically
        different (all of one sex or race, all left-handed, and so forth).
        That's just one of the well-known phenomena of probability theory
        ("drawing small samples without replacement"). So even if
        innate tendencies toward cooperation were only prominent in 10 percent
        of the population, after parcellation many subpopulations would have
        none and some might have a majority.
        These "island" subpopulations are not competing with one
        another like football teams, thanks to those resource-free gaps; they
        are fighting the environment in the cool-crash-and-burn case, and most
        subpopulations disappear with time. When conditions allow populations to
        re-expand, there will be, after expansion into interbreeding
        "continental" populations, a higher proportion of those
        "genes" that helped some subpopulations survive the
        downsizing.
        This, too, is susceptible to the "Why us?" objection as it
        would seem to apply to many mammalian species, and the answer may lie in
        what is shared. Groups with a majority of cooperation genes
        might spend less time arguing (and thereby wasting time which could be
        spent in locating more food) and fighting (thereby both losing time and
        risking injury) during the downsizing. This is particularly attractive
        because of the long growth curve for cooperation, as noted earlier. Or
        what's shared is language, where in order to realize its benefits, you
        might need a sizeable proportion of those with beginner's traits.
        Or, as Derek Bickerton has proposed (see Calvin & Bickerton
        1999), cognitive capacities (mental categories for giver, recipient,
        beneficiaries, type of action, and so on) to keep rough track of
        freeloading tendencies might allow "Who owes what to whom" to
        find another use, namely saying "Who did what to whom."
        Solving the cheater problem in reciprocal altruism could thus be the
        "wheelchair" that paid for the argument-structure scheme of
        handing syntax, until structured language started earning its own way.
        This by itself would constitute a large step up in intelligence.
        
        
        Precision Throwing as Curb Cut
        
 
        
        
        Another ape-to-human improvement with a long growth curve is the
        precision throwing which is so handy for expanding hunting abilities
        beyond that seen in the other predators. Hunting herd animals also has a
        link to cooperation, as any one prey animal is usually too much for one
        hunter to eat; one simply has to give away most of it as the chimpanzees
        do ("tolerated scrounging") and hope for reciprocation when
        someone else gets lucky. As Frans de Waal (1996) observed:
        
          If carnivory was indeed the catalyst for the
          evolution of sharing, it is hard to escape the conclusion that human
          morality is steeped in animal blood. When we give money to begging
          strangers, ship food to starving people, or vote for measures that
          benefit the poor, we follow impulses shaped since the time our
          ancestors began to cluster around meat possessors. At the center of
          the original circle, we find a prize hard to get but desired by
          many... this small, sympathetic circle grew steadily to encompass all
          of humanity -- if not in practice then at least in principle.... Given
          the circle's proposed origin, it is profoundly ironic that its
          expansion should culminate in a plea for vegetarianism.
        
        And, of course, hunting was one of the only solutions to an
        environment where, for a few centuries, you either had to eat grass or
        eat an animal that ate grass.
        The side-of-the-barn accuracy needed for flinging branches into
        waterhole herds may not have much of a growth curve by itself (it
        doesn't matter which one you trip), but it could have gotten hunters
        onto the bottom on the precision-throwing growth curve. Being able to
        hit smaller herds has an even higher payoff. So does throwing from
        farther away (herds eventually become wary), which also reduces risk to
        the hunter. One can use other projectiles, such as rocks. One can become
        accurate enough to hit lone animals. Then there are spears, and their
        augmentation by launching sticks, and so forth. Each improvement has an
        additional payoff: more days that you and your dependents can eat nice
        sterile, high-calorie, low-toxicity fresh meat. (Cooking has made the
        world much safer for vegetarians and scavengers.)
        It's easy to see how natural selection could have repeatedly improved
        throwing, but what does it have to do with higher intellectual function?
        As noted, all of the ballistic movements require much detailed planning
        during "get set" as feedback is too slow. While many ballistic
        movements have some payoffs even when stereotyped, the hominid hunter
        cannot function like the frog throwing its tongue when a fly is heading
        into its "gunsight." There is no standard throw because of the
        "approach distance" problem; each throw is a somewhat novel
        problem in both elevation and range, even if using a standard projectile
        size and weight.
        And, beyond planning and versatility demands, there is the problem of
        timing jitter (Calvin 1983). If you throw at a rabbit-sized target a
        car-length away, and release 5 milliseconds too soon, you'll overshoot
        the target. At two car lengths away, the launch window shrinks
        eight-fold. Redundant motor programs, each with independent noise, can
        solve this double-the-distance problem by using 64 times as many motor
        programs in parallel.
        Even the four-fold increase in the number of neocortical neurons
        during hominid evolution cannot solve this jitter problem (by itself, it
        would only buy a 25 percent increase in approach distance). Many-fold
        increases in parallelism can only be done by temporarily borrowing
        helpers from association cortex during "get set."
        I have developed this jitter-reduction idea elsewhere (Calvin 1983,
        1993, 1996b). What's important for intelligence is to recall Kimura's
        (1993) result from aphasics, that most also suffered from hand-arm
        sequencing problems when confronted with novel sequencing tasks (apraxia)
        - and to recall Ojemann and Mateer's (1979) result from the perisylvian
        core of language cortex, about the overlap of nonlanguage sensory and
        motor sequencing tasks. If there is a common neural sequencing machinery
        for mouth-and-face, hand-and-arm, sensory-and-motor, language cortex is
        an obvious candidate (Calvin & Ojemann 1994).
        And this can explain how a curb cut paid for by natural selection for
        precision throwing might greatly augment planning on other time scales.
        The structured aspect of the higher intellectual functions could easily
        arise from the nested embedding aspect of throwing: the shoulder motion
        is atop the forwards motion of the trunk; planning the elbow rotation
        needs to similarly work from the velocity of the upper arm; the wrist
        flip needs to be planned in light of the prediction of all those
        compounded motions controlling the lower arm's velocity, and so forth.
        All of the coordination must be done in advance, tweaking the parameters
        to find one of the dozens of possible combinations that will hit the
        target amidst a sea of solutions that will miss. This is nested
        embedding of much the same sort as shown in those binary diagrams of
        phrase structure, the other major way of doing syntax (see Calvin &
        Bickerton 1999). As you assemble words to find a coherent sentence to
        speak, you grapple with a problem analogous to novel hand-arm
        sequencing.
        Whatever paid for it in natural selection terms (and I assume it was
        different things at different times), such a multiple-use neural
        sequencer would have major implications for the structured higher
        intellection functions, and thus for intelligence.
        
        
        The Pump Run by Bistable Climate
        
 
        
        
        Pumping up intelligence is thus a real possibility - even though the
        natural selection that paid for it may be as remote as wheelchairs are
        from skateboards. Higher intellectual functions may have some silent,
        nonintellectual partners, those novel ballistic movements.
        Ignoring compound interest considerations for a moment, how many
        strokes on the pump, and of what size, would be sufficient to produce
        the many-fold increases in the mental functions that separate the Pan
        and Homo species? There were several dozen biphasic
        cooling-warming events (each lasting between 70-1500 years) in the last
        ice age alone, between 117,000 years and 11,000 years ago. There were
        dozens of ice ages and, although the high-time-resolution records do not
        extend to cover them yet, cores with thousand-year resolution can pick
        up the longer-lasting ones. These longer-lasting biphasic events have
        now been tracked back to 1.1 million years ago. From what we know of the
        oceanographic mechanisms (see Broecker 1997, 1999; Calvin 1998a), I
        would guess that some will be found between 2 and 3 million years ago --
        but not in the period before the Isthmus of Panama forced the major
        detours in ocean currents.
        So there are hundreds of events of pretty much the same type each
        time: abrupt cooling, crashing populations, and burning ecosystems. This
        suggests that a one percent increment each time might be sufficient.
        However, there were likely many more events (and so even smaller
        increments might suffice) because the typical abrupt cooling or warming
        chattered between modes like an old fluorescent light tube before
        finally settling down in the new mode. Typically, there would be a
        century where the temperature and rainfall whipped back and forth
        between modern and ice-age values a few times, where vast storms churned
        a lot of dust into the atmosphere (the isotopic signature of the dust in
        the Greenland cores suggests that much came from the Great Gobi Desert).
        Such flickering climate would have run the population
        contraction-expansion cycle a few times within a single "madhouse
        century."
        This type of pumping and multiple use shows how big steps up in
        functionality (say, from unstructured protolanguage to structured
        syntax) can arise from a series of small changes in nonintellectual
        functions. It may be that something else from that bookshelf of
        plausible suggestions will prove to run the evolutionary ratchet more
        quickly than my combination of grass, throwing, and cooperation. But if
        we are to ever give an explanation for how an ape can turn into a human,
        we will likely have to address the profound challenges and unusual
        opportunities given our ancestors by the fickle climate.
        
        
        Acknowledgments
        During the two decades that this theory has been
        under construction, I have profited from numerous suggestions. The most
        recent extensions of the theory have benefitted much from a stay at the
        Rockefeller Foundation's Bellagio Center and from workshops organized by
        the LaJolla Origins of Humans group (sponsored by the Preuss Foundation
        and the Mathers Foundation) and by the Center for Human Evolution at the
        Foundation for the Future.
        
        
        
        
        Afterword: The Future's Intelligence Test
        for Humans
        
        
        It has been 8,200 years since an abrupt cooling of even half the
        magnitude discussed here (the Little Ice Age starting about 700 years
        ago was an order of magnitude smaller). Everything we know about the
        geophysical mechanisms (see Broecker 1999, Calvin 1998a) suggests that
        another one could easily happen - indeed, that our greenhouse-effect
        warming could trigger an abrupt cooling in several different ways.
        Because such a cooling would occur too quickly for us to make
        readjustments in agricultural productivity and associated supply lines,
        it would be a potentially civilization-shattering affair, likely to
        cause a population crash far worse than those seen in the wars and
        plagues of history.
        The best understood part of the flip-flop tendencies involves what
        happens to the warm Gulf Stream waters, with the flow of about a hundred
        Amazon Rivers, once they split off Ireland into the two major branches
        of the North Atlantic Current. They sink to the depths of the
        Greenland-Norwegian Sea and the Labrador Sea because so much evaporation
        takes place (warming up the cold dry winds from Canada, and eventually
        Europe, so that it is unlike Canada and Siberia) that the surface waters
        become cold and hypersaline - and therefore more dense than the
        underlying waters. At some sinking sites, giant whirlpools 15 km in
        diameter can be found, carrying surface waters down into the depths.
        Routinely flushing the cold waters in this manner makes room for more
        warm waters to flow far north.
        But this sinking mechanism can fail if fresh water accumulates on the
        surface, diluting the dense waters. The increased rainfall that occurs
        with global warming causes more rain to wall into the oceans at the high
        latitudes. Ordinarily, rain falling into the ocean is not a problem --
        but at these sites in the Labrador and Greenland-Norwegian Seas, it can
        be catastrophic. So can meltwater from nearby Greenland ice cap,
        especially when it comes out in surges. By shutting down the
        high-latitude parts of this "Nordic Heat Pump," these
        consequences of global warming can abrupt change Europe's climate. If
        Europe's agriculture reverted to the productivity of Canada's (at the
        same latitudes but lacking a preheating for winds off the Pacific
        Ocean), 22 out of 23 Europeans would starve.
        The surprise was that it isn't just Europe that gets hit hard. Most
        of the habitable parts of the world have similarly cooled during past
        episodes. Another failure would cause a population crash that would take
        much of civilization with it, all within a decade.
        Ways to postpone such a climatic shift are conceivable, however --
        cloud-seeding to create rain shadows in critical locations are just one
        possibility. Although we can't do much about everyday weather or
        greenhouse warming, we may nonetheless be able to stabilize the climate
        enough to prevent an abrupt cooling.
        Devising a long-term scheme for stabilizing the flushing mechanism
        has now become one of the major tasks of our civilization, essential to
        prevent a drastic downsizing whose wars over food would leave a world
        where everyone hated their neighbors for good reasons. Human levels of
        intelligence allow us both foresight and rational planning. Civilization
        has enormously expanded our horizons, allowing us to look far into the
        past and learn from it. But it remains to be seen whether humans are
        capable of passing this intelligence test that the climate sets for us.