Category Archives: Fruit

Botany lab of the month – August edition: Rocky Top Corn Soup

It’s August, and everyone from the American Midwest knows that late summer means fresh sweet corn, and a lot of it. When I was growing up in Indiana, every few days during corn season we would pick up a dozen ears from my family’s favorite roadside stand, just hours after harvest, and cook them right away, before the kernels could start converting their sugar into starch. 

Corn season typically peaked the final week of the Indiana State Fair, which always fell between my sister’s birthday and mine. We felt like the whole world was celebrating with us since the Indiana State Fair really is just a giant party, with rides and games and food and music and anatomically impressive hogs. So every year we went to revel in the indulgent atmosphere of the fair alongside thousands of unbridled Hoosiers from all over the state sweating in tank tops and showing off their best pickles. But I most looked forward to the corn. The fair was littered with vendors serving huge ears of fresh-picked local corn straight from a grate set up over a large open flame. As soon as the charred husks were cool enough to peel back into a handle, we sidled up next to our fellow Hoosiers at a trough (literally a trough) of hot melted butter and plunged the roasted ears into it, right up to the hilt. Then we gave them a generous coating of salt from oversized aluminum shakers, passed from hand to greasy hand around the trough and down the line to us. Best birthdays ever.

For this month’s botany lab, I have created a cold summer soup that is as much a celebration of decadent State Fair food as an homage to the millennia of cultivation and adaptation that makes that food possible. The soup features corn four ways, and its various ingredients are available to us only because people have been growing corn (more accurately, maize) and creating distinct varieties for a very long time over an unusually wide geographical area. Maize as a crop goes back to Mesoamerica about 9 thousand years ago, and it had become a substantial part of people’s diets there by about 4500 years ago (Kennet et al. 2020). The spread of maize throughout much of North America was slow because the right mutations had to arise before Native Americans could select for genotypes that were well suited to the local day length, season length, and altitude (Doebley 1990). In the process, people created the varieties we know now, including those in the soup: sugary sweet corn, eaten while immature; dent corn, with soft starchy kernels good for grinding into fine cornmeal, masa, and grits; and flint, with hard round starchy kernels that can be popped or ground into polenta.

Photo of a bowl of cold corn soup with bourbon-infused grits croutons and popcorn
Cold corn soup with bourbon-infused grits croutons and popcorn

The Indiana State Fair and its unabashed celebration of big agriculture most probably sits atop ancient hand-tended corn fields. It is important to recognize that the Fairgrounds occupy land that was traditionally held by the Miami Tribe of Native Americans and lost through a series of coercive treaties in the 19th century. Many members were forced to relocate, but not all, and the Miami Nation of Indiana maintains a cultural presence in the state. The many indigenous tribes throughout the region have a long tradition of agriculture, and as far back as a thousand years ago people would have made corn (Zea mays) the main staple of their diet (Emerson et al. 2020).

Now that corn is big business in the US, dent corn is the most widely grown type because it is used for animal feed, corn oil, high fructose corn syrup, and the many processed foods that have become modern dietary staples. The luckiest dent corn, however, ends up as mash to be fermented into bourbon, which brings me to the name of the soup.

photo of the horse Rocky Top. Credit Sara Baggett Preston
Good old Rocky Top. Photo: Sara Baggett-Preston

When I was about to turn 13, my mother, who grew up riding, brought a horse into the family. He was a big palomino named Rocky Top, with a mane the color of corn silk and some Tennessee Walker genes that occasionally showed themselves. Naturally, the Osborne Brothers’ song “Rocky Top” was a favorite in our house, especially since my father was, as he described himself, a “tolerable” bluegrass musician who could sometimes be coaxed into singing it.

Corn won’t grow at all on Rocky Top,
Dirt’s too rocky by far.
That’s why all the folks on Rocky Top
Get their corn from a jar

In the song, Rocky Top is an idyl somewhere down in the Tennessee hills. Personally, when I drink corn from a jar, I choose a nice aged bourbon from Kentucky, but that’s beside the point. As long as it’s mostly corn, with just a bit of rye and barley, I’m happy.

This cold late-summer soup features corn four ways, with different flavors and botanical properties that will be obvious as you prepare it. Observation notes are included in the recipe. Like vendors at the Indiana State Fair, this soup plays around in the corners of midwestern and southern food traditions and comes up with something new and delicious. The fresh raw corn base is almost dessert like while croutons of bourbon-soaked grits add intense corn flavor infused with smoke and caramel, and a scattering of buttered popcorn balances the sweet creamy texture of the soup with salty crunch. The soup’s name, of course, honors the inimitable Rocky Top, who was a beloved equine member of our family for over thirty years. The croutons and popcorn might even remind you of rocky outcrops in the Tennessee hills.

Rocky Top soup with corn four ways

Serves 4-6

The preparation is simple, but leave ample time to cook the stock and grits and to chill the ingredients at various stages.

Ingredients

6 large or 8 medium ears of fresh sweet corn in the husk, preferably from a local farm stand or farmers market and as recently picked as possible

1/2 cup of uncooked grits, stone-milled if available, but definitely not “quick”

1/2 cup bourbon (corn in a jar)

1/4 cup unpopped popcorn

Salt (both regular and smoked, if you’ve got it)

Black pepper

White pepper (optional)

1 stick of butter (or olive oil for a vegan version)

Cold soup base

This part of the recipe is quick; my observation notes are long.

1. Before removing the husks, use a large knife to cut the top inch or two from each ear. If insect larvae have gotten into the ears, this is where they will be, and you may prefer not to see them.

2. Remove the husks, which are leaves enclosing the bud of a giant flowering stalk. Notice their shape and the arrangement of their veins, then take a look at the short bit of stem at the base of the ear to see its scattered vascular bundles. Corn (Zea mays) is typical of monocots in having long narrow leaves with parallel veins and vascular bundles scattered throughout its stems (not in a clear ring). Wash and save the tender inner layer of husks for the stock.

Click to enlarge

3. Look closely at the corn silk and notice that the strands seem to originate from between the kernels. That is because each one is (or was) attached to the top side (the side facing the tip end) of a single kernel. Amazingly, a strand of silk is the extremely long stigma and style through which germinated pollen grains traveled to fertilize the cells inside what is now a kernel. Sometimes you can see that the silk strand is rough or slightly hairy on one side, the better to capture pollen with. (For more details, see Jeanne’s terrific essay, Super Styled.)

Corn silk emerges from between kernels
Corn silk emerges from between kernels
Close up of corn silk, showing pollen-catching hairs
Close up of corn silk, showing pollen-catching hairs. Click to enlarge.

4. Rub the ear under water to remove as much silk as possible. Their race is run, and their job is done.

5. Take a moment to appreciate that each kernel of corn on a cob was once a flower, embedded alongside other flowers in a thick flowering axis. The flowers never had functioning sepals or petals or stamens. (Separate stamen-bearing flowers make the pollen and are found in a tassel on top of the plant). Each flower was essentially a single pistil: an ovary with a style and stigma (those silks!) long enough to protrude beyond the husks where it pollen could find it. The mature product of an ovary is a fruit, so it follows that a kernel of corn is a fruit, not a seed. The fruit functions as a seed, however, because it is essentially just a thin wall fused tightly to the single large seed inside. This type of fruit is called a caryopsis (or, simply, a grain). Fresh corn on the cob is immature, and the fruits are soft, but they would become hard if allowed to mature.

The attachment points of the corn silks (super long styles and stigmas!) are clearest where kernels are uncrowded, near unpollinated flowers. Click to enlarge

6. Cut the kernels off the cobs and into a bowl. First, cut the cobs in half, then stand them on their cut ends and run a large knife down the sides to remove the kernels. When all kernels have been removed, pull the knife blade across the cobs over the bowl to pull out any residual “milk” (actually endosperm, described below).

kernels cut from a cob to show embryo
The corn embryo is embedded in sweet soft endosperm. The kernel is surrounded by a fringe of paleas, lemmas, and glumes.

7. Notice the very small opaque flattened round structures that pop out of the cut kernels and milked juice. These are embryos. The milky juice is endosperm, the tissue that would supply the embryo with energy and nutrients during germination. At this stage, most of the endosperm is soft and some is still liquid. The liquid portion contains many nuclei because it has not yet been divided into walled cells. Refrigerate the bowl of kernels until the stock has been made and cooled.

8. Look closely at one of the empty cobs. Notice that the sockets that once held kernels are ringed with short papery ruffles. These structures – paleas, lemmas, and glumes– are evidence that the corn cob is much more complicated than it seems. Those empty sockets held not one but two corn flowers, one of which simply never developed. The flower pairs were borne on a very tiny branch with two short glumes at its base. Each flower, in turn, was enclosed by a pair of thin structures, one palea and one lemma. The raggedy ruffles are glumes and paleas and lemmas and are left behind when you eat corn off the cob or shave off the kernels with a knife.

9. Place the cobs and the reserved tender husks into a saucepan, add water to barely cover them, and simmer for about 30 minutes to make a stock. Remove the cobs and husks and allow the stock to cool to room temperature.

10. Move the kernels and milky endosperm from the bowl into a blender and add about 1/4 cup of the stock. Blend very well until the soup is a fine silky purée. Add small amounts of the stock as needed to ease the blending and achieve the consistency you prefer. Salt very sparingly; you want to retain the grassy sweetness of the raw fresh corn. Chill thoroughly.

This soup should be made no more than a day ahead for peak flavor. It is raw and may start to ferment after several days.

Grits croutons

In a pinch, you can use polenta, but the texture will be less interesting and the cuisine will be less American. 

1. Place the grits and the bourbon in a heavy-bottomed saucepan, and allow the grits to soak for 30 minutes.

2. Notice that grits, especially traditional stone milled grits, vary much more in particle size than polenta does. Could that be why most people treat “grits” as plural and “polenta” as a singular mass noun? It won’t be obvious, but grits are generally made of dent corn and polenta is made of harder flint corn.

4. Add a tablespoon of butter, a teaspoon of salt, and several grinds of white pepper (about 1/4 teaspoon). If you have smoked salt available, use it here.

3. Cook according to the directions for your particular grits. If you have leftover corn cob stock, use it, supplementing with water if needed.

4. Continue to cook and stir the grits until they are thoroughly done and very thick, like mashed potatoes. You may need to cook longer than directed to get the grits thick enough.

5. Spread the grits into a couple of buttered loaf pans or a square cake pan and chill them for at least an hour, until they are well set. The grits should be about half an inch thick in the pan.

6. Use a table knife to cut the chilled grits into one-inch squares and turn them out into a roasting pan. Toss them with soft butter or olive oil and bake them at 350º for 20 minutes or until crispy on the outside. (It is also possible to fry them in a pan, but they tend to fall apart because they are more fragile than polenta squares.)

7. Croutons should be room temperature or warm but not hot when you serve the soup. You want to keep the soup cool. Croutons may be reheated if needed.

Popcorn topping

1. Notice that popcorn is hard because it has been allowed to mature before harvest, and the once liquid endosperm has been divided into separate cells containing the previously free floating nuclei. Popcorn is a variety of flint corn, so it has a round end without the depressed center seen in dent corn. The nutritive endosperm of popping corn is much starchier than sweet corn would ever get. Although popcorn seems very dry, there is some residual water inside. When heated, that water becomes steam and swells the starch until it bursts the kernel open.

2. Pop the popcorn using your favorite method. Admire the fluffy white expanded endosperm.

2. Butter and salt the popcorn generously, using about 4 tablespoons of melted butter.

Assembling the soup

1. Pour the chilled soup into bowls

2. Scatter several grits croutons over the soup. Most will sink

3. Top generously with popcorn and grind some black pepper over the top

4. Serve immediately to maintain the contrasts in temperature and texture

References and resources

My favorite sources of stone-milled grits are Anson Mills and Nora Mills. Anson Mills also sells popcorn and their website has several interesting pages about the history and botany of the foods they mill.

Doebley, J. (1990). Molecular evidence and the evolution of maize. Economic Botany, 44(3), 6-27.

Doebley, J. F., Goodman, O. M., & Stuber, C. W. (1986). Exceptional genetic divergence of northern flint corn. American Journal of Botany, 73(1), 64-69.

Emerson, T. E., Hedman, K. M., Simon, M. L., Fort, M. A., & Witt, K. E. (2020). Isotopic confirmation of the timing and intensity of maize consumption in greater Cahokia. American Antiquity, 85(2), 241-262.

Kennett, D. J., Prufer, K. M., Culleton, B. J., George, R. J., Robinson, M., Trask, W. R., … & Gutierrez, S. M. (2020). Early isotopic evidence for maize as a staple grain in the Americas. Science Advances, 6(23), eaba3245.

Nickerson, N. H. (1954). Morphological analysis of the maize ear. American Journal of Botany, 87-92.

Weatherwax, P. (1916). Morphology of the flowers of Zea mays. Bulletin of the Torrey Botanical Club, 43(3), 127-144.

Mberkery1, CC BY-SA 4.0 https://creativecommons.org/licenses/by-sa/4.0, via Wikimedia Commons

The Botanist Stuck in the Kitchen With You (and Peas)

I am about to start an 8th week of online teaching and my county’s 11th week of sheltering in place. While the (essential and life saving) sheltering is getting really old, the academic quarter has sped by as usual, along with its relentless parade of deadlines and grading. Our current crisis may have no definite end, but the academic quarter must wrap up on time, ready or not.

Some people are reporting really vivid dreams right now, however, for me, the most noticeable side effect of working and teaching from home is that I never stop thinking about it. Like midway through a Saturday night screening of Reservoir Dogs when I was suddenly reminded of peas and the upcoming class meeting on fruit. Can I do this online? We’ll just have to see, won’t we?

Oh, and don’t be a Mr. Pink.

Apologies to Stealers Wheel, the terrific Michael Madsen, and his PSA on sheltering.

peeeees.003.jpeg

Closeup of sugar snap pea flower with tiny developing fruit.

Sage, rosemary, and chia: three gifts from the wisest genus (Salvia)

This essay is our annual contribution to the Advent Botany essay collection curated by Alastair Cullham at the University of Reading. We highlight three charismatic species in the large genus Salvia (in the mint family, Lamiaceae): rosemary, sage, and chia.

Two Christmases ago we pointed out the current fad in decorating pineapples for Christmas. This year, some of our gentle readers may come across potted rosemary bushes that are trimmed into a cone to resemble a conifer. These are pleasant and ostensibly can be kept alive after the holiday season.

A rosemary shrub trimmed into a conifer shape. Photo from Pottery Barn.

A perhaps less pleasant holiday botanical encounter may include a Christmas tree-shaped Chia Pet.

Christmas tree Chia Pet. Photo from Amazon.

As far as Chia Pets go, this one is fairly innocuous. In my view, however, its only saving grace is that the chia plant itself is a fabulous taxon (Salvia hispanica), as is the rest of its large genus, Salvia, which also happens to include rosemary (Salvia rosmarinus). Rosemary of course is much more likely to make a holiday appearance as a culinary ingredient than a decoration, lovely as it is. In the kitchen it is frequently joined with its congener Salvia officinalis, usually just called garden sage. That the genus Salvia is responsible for half the taxa in the title of a Simon & Garfunkel album (Parsley, Sage, Rosemary and Thyme), notwithstanding that Art Garfunkel looks like a Chia Pet on the cover, could provide enough taxonomic joy to justify leaving this examination of these plants here. The name “sage”, however, implies wisdom, and so like the wise men of old, I shall persevere.

Parsley, Sage, Rosemary and Thyme album cover by Simon & Garfunkel (1966)

We’ll start by addressing the taxonomic elephant in the room that might otherwise distract learned readers: rosemary was only brought into the Salvia fold in 2017. Before then it was in its own small genus: Rosmarinus. The reason Rosmarinus is now Salvia is that the speciose Salvia was found to be paraphyletic: the pre-2017 conscription of the nearly 1000 species in the genus did not include all of the descendants of their most recent common ancestor. When the relationships between all the Salvia species and their closest relatives were plotted on a single phylogenetic tree, it was obvious that Rosmarinus and a few other genera should more naturally be considered Salvia, and Salvia was revised accordingly.

Rosemary (Salvia rosmarinus)

Another taxonomic bookkeeping item is to clarify that the sages in Salvia are only distant relatives of the sagebrushes and sageworts in the genus Artemisia, which is in the sunflower family Asteraceae (please see our Artemisia essay for more information about that genus, which includes the herb tarragon). The phylogenetic relationships of the major groups in Salvia from the most recent revision (Drew et al., 2017) is shown below.

Figure 2 from Drew et al. (2017): “(A) Composite chronogram of subtribe Salviinae (which contains Salvia and related taxa) based on chloroplast DNA sequences from previous molecular phylogenetic analyses. Asterisks denote nodes with low support and/or conflicting resolution among previous analyses. Salvia nomenclature follows subgeneric clades described here, including three tentatively named clades that await proper circumscription. Calibrations based on Drew & Sytsma (2012; See supplementary figure S4) (B) Circle cladogram framed on larger chronogram with weakly supported nodes collapsed, depicting species diversity and generalized staminal types within each clade of Salvia; modified after Walker & Sytsma (2007) and Walker et al. (2015).” S. elegans (pineapple sage), S. sclarea (clary sage), and S. hispanica (chia) are in the American subgenus Calosphace. Rosemary is in its own subgenus, Rosmarinus.

The phylogenetic diagram above (from Drew et al., 2017) shows locations where the flower anther structure evolved into a lever-like mechanism that aids in bee pollination by physically moving the two stamens into contact with the bee’s back when a bee enters the flower (see illustration below from Walker, Sytsma, Treutlein, & Wink, 2004).

Figure 2 from Walker et al 2004: “Flower and pollination of Salvia pratensis (Salvia clade I). A flower without the lever mechanism activated (A). As the pollinator enters the flower (B), the pollen is deposited on the back of the pollinator. As the pollinator enters an older flower (stamens removed from sketch, but remain present in flower) pollen is transferred (C). The posterior anther thecae forming the lever can be fused or free and in the subg. Leonia, produce fertile pollen”

The lever mechanism independently evolved three times within Salvia. Each of these evolutionary events was followed by rapid and prolific speciation driven by this innovation in pollination biology (Drew et al., 2017): the advent of the lever mechanism led to the radiation of around 500 species in the subgenus Calosphace in Central and South America; around 250 species evolved soon after the advent of the lever mechanism in the Salvia officinalis clade in the Mediterranean and Western Asia; and around 100 species radiated following the lever in Far East Asia in the Salvia glutinosa clade.

Sage (Salvia officinalis) flowering on my deck this summer

The bee-pollinated Salvia flowers are distinct from those pollinated by hummingbirds, which are more elongate and often red, like the flowers of pineapple sage (S. elegans), and have either evolutionarily lost the staminal lever mechanism or never had it in the first place.

Pineapple sage (Salvia elegans)

The parsley, sage, rosemary, and thyme made famous by Simon & Garfunkel started their culinary careers in Europe. All but parsley are in the mint family (Lamiaceae; see our carrot top essay for a discussion of fun chemical relationships between the flavor compounds in the mint family and the parsley family, Apiaceae). This points to the profusion of aromatic mint family species common to the rocky shrublands covering much of Europe and western Asia (Rundel et al., 2016; Vargas, Fernández-Mazuecos, & Heleno, 2018).

Called “tomillar” in spanish, literally a field of wild thyme (Thymus vulgaris) and associated species growing in the Orusco de Tajuña hills (near Madrid. Spain). Other edible Lamiaceae can be found in this plant community, including Salvia rosmarinus, and Lavandula latifolia (a lavendar). Photo by Julia Chacón-Labella.

That broad area is one of the centers of Salvia species diversity, but the genus is globally widespread. The genus probably originated and dispersed first from African and then the Mediterranean (see the figure of Salvia distribution and putative dispersal history below from Will & Claßen-Bockhoff, 2017), but the full story of dispersal and species radiation within the genus requires more elucidation.  Numerous species of Salvia are utilized as culinary or medicinal herbs or garden ornamentals throughout its range.

Fig. 8 from Will et al. 2017: “Salvia s.l. in time and space. A: Distribution of Salvia s.l., putative migration routes and fossil sites; BLB = Bering Land Bridge; D = Dorystaechas; M= Meriandra; NALB = North Atlantic Land Bridge; P = Perovskia; R = Rosmarinus; Z = Zhumeria; white arrows indicate repeated colonization of S Africa and dispersal from the Eastern Cape to Madagascar; hatched arrows (dark grey) indicate the repeated colonization of the Canary Islands from two different mainland sources; red arrow illustrate the dispersal from East Asia to Eurasia reflected by S. glutinosa; black arrows correspond to dispersal events from the OW to America reflected by two distinct lineages; ? = route uncertain; template of the map provided by the German earth science portal (www.mygeo.info). B: Simplified phylogenetic tree; nodes discussed in the text are indicated by capital letters; colors reflect distribution areas. (For interpretation of the references to color in this figure legend, the reader is referred to the web version of this article.)”

The phylogeny above shows the large number of American taxa in subgenera Calosphace and Audibertia. While many of these species have also been used as aromatic herbs and traditional medicines, the most famous of the American Salvias, chia, is known for its nutritious seeds (Jenks & Kim, 2013a). Chia is a name given to two species of Salvia: S. columbariae and S. hispanica. S. columbariae ranges from southern California to central Mexico, at which point the range of S. hispanica begins and extends to Guatemala. Indigenous groups throughout that range historically used both species of chia as a pre-Columbian staple food source. The Aztecs cultivated it, and 16th century Spanish codices indicate it may have been as widely utilized as maize (Cahill, 2003).

Chia nutlets (S. hispanica) and a dried sage (S. officinalis) leaf for scale

Technically, the chia “seeds” you can buy in the store (or harvest yourself) are fruits. The Salvia fruit, like those of all mint family species, is called a schizocarp. The ovary inside the flower has four chambers, called locules. Each locule matures into an independent, indehiscent nutlet. The shell (pericarp) of the nutlet is stratified into the same categories of outer fruit layers as are more familiar fleshy fruits (cuticle, epicarp, mesocarp, endocarp; see our pomegranate or apple essay for more details about fruit structure), but in the Salvia nutlet the outer fruit layers are dry and compressed and inseparable from the single seed inside the fruit (Capitani, Ixtaina, Nolasco, & Tomás, 2013). Salvia nutlets mature inside of papery fused calyces (see the photo below of sage nutlets and their cup-like persistent calyces).

Sage (Salvia officinalis) leaves and nutlets inside of papery, fused persistent calyces.

The word “chia” is derived from the Aztec language Nahuatl word for “oily,” a name bestowed because chia seeds do have a high oil content (Cahill, 2003). Chia oil is rich in the omega-3 fatty acid alpha-linolenic acid, which has contributed to its recent fame as a modern health food. High alpha-linolenic acid content may be a general feature of the genus: other Salvia species, including S. officinalis, garden sage, have been shown to have high alpha-linolenic acid content in their seeds (Ben Farhat, Chaouch -Hamada, & Landoulsi, 2015).

Chia nutlets are also known for the gooey mucilage they exude when wet. This polysaccharide matrix is used as a food binder and thickener (Google “vegan egg replacement”). The production of mucilaginous diaspores (the dispersing agent, a fruit or a seed) is called myxocarpy. As Katherine discusses in her essay on okra, the flagship mucilaginous food plant, the purpose of the mucilage is likely water retention in the arid regions where these plants tend to come from. The mucilage might also act as a glue to bind the nutlet to the soil or to a dispersing animal’s fur—or to the terracotta substrate of a Chia Pet. Myxocarpy is most common in plants with small seeds growing in dry, arid areas, like those where Salvia species have radiated (Ryding, 1992).

Sage growing in coastal California, a Mediterranean-type ecosystem

Within the mint family, myxocarpy only occurs in the subfamily Nepetoideae. The subfamily, incidentally, gets its name from the catnip genus, Nepeta. Most if not all of the familiar edible herbs from the mint family are in this subfamily. Katherine has taken advantage of myxocarpy in this clade by serving soaked black basil (Ocimum basilicum) nutlets as a basil-scented vegan “caviar.”

Cat in the catnip (Nepeta cataria)

Salvia aroma and flavor–and I think the psychoactive properties of catnip for cats and known hallucinogen Salvia divinorum–comes from the terpenoids and phenolics that comprise their essential oils. The terpenoids are synthesized and stored in special glandular trichomes on the leaf surface (Schuurink & Tissier, 2019). Trichomes are hair-like extensions of the epidermis, although the glandular trichomes full of essential oil look more like water balloons than hair. Salvia species have other types of trichomes in addition to the glandular trichomes that are indeed much more hair-like and give the leaves of some Salvia a downy or prickly appearance (Kamatou et al., 2006).

Scanning electron micrograph (SEM) of a rosemary leaf. Spherical oil-filled glandular trichomes are found amongst the branched hair-like trichomes covering the lower surface of the leaf, which has a greater profusion of hairs and glands than the upper surface. When the glands are damaged or broken the aromatic essential oil is released. Magnification: x1550 (x381 at 10cm wide). Photo from https://psmicrographs.com/sems/flowers-plants/

We discussed trichome function extensively in one of our kiwi essays. The hair-like trichomes may serve the leaf by protecting it from excess solar radiation and wind and otherwise creating a more mild microclimate at the leaf surface to help it retain water.

Rosemary

Terpenoid biosynthesis requires numerous steps in which intermediate chemical products are modified by a series of specific enzymes and other proteins. Small changes in the genes responsible for those proteins can lead to big qualitative changes in the final terpenoid mix in the essential oil of a given taxon. We mammals are adept at discerning aroma differences between chemically similar terpenoids. For example, in on our carrot top essay we discussed the case of spearmint and caraway. The respective versions of the terpenoid carvone that characterize the essential oils of those plants differ only in the physical configuration of the same chemical elements, but they smell radically differently to us.

Clary sage (S. sclarea)

The function of the essential oil in the glandular trichomes, however, is not to improve human well being. Plants synthesize those lovely terpenoids as chemical defense against insect herbivores and microbial pathogens.  When the hair-like trichomes fail to stop the intruders, the glandular trichomes will explode on contact, drenching the would-be attackers in a caustic-but-fragrant deluge.

rosemary

The pharmacopeia of terpenoid aromas present in the mint family—bring to mind the scents of sages, rosemary, lavender, peppermint, spearmint, savory, thyme, oregano, marjoram, shiso, basil—owes its evolutionary origins certainly in part at least to the various selection pressures imposed on those herbal taxa by their pests. Within even commonly grown domesticated Salvia species, essential oil constituent variation leads to dramatic differences in aroma. For example, consider the differences among rosemary, garden sage, clary sage (S. sclarea), and pineapple sage (Salvia elegans), which has a notably fruity smell. The fruitiness is due in part to the presence of the terpenoids charcteristic of citrus, which are widespread across plants.

Garden sage (Salvia officinalis)

The Roman historian and natural scientist Pliny the Elder coined the name Salvia, which is derived from the Latin salvare, meaning to heal and save, and salvus, meaning uninjured or whole. The common English name “sage” of these plants ultimately comes from this same Latin root. In Pliny the Elder’s time, the Mediterranean Salvia species were considered healing herbs, good for treating colds and a variety of ailments. Salvia feature prominently in the ethnomedicine of every region in which it is found (South Africa: (Kamatou et al., 2006); Central and South America: (Jenks & Kim, 2013b)). There is a Chinese proverb that asks “How can a man grow old who has sage in his garden?” I do not know which Salvia species would have been responsible for this proverb. There are over a hundred species of Salvia species native to China, and the Mediterranean import Salvia officinalis is grown throughout the country.

Bundle of dried sage, recently, recently, in Alaska

The health and wellness meaning of “sage” is etymologically independent from its other definition as a wise thing or wise person. This second meaning ultimately comes from the Latin sapere, to know or taste. I personally enjoy conflating these meanings, tying wisdom and well-being to the plant. I like that the Salvia officinalis that grew on a pot on my deck this summer and that will season comfort food this winter is a descendent from the plants that healer contemporaries of Pliny the Elder would have searched for amidst sun-drenched rocks in the Mediterranean hills.

Salvia in macarons at my local bakery (Fire Island) this week: blackberry-sage and rosemary-merlot.

Simon & Garfunkel close the Parsley, Sage, Rosemary and Thyme album with the song “7 O’Clock News/Silent Night,” in which they juxtapose jarring newscasts from the Nixon and Johnson era with the Christmas carol. This holiday season has felt a bit like that song to me, like concerted effort is required to prevent awful, omnipresent news from drowning out the joy and solemnity of marking the darkest time of the year. But perhaps honoring traditions always involves this element of deliberately carving out the space in which to do so. Perhaps sprinkling rosemary and sage into a holiday stew or stuffing can be a radical act, a defiant embrace of old wisdom to fortify ourselves to stand with each other and create something beautiful in the cold. Regardless, insane amounts of butter will be involved, at least at my house. And when the January 2nd resolutions to “eat better” come around, chia will be there.

References

Ben Farhat, M., Chaouch -Hamada, R., & Landoulsi, A. (2015). Oil yield and fatty acid profile of seeds of three Salvia species. A comparative study. Herba Polonica, 61(2), 14–29. doi:10.1515/hepo-2015-0012

Cahill, J. P. (2003). Ethnobotany of Chia, Salvia hispanica L. (Lamiaceae). Economic Botany, 57(4), 604–618. doi:10.1663/0013-0001(2003)057[0604:EOCSHL]2.0.CO;2

Capitani, M. I., Ixtaina, V. Y., Nolasco, S. M., & Tomás, M. C. (2013). Microstructure, chemical composition and mucilage exudation of chia ( Salvia hispanica L.) nutlets from Argentina. Journal of the Science of Food and Agriculture, 93(15), 3856–3862. doi:10.1002/jsfa.6327

Drew, B. T., González-Gallegos, J. G., Xiang, C. L., Kriebel, R., Drummond, C. P., Walker, J. B., & Sytsma, K. J. (2017). Salvia united: The greatest good for the greatest number. Taxon, 66(1), 133–145. doi:10.12705/661.7

Jenks, A. A., & Kim, S. C. (2013a). Medicinal plant complexes of Salvia subgenus Calosphace: An ethnobotanical study of new world sages. Journal of Ethnopharmacology, 146(1), 214–224. doi:10.1016/j.jep.2012.12.035

Jenks, A. A., & Kim, S. C. (2013b). Medicinal plant complexes of Salvia subgenus Calosphace: An ethnobotanical study of new world sages. Journal of Ethnopharmacology, 146(1), 214–224. doi:10.1016/j.jep.2012.12.035

Kamatou, G. P., van Zyl, R. L., van Vuuren, S. F., Viljoen, A., Figueiredo, A. C., Barroso, J. G., … Tilney, P. M. (2006). Chemical composition, leaf trichome types and biological activities of the essential oils of four related Salvia Species indigenous to Southern Africa Analysis of plant volatile using 2D gas chromatography View project Chemometrics View project. Journal of Essential Oil Research. Retrieved from https://www.researchgate.net/publication/236850867

Rundel, P. W., Arroyo, M. T. K., Cowling, R. M., Keeley, J. E., Lamont, B. B., & Vargas, P. (2016). Mediterranean Biomes: Evolution of Their Vegetation, Floras, and Climate. Annual Review of Ecology, Evolution, and Systematics, 47, 383–407. doi:10.1146/annurev-ecolsys-121415-032330

Ryding, O. (1992). Pericarp structure and phylogeny within Lamiaceae subfamily Nepetoideae tribe Ocimeae. Nordic Journal of Botany, 12(3), 273–298. doi:10.1111/j.1756-1051.1992.tb01304.x

Schuurink, R., & Tissier, A. (2019). Glandular trichomes: micro-organs with model status? The New Phytologist, nph.16283. doi:10.1111/nph.16283

Vargas, P., Fernández-Mazuecos, M., & Heleno, R. (2018). Phylogenetic evidence for a Miocene origin of Mediterranean lineages: species diversity, reproductive traits and geographical isolation. Plant Biology, 20, 157–165. doi:10.1111/plb.12626

Walker, J. B., Sytsma, K. J., Treutlein, J., & Wink, M. (2004). Salvia (Lamiaceae) is not monophyletic: implications for the systematics, radiation, and ecological specializations of Salvia and tribe Mentheae. American Journal of Botany, 91(7), 1115–1125. doi:10.3732/ajb.91.7.1115

Will, M., & Claßen-Bockhoff, R. (2017). Time to split Salvia s.l. (Lamiaceae) – New insights from Old World Salvia phylogeny. Molecular Phylogenetics and Evolution, 109, 33–58. doi:10.1016/j.ympev.2016.12.041

 

The Chestnut Song

“The Christmas Song” tops the charts every December, but there’s lots more to know about those chestnuts roasting on an open fire. We peel back the layers in this essay, which is one of our two contributions to this year’s Advent Botany holiday collection.

I first tried chestnuts when I was a student in Paris. The holiday season was peaceful that year, as it should be, and I’ve cherished my memories of it all the more as intense protests are spreading through France, and violence has shattered a Christmas market in Strasbourg. In that long-ago December, though, my most consuming emotion was a kind of double nostalgia. I missed home, and yet I wasn’t quite ready to leave that beautiful city behind. As I walked for hours and hours gathering last looks, it was thrilling to get caught up in the sudden early darkness of winter and the elaborate holiday windows of the grand old department stores. During one evening promenade, I saw a street merchant who had anchored himself in the middle of the streaming agitated crowd and was patiently tending a pan of marrons grillés, freshly roasted chestnuts. The scene was so sepia toned, so achingly 19th century, that I had to have some, just to glut my sentimentality. I bought a newspaper cone of the hot aromatic nuts and managed to peel one with my cold fingers right there on the sidewalk. Continue reading

Kiwifruit 2: Why are they green?

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

Kiwifruit 1: Why are they so fuzzy?

Kiwifruit is not covered in hairs. It’s covered in trichomes. And only if you’re talking about green Actinidia chinensis var. deliciosa. But, why? One answer is: pretty much to keep it from drying out. Another is: because it’s a polyploid from western China and was kind of chosen at random to be the most commonly grown kiwifruit, and they’re not all fuzzy. Those aren’t mutually exclusive answers. Put on your ecophysiology hats and grab a paring knife.

Think of fruit growth as a balancing act between ingoing and outgoing fluxes. When the balance is positive, fruits grow. When it is negative, they shrink—or shrivel. The main fluxes in question are carbon and water, which enter the fruit from the xylem and phloem of the plant vascular system. Water is lost mainly to the atmosphere via transpiration (evaporative water lost through stomata and other pores and from the skin surface). Keeping the ledger positive isn’t an easy job for a fruit. Hot, dry, and windy weather encourages transpiration and thereby increases the odds that a fruit will experience water stress. Excessive sunlight may cause sunburn. Fruits also need to avoid attack from pathogens and herbivores before the seeds within mature. A fruit’s skin—its cuticle and epidermis—is its first line of defense against abiotic and biotic threats. Some fruits resort to creative coverings to get the job done.

Here I’ll take a close look at the skin of kiwifruits. Why, exactly, are they so fuzzy?

A heart-shaped green kiwifruit (Actinidia chinensis var. deliciosa), covered in fuzzy trichomes

Continue reading

Pirates of the Carob Bean

Maybe the name takes you back to gentler days of Moosewood Cookbook and the dusty spicy local co-op. Or maybe you were a kid back then and fell for a chocolate bait-and-switch. Whether you are sweetly nostalgic or wary and resentful, it’s worth giving carob another chance. Katherine argues that it’s time to pull this earthy crunchy 70’s food into the superfood age. She offers foraging tips and recipes to help you get to know carob on its own terms.

From November through January, the carob trees in my neighborhood dangle hard, lumpy, dark brown fruits resembling lacquered cat turds. They are delicious and nutritious and of course I collect them. I am, without apology, a pod plundering, legume looting, pirate of the carob bean. CarobPiratesIf you seek adventure and happen to live in California, Arizona, or on the Mediterranean coast, you can probably pilfer some carob fruits yourself and play with them in your kitchen. If you lack local trees or the pirate spirit, you can order carob powder and even whole carob beans with one simple click.

Although plundering season begins just as the year is ending, I always wait until January to gather carob fruits for two reasons. First, carob functions mainly as a healthful chocolate substitute, and during the holiday season, fake chocolate just seems sad. In January, however, eating locally foraged carob feels virtuous and resourceful. Second, November and December are when my local carobs make the flowers that will produce the next year’s crop, and those flowers smell like a pirate’s nether parts after a shore leave. Or so I imagine, and not without precedent. A man who should have been inured to such salty smells, Pliny the Elder, natural historian and commander in the Roman Imperial Navy, described the flowers as having “a very powerful odor.” It’s not clear why these flowers have a sort of seaman scent, since the main volatiles wafting from the flowers – linalools and farnesene – smell like lilies and gardenias (Custódio et al., 2006). In any case, I keep my distance until the flowers have finished mating season.

Carob trees

Despite their stinky flowers, carobs make great street trees and produce a valuable crop in many Mediterranean-type climates. They are beautiful, tolerant of dry and poor soils, pest resistant, and tidy. Carobs are legumes – like familiar peas and beans – but they belong to a different branch of the legume family (Caesalpinioideae), one that contains mostly trees and woody shrubs with tough inedible fruit (Legume Phylogeny Working Group, 2017). Carob pods look about as edible as Jack Sparrow’s boots, and the species’ scientific name, Ceratonia siliqua, means “horny long pod,” which well captures the intimidating nature of their leathery fruit. But as you will see below, the fruits are easy to harvest and process, and their sweet pulp is worth seeking out. Continue reading

A holiday pineapple for the table

This deep dive into pineapple anatomy is our contribution this year to the very fun Advent Botany essay collection, a celebration of plants that are at least somewhat tangentially connected to the winter holidays. In previous years we’ve contributed essays on figs, peppermint, and sugar.

December is the time to bring out the fancy Christmas china, polish the silver pitchers, and . . . bedeck your best bromeliads. In 2017, as in 1700, no proper hostess can be without a pineapple for her centerpiece. Here we unpack the botany of pineapple, which is as complicated and fabulous as its cultural history. A proper hostess, after all, should also be able to dazzle her guests with tales of tropical fruit morphology. Continue reading

Botany Lab of the Month: Jack-O-Lantern

Happy National Pumpkin Day! Turn carving your Halloween Jack-O-Lantern into a plant dissection exercise.

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The first Jack-O-Lanterns were carved out of turnips in 17th-century Ireland. While the large, starchy hypocotyls (fused stem and taproot) of cruciferous vegetables are anatomically fascinating, this post will be about the stuff you are more likely cutting through to make a modern Jack-O-Lantern out of squash. Continue reading

Preserving diversity with some peach-mint jam

We are knee deep in peach season, and now is the time to gather the most diverse array of peaches you can find and unite them in jam. Katherine reports on some new discoveries about the genetics behind peach diversity and argues for minting up your peach jam.

Jam inspiration

Fresh peaches at their peak are fuzzy little miracles, glorious just as they are. But their buttery mouthfeel and dripping juice are lost when peaches are processed into jam and spread across rough toast. To compensate for textural changes, cooked peaches need a bit more adornment to heighten their flavor, even if it’s only a sprinkling of sugar. Normally I am not tempted to meddle with perfection by adding ginger or lavender or other flavors to peach jam. This year, however, as I plotted my jam strategy, the unusual juxtaposition of peach and mint found its way into my imagination over and over again, like the insistent echo of radio news playing in the background. Peach and mint, peach and mint, peach and mint – almost becoming a single word. To quiet the voice in my head I had to make some peach-mint jam. The odd combination turned out to be wonderful, and I’m now ready to submit the recipe to a candid world. As we will see below, it’s not without precedent. Mmmmmmpeachmint jam. Continue reading