Why are some kiwifruits green when they are ripe? Or avocados or honeydew melons? The answer involves genetic accidents, photosynthesis, hidden pigments, and the words “monkey peach.”

In our kiwifruit fuzziness essay we described how the type and density of trichomes—the hairlike projections from the fruit’s skin that create the fuzziness—in the Actinidia chinensis species complex is correlated with the habitat in China to which a particular population is adapted and the ploidy level of its genome. Only polyploid (having multiple genome copies) Actinidia chinensis occupy the harshest environments—the high, arid reaches of western China—and have the highest trichome density and the longest trichomes. And those fuzzy, resilient, polyploid kiwifruits are all green on the inside (1). They are the plant kingdom’s version of an unshaved vegan after backcountry skiing for a week. The hardy plant had no trouble growing outside its plateau of origin and became the most common commercial kiwifruit in the world (A. chinensis var. deliciosa), followed closely by yellow-fleshed (“golden”), less fuzzy variants of the same species (A. chinensis var. chinensis).

An expanded view of the dozens of Actinidia species reveals orange, red, and purplish pigments that color fruits in the genus. While beautiful, this warm palette strikes me as noteworthy only in contrast to the bright green displayed by the fuzzy A. chinensis var. deliciosa that initially grabbed my attention, and, later, in green kiwiberries (A. arguta). A non-green (for lack of better terminology, “colorful”) ripe fruit, after all, is a common end point for species with fleshy fruit.

Fig. 1 from Crowhurst et al. (2008) of some fruit diversity in the kiwifruit genus Actinidia. We describe the botany and anatomy of kiwifruits in our kiwifruit fuzziness essay.

It is not difficult, however, to bring to mind other examples of species with green-ripe fruit: avocado, green grapes, some citrus, honeydew melon (I’m specifically thinking here of the pericarp or mesocarp tissue under the skin and exclude from this discussion immature fruits that lose their greenness when fully ripe, like green beans and olives). Green ripe fruit, then, in Actinidia and other taxa, seems to me to be something to explain. What, if any, function might it serve, and what are the mechanisms responsible?

While the literature on the subject is far from exhaustive, there is a fairly pedestrian explanation at least for the mechanism, if not any adaptive function, of unusually green fruit flesh outside of Actinidia: fruits start green, and straightforward mutations in a few key genes cause them to remain so. Like that intrepid, hirsute montane vegan, though, Actinidia performs the task a little differently, and it is a bit of a mystery. To understand why that is, we need some backstory on pigments in fruit and how and why they change as fruit ripens, with a focus on Actinidia. Continue reading →