successful gene suddenly became lethal, and the lethal mutant became successful. Over 10,000 years ago, that unconscious selection for nonshat-tering wheat and barley stalks was apparently the first major human 'improvement' in any plant. That change marked the beginning of agriculture in the Fertile Crescent. The second type of change was even less visible to ancient hikers. For annual plants growing in an area with a very unpredictable climate, it HOWTO MAKE AN ALMOND • 121 could be lethal if all the seeds sprouted quickly and simultaneously. Were that to happen, the seedlings might all be killed by a single drought or frost, leaving no seeds to propagate the species. Hence many annual plants have evolved to hedge their bets by means of germination inhibitors, which make seeds initially dormant and spread out their germination over several years. In that way, even if most seedlings are killed by a bout of bad weather, some seeds will be left to germinate later. A common bet-hedging adaptation by which wild plants achieve that result is to enclose their seeds in a thick coat or armor. The many wild plants with such adaptations include wheat, barley, peas, flax, and sunflowers. While such late-sprouting seeds still have the opportunity to germinate in the wild, consider what must have happened as farming developed. Early farmers would have discovered by trial and error that they could obtain higher yields by tilling and watering the soil and then sowing seeds. When that happened, seeds that immediately sprouted grew into plants whose seeds were harvested and planted in the next year. But many of the wild seeds did not immediately sprout, and they yielded no harvest. Occasional mutant individuals among wild plants lacked thick seed coats or other inhibitors of germination. All such mutants promptly sprouted and yielded harvested mutant seeds. Early farmers wouldn't have noticed the difference, in the way that they did notice and selectively harvest big berries. But the cycle of sow / grow / harvest / sow would have selected immediately and unconsciously for the mutants. Like the changes in seed dispersal, these changes in germination inhibition characterize wheat, barley, peas, and many other crops compared with their wild ancestors. The remaining major type of change invisible to early farmers involved plant reproduction. A general problem in crop development is that occasional mutant plant individuals are more useful to humans (for example, because of bigger or less bitter seeds) than are normal individuals. If those desirable mutants proceeded to interbreed with normal plants, the mutation would immediately be diluted or lost. Under what circumstances would it remain preserved for early farmers? For plants that reproduce themselves, the mutant would automatically be preserved. That's true of plants that reproduce vegetatively (from a tuber or root of the parent plant), or that are hermaphrodites capable of fertilizing themselves. But the vast majority of wild plants don't reproduce I 2. 1 •GUNS,GERMS, AND STEEL that way. They're either hermaphrodites incapable of fertilizing themselves and forced to interbreed with other hermaphrodite individuals (my male part fertilizes your female part, your male part fertilizes my female part), or else they occur as separate male and female individuals, like all normal mammals. The former plants are termed self-incompatible hermaphrodites; the latter, dioecious species. Both were bad news for ancient farmers, who would thereby have promptly lost any favorable mutants without understanding why. The solution involved another type of invisible change. Numerous plant mutations affect the reproductive system itself. Some mutant individuals developed fruit without even having to be pollinated, resulting in our seedless bananas, grapes, oranges, and pineapples. Some mutant hermaphrodites lost their self-incompatibility and became able to fertilize themselves—a process exemplified by many fruit trees such as plums, peaches, apples, apricots, and cherries. Some mutant grapes that normally would have had separate male and female individuals also became self-fertilizing hermaphrodites. By all these means, ancient farmers, who didn't understand plant reproductive biology, still ended up with useful crops that bred true and were worth replanting, instead of initially promising mutants whose worthless progeny were consigned to oblivion. Thus, farmers selected from among individual plants on the basis not only of perceptible qualities like size and taste, but also of invisible features like seed dispersal mechanisms, germination inhibition, and reproductive biology. As a result, different plants became selected for quite different or even opposite features. Some plants (like sunflowers) were selected for much bigger seeds, while others (like bananas) were selected for tiny or even nonexistent seeds. Lettuce was selected for luxuriant leaves at the expense of seeds or fruit; wheat and sunflowers, for seeds at the expense of leaves; and squash, for fruit at the expense of leaves. Especially instructive are cases in which a single wild plant species was variously selected for different purposes and thereby gave rise to quite different-looking crops. Beets, grown already in Babylonian times for their leaves (like the modern beet varieties called chards), were then developed for their edible roots and finally (in the 18th century) for their sugar content (sugar beets). Ancestral cabbage plants, possibly grown originally for their oily seeds, underwent even greater diversification as they became variously selected for leaves (modern cabbage and kale), stems (kohlrabi), buds (brussels sprouts), or flower shoots (cauliflower and broccoli). So far, we have been discussing transformations of wild plants into HOWTO MAKE AN ALMOND • 1x3 crops as a result of selection by farmers, consciously or unconsciously. That is, farmers initially selected seeds of certain wild plant individuals to bring into their gardens and then chose certain progeny seeds each year to grow in the next year's garden. But much of the transformation was also effected as a result of plants' selecting themselves. Darwin's phrase 'natural selection' refers to certain individuals of a species surviving better, and / or reproducing more successfully, than competing individuals of the same species under natural conditions. In effect, the natural processes of differential survival and reproduction do the selecting. If the conditions change, different types of individuals may now survive or reproduce better and become 'naturally selected,' with the result that the population undergoes evolutionary change. A classic example is the development of industrial melanism in British moths: darker moth individuals became relatively commoner than paler individuals as the environment became dirtier during the 19th century, because dark moths resting on a dark, dirty tree were more likely than contrasting pale moths to escape the attention of predators. Much as the Industrial Revolution changed the environment for moths, farming changed the environment for plants. A tilled, fertilized, watered, weeded garden provides growing conditions very different from those on a dry, unfertilized hillside. Many changes of plants under domestication resulted from such changes in conditions and hence in the favored types of individuals. For example, when a farmer sows seeds densely in a garden, there is intense competition among the seeds. Big seeds that can take advantage of the good conditions to grow quickly will now be favored over small seeds that were previously favored on dry, unfertilized hillsides where seeds were sparser and competition less intense. Such increased competition among plants themselves made a major contribution to larger seed size and to many other changes developing during the transformation of wild plants into ancient crops. What accounts for the great differences among plants in ease of domestication, such that some species were domesticated long ago and others not until the Middle Ages, whereas still other wild plants have proved immune to all our activities? We can deduce many of the answers by examining the well-established sequence in which various crops developed in Southwest Asia's Fertile Crescent. It turns out that the earliest Fertile Crescent crops, such as the wheat 124 ' GUNS, GERMS, ANDsteel
