Summer of Weeds: Plantains

This is a revised version of two ethnobotany posts that appeared previously on Awkward Botany: White Man’s Foot, part one and part two. Plantains have a long history of ethnobotonical uses, as well as a bad reputation of being pesky, hard-to-eliminate weeds. The two most common introduced plantain species in North America are broadleaf plantain (Plantago major) and lanceleaf plantain (Plantago lanceolata). Wherever our daily travels take us, chances are there is a plantain growing nearby.

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Plantago major is in the plantain family (Plantaginaceae) a family that consists of at least 90 genera, including common ornamental plants like Veronica (speedwells), Digitalis (foxgloves), and Antirrhinum (snapdragons). The genus Plantago, commonly known as plantains, consists of around 200 species distributed throughout the world in diverse habitats. Most of them are herbaceous perennials with similar growth habits.

Originating in Eurasia, P. major now has a cosmopolitan distribution. It has joined humans as they have traveled and migrated from continent to continent and is now considered naturalized throughout most temperate and some tropical regions. P. major has a plethora of common names – common plantain being the one that the USDA prefers. Other names include broadleaf plantain, greater plantain, thickleaf plantain, ribgrass, ribwort, ripplegrass, and waybread. Depending on the source, there are various versions of the name white man’s foot. Along the same line, a common name for P. major in South Africa is cart-track plant.

common plantain (Plantago major)

Common plantain starts by forming a rosette of broad leaves usually oriented flat against the ground. The leaves are egg-shaped with parallel veins; occasionally, leaf margins are faintly toothed. The inflorescence is a leafless spike up to 20 centimeters tall or taller with several tiny flowers that are a dull yellow-green-brown color. The flowers are wind pollinated and highly prone to self-pollination. The fruits are capsules that can contain as many as 30 seeds; an entire plant can produce as many as 15,000 seeds. The seeds are small, brown, sticky, and easily transported by wind or by adhering to shoes, clothing, animals, and machinery. They require light to germinate and can remain viable for up to 60 years.

Common plantain prefers sunny sites but can also thrive in part shade. It adapts to a variety of soil types but performs best in moist, clay-loam soils. It is often found in compacted soils and is very tolerant of trampling. This trait, along with its low-growing leaves that easily evade mower blades, explains why it is so common in turf grass. It is highly adaptable to a variety of habitats and is particularly common on recently disturbed sites (both natural and human caused). It is an abundant urban and agricultural weed.

Illustration of three Plantago species from Selected Weeds of the United States (Agriculture Handbook No. 366) circa 1970

Even though it is wind pollinated, its flowers are visited by syrphid flies and various bee species which feed on its pollen. Several other insects feed on its foliage, along with a number of mammalian herbivores. Cardinals and other bird species feed on its seeds.

Humans also eat plantain leaves, which contain vitamins A, C, and K. Young, tender leaves can be eaten raw, while older leaves need to be cooked as they become tough and stringy with age. The medicinal properties of  P. major have been known and appreciated at least as far back as the Anglo-Saxons, who likely used a poultice made from the leaves externally to treat wounds, burns, sores, bites, stings, and other irritations. It has also been used to stop cuts from bleeding and to treat rattlesnake bites. Apart from external uses, the plant was used internally as a pain killer and to treat ulcers, diarrhea, and other gastrointestinal issues.

P. major has been shown to have antibacterial, anti-inflammatory, antioxidant, and other biological properties; several chemical compounds have been isolated from the plant and deemed responsible for these properties. It is for this reason that P. major and other Plantago species have been used to treat such a diverse number of ailments. The claims are extensive and worth exploring. You can start by visiting the following sites:

Excerpts about plantains from The Book of Field and Roadside by John Eastman

Concerning their cosmopolitan nature: “Although both plantains [P. major and P. lanceolata] are Eurasian natives, they have long been thoroughly naturalized global residents; the designation ‘alien’ applies to them in the same sense that all white and black Americans are alien residents.”

In which I learned a new term: “Both species are anthropophilic (associate with humans); they frequent roadsides, parking areas, driveways, and vacant lots, occurring almost everywhere in disturbed ground. Where one species grows, the other can often be found nearby.”

Illustration by Amelia Hansen from The Book of Field and Roadside by John Eastman

Medicinal and culinary uses according to Eastman: “Plantains have versatile curative as well as culinary properties; nobody need go hungry or untreated for sores where plantains grow. These plants contain an abundance of beta carotene, calcium, potassium, and ascorbic acid. Cure-all claims for common plantain’s beneficial medical uses include a leaf tea for coughs, diarrhea, dysentery, lung and stomach disorders, and the root tea as a mouthwash for toothache. … Their most frequent and demonstrably effective use as a modern herb remedy, however, is as a leaf poultice for insect bites and stings plus other skin irritations. The leaf’s antimicrobial properties reduce inflammation, and its astringent chemistry relieves itching, swelling, and soreness.”

Even the seeds are “therapeutic”: “The gelatinous mucilage surrounding seeds can be readily separated, has been used as a substitute for linseed oil. Its widest usage is in laxative products for providing bulk and soluble fiber called psyllium, mainly derived from the plantain species P. ovata and leafy-stemmed plantain (P. psyllium), both Mediterranean natives.”

An excerpt from Weeds: In Defense of Nature’s Most Unloved Plants by Richard Mabey

“Plantain, ‘the mother of worts,’ is present in almost all the early prescriptions of magical herbs, back as far as the earliest Celtic fire ceremonies. It isn’t clear why such a drab plant – a plain rosette of grey-green leaves topped by a flower spike like a rat’s-tail – should have had pre-eminent status. But its weediness, in the sense of its willingness to tolerate human company, may have had a lot to do with it. The Anglo-Saxon names ‘Waybroad’ or ‘Waybread’ simply mean ‘a broad-leaved herb which grows by the wayside.’ This is plantain’s defining habit and habitat. It thrives on roadways, field-paths, church steps. In the most literal sense it dogs human footsteps. Its tough, elastic leaves, growing flush with the ground, are resilient to treading. You can walk on them, scuff them, even drive over them, and they go on living. They seem to actively prosper from stamping, as more delicate plants around them are crushed. The principles of sympathetic magic, therefore, indicated that plantain would be effective for crushing and tearing injuries. (And so it is, to a certain extent. The leaves contain a high proportion of tannins, which help to close wounds and halt bleeding.)”

Seed Dispersal via Caching – The Story of Antelope Bitterbrush

Generally speaking, individual plants produce an enormous amount of seeds. This may seem like a huge waste of resources, but the reality is that while each seed has the potential to grow into an adult plant that will one day produce seeds of its own, relatively few may achieve this. Some seeds will be eaten before they get a chance to germinate. Others germinate and soon die from lack of water, disease, or herbivory. Those that make it past the seedling stage continue to face similar pressures. Reaching adulthood, then, is a remarkable achievement.

Antelope bitterbrush is a shrub that produces hundreds of seeds per individual. Each seed is about the size of an apple seed. Some seeds may be eaten right away. Others fall to the ground and are ignored. But a large number are collected by rodents and either stored in burrows (larder hoarding) or in shallow depressions in the soil (scatter hoarding). It is through caching that antelope bitterbrush seeds are best dispersed. When rodents fail to return to caches during the winter, the seeds are free to sprout in the spring. Some of the seedlings will dry out and others will be eaten, but a few will survive, making the effort to produce all those seeds worth it in the end.

Fruits forming on antelope bitterbrush (Purshia tridentata)

Antelope bitterbrush (Purshia tridentata) is in the rose family and is often simply referred to as bitterbrush. It occurs in grasslands, shrub steppes, and dry woodlands throughout large sections of western North America. It is a deciduous shrub that generally reaches between three and nine feet tall but can grow up to twelve feet. It has wedge-shaped leaves that are green on top, grayish on bottom, and three-lobed. Flowers are yellow, strongly fragrant, and similar in appearance to others in the rose family. Flowering occurs mid-spring to early summer. Fruits are achenes – single seeds surrounded by papery or leathery coverings. The covering must rot away or be removed by animals before the seed can germinate.

Bitterbrush is an important species for wildlife. It is browsed by mule deer, pronghorn antelope, bighorn sheep, and other ungulates, including livestock. It provides cover for birds, rodents, reptiles, and ungulates. Its seeds are collected by harvester ants and rodents, its foliage is consumed by tent caterpillars and other insects, and its flowers are visited by a suite of pollinators. For all that it offers to the animal kingdom, it also relies on it for pollination and seed dispersal. The flowers of bitterbrush are self-incompatible, and if it wasn’t for ants and rodents, the heavy seeds – left to rely on wind and gravity – would have trouble getting any further than just a few feet from the parent plant.

Antelope bitterbrush (Purshia tridentata) in full bloom – photo credit: wikimedia commons

In a study published in The American Naturalist (February 1993), Stephen Vander Wall reported that yellow pine chipmunks were the primary dispersal agents of bitterbrush seeds in his Sierra Nevada study area. The optimal depth for seedling establishment was between 10-30 millimeters. Seeds that are cached too near the surface risk being pushed out of the ground during freeze and thaw cycles where they can desiccate upon germination. Cached bitterbrush seeds benefit when there are several seeds per cache because, as Vander Wall notes, “clumps of seedlings are better able to push through the soil and can establish from greater depths than single seedlings.”

Another study by Vander Wall, published in Ecology (October 1994), reiterated the importance of seed caching by yellow pine chipmunks in the establishment of bitterbrush seedlings. Seed caches, which consisted of anywhere from two to over a hundred seeds, were located as far as 25 meters from the parent plant. Cached seeds are occasionally moved to another location, but Vander Wall found that even these secondary caches produce seedlings. Of course, not all of the seedlings that sprout grow to maturity. Vander Wall states, “attrition over the years gradually reduces the number of seedlings within clumps.” Yet, more than half of the mature shrubs he observed in his study consisted of two or more individuals, leading him to conclude that “they arose from rodent caches.”

A study published in the Journal of Range Management (January 1996) looked at the herbivory of bitterbrush seedlings by rodents. In the introduction the authors discuss how “rodents [may] not only benefit from antelope bitterbrush seed caches as a future seed source, but also benefit from the sprouting of their caches as they return to graze the cotyledons of germinating seeds.”  In this study, Ord’s kangaroo rats, deer mice, and Great Basin pocket mice were all observed consuming bitterbrush seedlings, preferring them even when millet was offered as an alternative. The two species of mice also dug up seedlings, possibly searching for ungerminated seeds. Despite seed dispersal via caching, an overabundance of rodents can result in few bitterbrush seedlings reaching maturity.

A cluster of antelope bitterbrush seedlings that has been browsed. “Succulent, young seedlings are thought to be important in the diets of rodents during early spring because of the nutrients and water they contain.” — Vander Wall (1994)

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Photos of antelope bitterbrush seedling clusters were taken at Idaho Botanical Garden, where numerous clusters are presently on display along the pathways of the native plant gardens and the adjoining natural areas. 

Beavers and Water Lilies – An Introduction to Zoochory

Beavers are classic examples of ecosystem engineers. It is difficult to think of an animal – apart from humans – whose day-to-day activities have more impact on the landscape than beavers. Their dam building activities create wetlands that are used by numerous other species, and their selective harvesting of preferred trees affects species composition in riparian areas. And that’s just the start. Their extensive evolutionary history and once widespread distribution has made them major players in the landscape for millions of years.

Today, the beaver family (Castoridae) consists of just two extant species: Castor fiber (native to Eurasia) and Castor canadensis (native to North America). Both species were hunted by humans to the brink of extinction but, thanks to conservation efforts, enjoy stable populations despite having been eliminated from much of their historical ranges. Before the arrival of Europeans, North American beavers are estimated to have been anywhere from 60 million to 400 million strong. Extensive trapping reduced the population to less than half a million. Today, 10 million or more make their homes in rivers, streams, and wetlands across the continent.

North American beaver (Castor canadensis) - photo credit: wikimedia commons

North American beaver (Castor canadensis) – photo credit: wikimedia commons

Beavers are herbivores, and they harvest trees and shrubs to build dams and lodges. Their interactions with plants are legion, and so what better way to introduce the concept of animal-mediated seed dispersal than beavers. Plants have several strategies for moving their seeds around. Wind and gravity are popular approaches, and water is commonly used by plants both aquatic and terrestrial. Partnering with animals, however, is by far the most compelling method. This strategy is called zoochory.

Zoochory has many facets. Two major distinctions are epizoochory and endozoochory. In epizoochory, seeds become attached in some form or fashion to the outside of an animal. The animal unwittingly picks up, transports, and deposits the seeds. The fruits of such seeds are equipped with hooks, spines, barbs, or stiff hairs that help facilitate attachment to an animal’s fur, feathers, or skin. A well known example of this is the genus Arctium. Commonly known as burdock, the fruits in this genus are called burs – essentially small, round balls covered in a series of hooks. Anyone who has walked through – or has had a pet walk through – a patch of burdocks with mature seed heads knows what a nuisance these plants can be. But their strategy is effective.

The burs of Arctium - photo credit: wikimedia commons

The burs of Arctium – photo credit: wikimedia commons

Endozoochory is less passive. Seeds that are dispersed this way are usually surrounded by fleshy, nutritious fruits desired by animals. The fruits are consumed, and the undigested seeds exit out the other end of the animal with a bit of fertilizer. Certain seeds require passage through an animal’s gut in order to germinate, relying on chemicals produced during the digestion process to help break dormancy. Other seeds contain mild laxatives in their seed coats, resulting in an unscathed passage through the animal and a quick deposit. Some plants have developed mutualistic relationships with specific groups of animals regarding seed dispersal by frugivory. When these animal species disappear, the plants are left without the means to disperse their seeds, which threatens their future survival.

Beavers rely on woody vegetation to get them through the winter, but in warmer months, when herbaceous aquatic vegetation is abundant, such plants become their preferred food source. Water lilies are one of their favorite foods, and through both consumption of the water lilies and construction of wetland habitats, beavers help support water lily populations. This is how John Eastman puts it in The Book of Swamp and Bog: “Beavers relish [water lilies], sometimes storing the rhizomes. Their damming activities create water lily habitat, and they widely disperse the plants by dropping rhizome fragments hither and yon.”

Fragrant water lily (Nympaea odorata) - photo credit: wikimedia commons

Fragrant water lily (Nymphaea odorata) – photo credit: wikimedia commons

The seeds of water lilies (plants in the family Nymphaceae) are generally dispersed by water. Most species (except those in the genera Nuphar and Barclaya) have a fleshy growth around their seeds called an aril that helps them float. Over time the aril becomes waterlogged and begins to disintegrate. At that point, the seed sinks to the bottom of the lake or pond where it germinates in the sediment. The seeds are also eaten by birds and aquatic animals, including beavers. The aril is digestible, but the seed is not.

In her book, Once They Were Hats, Frances Backhouse writes about the relationship between beavers and water lilies. She visits a lake where beavers had long been absent, but were later reintroduced. She noted changes in the vegetation due to beaver activity – water lilies being only one of many plant species impacted.

Every year in late summer, the beavers devoured the seed capsules [of water lilies], digested their soft outer rinds and excreted the ripe undamaged seeds into the lake. Meanwhile, as they dredged mud from the botom of the lake for their construction projects, they were unintentionally preparing the seed bed. Seeing the lilies reminded me that beavers also inadvertantly propagate willows and certain other woody plants. When beavers imbed uneaten sticks into dams or lodges or leave them lying on moist soil, the cuttings sometimes sprout roots and grow.

Other facets of zoochory include animals hoarding fruits and seeds to be eaten later and then not getting back to them, or seeds producing fleshy growths that ants love called elaiosomes, resulting in seed dispersal by ants. Animals and plants are constantly interacting in so many ways. Zoochory is just one way plants use animals and animals use plants, passively or otherwise. These relationships have a long history, and each one of them is worth exploring and celebrating.

What’s in a Packet of Wildflower Seeds? – An Introduction

Occasionally I receive packets of wildflower seeds from companies that are not in the business of growing plants. They are promotional items – encouraging people to plant flowers while simultaneously marketing their wares. Often the seed packet lacks a list of the seeds included in the mix, and so it remains unclear what “wildflowers” are actually in there. My guess is that most seed packets like this go unplanted, and those that do get planted, may go uncared for. After all, the company that supplied them isn’t all that concerned about what gets done with them anyway.

As it is, generic packets of wildflower seeds like this may not actually contain any wildflower seeds. The term wildflower generally refers to a flowering plant that grows in the wild and was not intentionally planted by humans. It is synonymous with native plant, but it can also refer to non-native plants that have become naturalized. By this definition, a packet of wildflower seeds should only include seeds of native or naturalized plants and should not include horticultural selections, hybrids, or cultivated varieties. Ideally, the seed mix would be specific to a particular region, as each region throughout the world has its own suite of native wildflowers.

With that in my mind, I was immediately curious about an unlabeled packet of wildflower seeds I recently received as a promotional item from a company that has nothing to do with plants. This is a company that ships items nationwide and around the world, which leads me to believe that hundreds of people received similar packets of seeds around the same time I did. The seed packet is not labeled for a particular region, so all of us likely received a similar mix of seeds. “Wildflowers” then, at least in this case, means a random assortment of flowering plants with questionable provenance and no sense of geographic location.

The seed packet in question.

The seed packet in question.

Curiousity is killing me; so I am determined to find out what is in this mysterious packet of seeds. Using a pair of magnifying glasses, I seperated the seeds into 26 groups. Each group, from as best as I can tell, should be a unique species (or at least from the same genus). The next step will be to grow the seeds out and see what they actually are. I have limited space and time, so this is going to take a while. Since “wildflower” is not an exact term, I have decided that in order to be considered a wildflower the plant will have to be native to North America. (I should probably say western North America or Intermountain West, since that is where I am located, but that’s pushing it.)

The amount of seeds that each of the 26 groups consists of varies greatly, from a single seed to 52 seeds. Some of the seeds may not be viable, and some of the seedlings are sure to perish along the way. Despite losses, it should be clear in the end what this packet of seeds mainly consists of and whether or not it is indeed a wildflower seed mix. If I were skilled at identifying species simply by observing their seeds, I might be able to avoid growing them out, but I am not confident enough to do that. However, one group of seeds is almost certainly calendula. Calendula is a genus native to parts of Asia, Europe, and North Africa that has been introduced to North America. So, we’re already off to a bad start.

seed packets_experiment

To be clear, I have no intention of disclosing or calling out the company that sent the seeds. This is all in good fun. No hard feelings. I’m satisfying my own curiosity, and perhaps yours, too. Until the next update (which could be a while), go run through a field of wildflowers. Enjoy yourself.

Ethnobotany: Cattails

“If you ever eat cattails, be sure to cook them well, otherwise the fibers are tough and they take more chewing to get the starchy food from them than they are worth. However, they taste like potatoes after you have been eating them for a couple weeks, and to my way of thinking are extremely good.”  – Sam Gribley in My Side of the Mountain by Jean Craighead George

franz

Illustration by Franz Anthony (www.franzanth.com)

Ask anyone to list plants commonly found in American wetlands, and you can guarantee that cattails will make the list nearly every time. Cattails are widespread throughout the Northern Hemisphere. They are so successful, that it is hard to picture a wetland without them. In her book, Braiding Sweetgrass, Robin Wall Kimmerer discusses this well known association:

Cattails grow in nearly all types of wetlands, wherever there is adequate sun, plentiful nutrients, and soggy ground. Midway between land and water, freshwater marshes are among the most highly productive ecosystems on earth, rivaling the tropical rainforest. People valued the supermarket of the swamp for the cattails, but also as a rich source of fish and game. Fish spawn in the shallows; frogs and salamanders abound. Waterfowl nest here in the safety of the dense sward, and migratory birds seek out cattail marshes for sanctuary on their journeys.

The two most abundant species of cattails in North America are Typha latifolia (common cattail) and Typha angustifolia (narrow leaf cattail). T. angustifolia may have been introduced from Europe. The two species also hybridize to form Typha x glauca. There are about 30 species in the genus Typha, and they share the family Typhaceae with just one other genus. The common names for cattail are nearly as abundant as the plant itself: candlewick, water sausage, corn dog plant, cossack asparagus, reedmace, nailrod, cumbungi, etc., etc.

Cattails have long, upright, blade-like leaves. As they approach the base of the plant, the leaves wrap around each other to form a tight bundle with no apparent stem. As Kimmerer puts it, this arrangement enables the plants to “withstand wind and wave action” because “the collective is strong.” Flowers appear on a tall stalk that reaches up towards the tops of the leaves. The inflorescence is composed of hundreds of separate male and female flowers. Male flowers are produced at the top of the stalk and female flowers are found directly below them. In the spring, the male flowers dump pollen down onto the female flowers, and wind carries excess pollen to nearby plants, producing what looks like yellow smoke.

After pollination, the male flowers fade away, leaving the female flowers to mature into a seed head. Just like the flowers, the seeds are small and held tightly together, maintaining the familiar sausage shape. Each seed has a tuft of “hair” attached to it to aid in wind dispersal. In The Book of Swamp and Bog, John Eastman writes about the abundant seeds (“an estimated average of 220,000 seeds per spike”) of cattail: “A quick experiment, one that Thoreau delighted to perform, demonstrates how tightly the dry seeds are packed in the spike – pull out a small tuft and watch it immediately expand to fill your hand with a downy mass.”

cattails bunch

cattail fluff

Because cattails spread so readily via rhizomes, prolific airborne seeds mostly serve to colonize new sites, away from the thick mass of already established cattails. The ability to dominate vast expanses of shoreline gives cattails an invasive quality that often results in attempts at removal. Various human activities may be aiding their success. Regardless, they provide food and habitat to numerous species of insects, spiders, birds, and mammals. A cattail marsh may not be diverse plant-wise, but it is teeming with all sorts of other life.

Ethnobotanically speaking, it is hard to find many other species that have as many human uses as cattails. For starters, nearly every part of the plant is edible at some point during the year. The rhizomes can be consumed year-round but are best from fall to early spring. They can be roasted, boiled, grated, ground, or dried and milled into flour. Starch collected from pounding and boiling the rhizomes can be used as a thickener. In the spring, young shoots emerging from the rhizomes and the tender core of the leaf bundles can be eaten raw or cooked and taste similar to cucumber. Young flower stalks can be boiled and eaten like corn on the cob and taste similar to artichoke. Pollen, which is high in protein, can be mixed with flour and used to make pancakes and baked goods, among other things. The seeds can be ground into flour or pressed to produce cooking oil.

Cattail leaves can be used to make cords, mats, baskets, thatch, and many other things. Kimmerer writes about the excellent wigwam walls and sleeping mats that weaved cattail leaves make:

The cattails have made a suburb material for shelter in leaves that are long, water-repellent, and packed with closed-cell foam for insulation. … In dry weather, the leaves shrink apart from one another and let the breeze waft between them for ventilation. When the rains come, they swell and close the gap, making the [wall] waterproof. Cattails also make fine sleeping mats. The wax keeps away moisture from the ground and the aerenchyma provide cushioning and insulation.

The fluffy seeds make great tinder for starting fires, as well as excellent insulation and pillow and mattress stuffing. The dry flower stalks can be dipped in fat, lit on fire, and used as a torch. Native Americans used crushed rhizomes as a poultice to treat burns, cuts, sores, etc. A clear gel is found between the tightly bound leaves of cattail. Kimmerer writes, “The cattails make the gel as a defense against microbes and to keep the leaf bases moist when water levels drop.” The gel can be used like aloe vera gel to soothe sunburned skin.

Eastman rattles off a number of commercial uses for cattail: “Flour and cornstarch from rhizomes, ethyl alcohol from the fermented flour, burlap and caulking from rhizome fibers, adhesive from the stems, insulation from the downy spikes, oil from the seeds, rayon from cattail pulp, …” To conclude his section on cattails he writes, “With cattails present, one need not starve, freeze, remain untreated for injury, or want for playthings.”

Additional Resources:

Year of Pollination: Botanical Terms for Pollination, part two

“The stage is set for reproduction when, by one means or another, compatible pollen comes to rest on a flower’s stigma. Of the two cells within a pollen grain, one is destined to grow into a long tube, a pollen tube, that penetrates the pistil’s tissues in search of a microscopic opening in one of the ovules, located in the ovary. … The second of a pollen grain’s cells divides to become two sperm that move through the pollen tube and enter the ovule.” – Brian Capon, Botany for Gardeners

“Once pollination occurs, the next step is fertilization. Pollen deposited on the sticky stigma generates a fine pollen tube that conveys the sperm through the style to the ovary, where the ovules, or eggs, have developed. After fertilization, the rest of the flower parts wither and are shed as the ovary swells with seed development.” – Rick Imes, The Practical Botanist

Pollination tells the story of a pollen grain leaving an anther by some means – be it wind, water, or animal – and finding itself deposited atop a stigma. As long as the pollen and stigma are compatible, the sex act proceeds. In other words, the pollen grain germinates. One of the pollen grain’s cells – the tube nucleus – grows down the length of the style, forming a tube through which two sperm nuclei can travel. The sperm nuclei enter the ovary and then, by way of a micropyle, enter an ovule. Inside the ovule is the female gametophyte (also referred to as the embryo sac). One sperm nucleus unites with the egg nucleus to form a zygote. The remaining sperm nucleus unites with two polar nuclei to form a triploid cell which becomes the endosperm. The sex act is complete.

The illustration on the left includes the cross-section of a pistil showing the inside the ovary where pollen tubes have made their way to the ovules. The illustration on the right shows pollen grains germinating on a stigma and their pollen tubes begining to work their way down the style. (photo credit: wikimedia commons)

The illustration on the left includes the cross section of a pistil showing the inside of the ovary where pollen tubes have made their way to the ovules. The illustration on the right shows pollen grains germinating on a stigma and pollen tubes as they work their way down the style. (image credit: wikimedia commons)

The zygote divides by mitosis to become an embryo. The endosperm nourishes the development of the embryo. The ovule matures into a seed, and the ovary develops into a fruit. During this process, the remaining parts of the flower wither and fall away. In some cases, certain flower parts remain attached to the fruit or become part of the fruit. The flesh of an apple, for example, is formed from the carpels and the receptacle (the thickened end of a flower stem – peduncle – to which the parts of a flower are attached).

As the seed matures, the endosperm is either used up or persists to help nourish the embryonic plant after germination. Mature seeds that are abundant in endosperm are called albuminous. Examples include wheat, corn, and other grasses and grains. Mature seeds with endosperm that is either highly reduced or absent are called exalbuminous – beans and peas, for example. Certain species – like orchids – do not produce endosperm at all.

The cross section of a corn kernel showing the endosperm and the embryo (image credit: Encyclopedia Britannica Kids)

The cross section of a corn kernel showing the endosperm and the embryo (image credit: Encyclopedia Britannica Kids)

It is fascinating to consider that virtually every seed we encounter is the result of a single pollen grain making its way from an anther to a stigma, growing a narrow tube down a style, and fertilizing a single ovule. [Of course there are always exceptions. Some plants can produce seeds asexually. See apomixis.] Think of this the next time you are eating corn on the cob or popcorn – each kernel is a single seed – or slicing open a pomegranate to reveal the hundreds of juicy seeds inside. Or better yet, when you are eating the flesh or drinking the milk of a coconut. You are enjoying the solid and liquid endosperm of one very large seed.

Much more can be said about pollination and the events surrounding it, but we’ll save that for future posts. The “Year of Pollination” may be coming to an end, but there remains much to discover and report concerning the subject. For now, here is a fun video to help us review what we’ve learned so far:

 

Also, take a look at this TED talk: The Hidden Beauty of Pollination by Louie Schwartzberg

And finally, just as the “Year of Pollination” was coming to an end I was introduced to a superb blog called The Amateur Anthecologist. Not only did it teach me that “anthecology” is a term synonymous with pollination biology, it has a great series of posts called “A Year of Pollinators” that showcases photographs and information that the author has collected for various groups of pollinators over the past year. The series includes posts about Bees, Wasps, Moths and ButterfliesFlies, and Beetles, Bugs, and Spiders.

Harvester Ants – Seed Predators and Seed Dispersers

“The abundance of ants is legendary. A worker is less than one-millionth the size of a human being, yet ants taken collectively rival people as dominant organisms on the land. …  When combined, all ants in the world taken together weigh about as much as all human beings.” – Journey to the Ants by Bert Hölldobler and Edward O. Wilson

Considering how abundant and widely distributed ants are, it is easy to imagine the profound role they might play in the ecosystems of which they are a part. In fact, in the epilogue to Hölldobler and Wilson’s popular book about ants (quoted above), they conclude that in a world without ants, “species extinction would increase even more over the present rate, and the land ecosystems would shrivel more rapidly as the considerable services provided by these insects were pulled away.” It is no doubt then that ants, through their myriad interactions with their surroundings, are key players in terrestrial ecosystems.

photo credit: www.eol.org

photo credit: www.eol.org

Harvester ants offer a prime example of the important roles that ants can play. In the process of collecting seeds for consumption, harvester ants can help shape the abundance and distribution of the plants in their immediate environment. They do this by selecting the types and amounts of seeds they collect, by abandoning seeds along their collection routes, and by leaving viable seeds to germinate in and around their nests. Hölldobler and Wilson have this to say about harvester ants:

[The] numerical success [of ants] has allowed them to alter not just their nest environments, but the entire habitats in which they live. Harvesting ants, species that regularly include seeds in their diet, have an especially high impact. They consume a large percentage of the seeds produced by plants of many kinds in nearly all terrestrial habitats, from dense tropical forests to deserts. Their influence is not wholly negative. The mistakes they make by losing seeds along the way also disperse plants and compensate at least in part for the damage caused by their predation.

There are more than 150 species of harvester ants, spanning at least 18 genera. They are found throughout the world (except extreme cold locales) and are particularly common in arid to semi-arid environments. Pogonomyrmex is one the largest genera of harvester ants with nearly 70 species occurring throughout North, Central, and South America. Messor is another large genus of harvester ants that mainly occurs in Europe, Asia, and Africa. Both of these genera build large nests and move massive amounts of soil in the process.

Seed dispersal by harvester ants (also known as diszoochory) is a type of secondary (or Phase II) seed dispersal. It is a case of serendipity, as the dispersal occurs largely by accident. Some plants, on the other hand, have developed a mutualistic relationship with ants, enlisting them to disperse their seeds by way of an elaiosome – a fleshy, nutritious structure attached to seeds that attracts ants. Seeds with such structures are picked up by ants and brought to their nests where the elaiosome is consumed and the seed is left to germinate. This form of ant-mediated dispersal is called myrmecochory and is typically not carried out by harvester ants.

photo credit: wikimedia commons

photo credit: wikimedia commons

Harvester ant colonies have both direct and indirect influences on their surrounding environments; however, there is a dearth of research elucidating the exact details of such influences. A paper published in the Annual Review of Ecology and Systematics in 2000 by MacMahon et. al. reviewed available studies concerning harvester ants and explored our current understanding of the influences that harvester ants (particularly those in the genus Pogonomyrmex) can potentially have on community structure and ecosystem functions. Following are some of the direct influences the authors listed:

  • Removal and consumption of seeds and other materials – The relative abundance of plant species can be affected by the selective removal of seeds. Harvester ants also collect leaves, twigs, pollen, flowers, vertebrate feces, and arthropod body parts.
  • Storage and rejection of seeds – Collected seeds can be dropped during transport, rejected after arriving at the nest, or abandoned in nest granaries. All result in the transport of seeds away from the parent plant and dispersal beyond the plants’ primary dispersal mechanisms.
  • Construction and maintenance of nests – All vegetation and debris is removed from the area immediately surrounding the nest including mature and emerging plants. This area is kept clear for the duration of the life of the colony and, in some cases, can be quite extensive.

Harvester ants can also influence soil properties and soil food webs within and in the vicinity of their nests. They bring large amounts of organic matter down into the soil and redistribute vast amounts of soil particles. Their actions also influence the amount of moisture in the soil surrounding their nests.

This is a mere distillation of the influences that harvester ants might have; see the paper by MacMahon et al. to learn more.

In an effort to better understand how the seed predation and seed dispersal behaviors of harvester ants might influence plant population dynamics, a research team in Spain used data obtained from field research to build a computer model that would predict changes over time. The study site was described as “open and heterogeneous shrubland” and the vegetation was stated to be in “a very early stage in the secondary succession” after being subject to “recurring fires.” The harvester ant colonies involved in the study consisted of three species in the genus Messor. The plant species selected for the study were three native shrubs whose seeds were known to be collected by the harvester ants. Each plant species differed slightly in the amount and size of seeds it produced and in its primary seed dispersal mechanism, which is important because the researchers hypothesized that “the effect of seed predation and seed dispersal may depend on plant attributes.”

Messor bouvieri (photo credit: www.eol.org)

Messor bouvieri (photo credit: www.eol.org)

Data obtained from simulated scenarios and field observations appeared to support this hypothesis; each shrub species interacted differently with the harvester ants. Coronilla minima benefited from “accidental” seed dispersal. Comparatively, it produces a high amount of large seeds, which are primarily dispersed by gravity. Despite predation, ant-mediated dispersal was an advantage. Dorycnium pentaphyllum produced the highest amount of seeds among the three shrub species; however, seed predation was found to have negative effects on its population dynamics. Its primary seed dispersal mechanism involves ballistics (the mechanical ejection of its seeds), so ant-mediated dispersal may not offer an advantage. Finally, Fumana ericoides, despite its limited primary seed dispersal and its comparatively low production of seeds was not affected by the actions of the harvester ants. The authors concluded that “some unknown factor is driving the population dynamics of this species, more than the action of ants.”

Studies such as this, while leaving many unanswered questions, help us understand the important role that harvester ants play in our world. Harvester ants, and ants in general, are truly among Earth’s most enthralling and influential creatures. Learn more about their complex behaviors and countless interactions with flora and fauna by checking out these three documentaries recommended by ANTfinity.