A Few Fun Facts About Pollen

Sexual reproduction in vascular plants requires producing and transporting pollen grains – the male gametophytes or sperm cells of a plant. These reproductive cells must make their way to the egg cells in or order to form seeds – plants in embryo. The movement of pollen is something we can all observe. It’s happening all around us on a regular basis. Any time a seed-bearing plant (also known as a spermatophyte) develops mature cones or flowers, pollen is on the move. Pollen is a ubiquitous and enduring substance and a fascinating subject of study. In case you don’t believe me, here are a few fun facts.

Bee covered in pollen – photo credit: wikimedia commons

Pollen is as diverse as the species that produce it. Pollen grains are measured in micrometers and are so tiny that the only reason we can see them with the naked eye is because they are often found en masse. Yet they are incredibly diverse in size, shape, and texture, and each plant species produces its own unique looking pollen. With the help of a good microscope, plants can even be identified simply by looking at their pollen. See images of the pollen grains of dozens of plant species here and here.

Pollen helps us answer questions about the past. Because pollen grains are so characteristic and because their outer coating (known as exine) is so durable and long-lasting, studying pollen found in sediments and sedimentary rocks helps us discover all sorts of things about deep time. The study of pollen and other particulates is called palynology. Numerous disciplines look to palynology to help them answer questions and solve mysteries. Its even used in forensics to help solve crimes. Criminals should be aware that brushing up against a plant in bloom may provide damning evidence.

Pollen oddities. While all pollen is different, some plants produce particularly unique pollen. The pollen grains of plants in the orchid and milkweed families, for example, are formed into united masses called pollinia. Each pollinium is picked up by pollinators and transferred to the stigmas of flowers as a single unit. A number of other species produce other types of compound pollen grains. The pollen grains of pines and other conifers are winged, and the pollen grains of seagrass species, like Zostera spp., are filamentous and said to hold the record for longest pollen grains.

The pollinia of milkweed (Asclepias spp.) look like the helicopter-esque fruits of maple trees. photo credit: wikimedia commons

Pollen tube oddities. In flowering plants, when pollen grains reach the stigma of a compatible flower, a vegetative cell within the grain forms a tube in order to transport the regenerative cells into the ovule. This tube varies in length depending on the length of the flower’s style. Because corn flowers produce such long styles (also know as corn silk), corn pollen grains hold the record for longest pollen tube, which can measure 12 inches or more. Species found in the mallow, gourd, and bellflower families produce multiple pollen tubes per pollen grain. Hence, their pollen is said to be polysiphonous.

Pollen is transported in myriad ways. Plants have diverse ways of getting their pollen grains where they need to be. Anemophilous plants rely on wind and gravity. They produce large quantities of light-weight pollen grains that are easily dislodged. Most of this pollen won’t make it, but enough of it will to make this strategy worth it. Hydrophilous plants use water and, like wind pollinated plants, may produce lots of pollen due to the unpredictably of this method. Some hydrophilous plants transport their pollen on the surface of the water, while others are completely submerged during pollination.

Employing animals to move pollen is a familiar strategy. Entomophily (insect pollination) is the most common, but there is also ornithophily (bird pollination) and chiropterophily (bat pollination), among others. Plants that rely on animals for pollination generally produce pollen grains that are sticky and nutritious. They attract animals using showy flowers, fragrance, and nectar. The bodies of pollinating insects have modifications that allow them to collect and transport pollen. Certain bees, like honey bees and bumblebees, have pollen baskets on their hind legs, while other bees have modified hairs called scopae on certain parts of their bodies.

Pollen is edible. Some animals – both pollinating and non-pollinating – use pollen as a food source. Animals that eat pollen are palynivores. Bees, of course, eat pollen, but lots of other insects do, too. Even some spiders, which are generally thought of as carnivores, have been observed eating pollen that gets trapped in their webs.

Pollen is thought to be highly nutritious for humans as well, and so, along with being taken as a supplement, it is used in all sorts of food products. To collect pollen, beekeepers install pollen traps on their beehives that strip incoming worker bees of their booty. Pollen from various wind pollinated plants, like cattails and pine trees, are also collected for human consumption. For example, a Korean dessert called dasik is made using pine pollen.

pine pollen – photo credit: wikimedia commons

Pollen makes many people sick. Hay fever is a pretty common condition and is caused by an allergy to wind-borne pollen. This condition is also known as pollinosis or allergic rhinitis. Not all flowering plants are to blame though, so here is a list of some of the main culprits. Because so many people suffer from hay fever, pollen counts are often included in weather reports. Learn more about what those counts mean here.

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Bats As Pollinators – An Introduction to Chiropterophily

Most plants that rely on animals to assist in pollination look to insects. In general, insects are abundant, easy to please, and efficient at transferring pollen. Because insect pollination is such a common scenario, it is easy to overlook pollination that is carried out by vertebrates. Birds are the most prominent pollinator among vertebrates, but mammals participate, too. The most common mammal pollinator is the bat.

About a fifth of all mammal species on the planet are bats, with species estimates numbering in the 1200-1300 range. Bats are the only mammals that can truly fly. They are not blind, nor are they flying rodents, and they are not going to suck your blood (except in very rare cases!). Most bats eat insects, but a small, significant group of them are nectarivorous. Their main food source is the nectar produced within flowers. In the process of feeding, these bats pollinate plants.

Out of 18 families in the order Chiroptera, only two include species with morphologies that set them apart as nectar-feeders. The family Pteropodidae, known commonly as Old World fruit bats or flying foxes, occurs in tropical and subtropical regions of Africa, Asia, Australia, Papa New Guinea, and the Pacific Islands. The family Phyllostomidae, known commonly as American leaf-nosed bats, occurs in tropical and subtropical regions of the Americas. For simplicity’s sake, the former are referred to as Old World bats, and the latter as New World bats. While both groups are similar in that they consist of species that feed on nectar, they are only distantly related, and thus the nectar feeding species in these families have distinct behavioral and morphological differences.

Grey headed flying fox photo credit: wikimedia commons

Grey headed flying fox (Pteropus poliocephalus), a floral visiting bat from Australia (photo credit: wikimedia commons)

More than 500 species of plants, spanning 67 plant families, are pollinated by bats. This pollination syndrome is known as chiropterophily. In general, flowers that use this approach tend to be white or dull in color, open at night, rich with nectar, and musty or rotten smelling. They are generally tubular, cup shaped, or otherwise radially symmetrical and are often suspended atop tall stalks or prominently located on branches or trunks. In a review published in Annals of Botany, Theodore Fleming, et al. state “flower placement away from foliage and nocturnal anthesis [blooming] are the unifying features of the bat pollination syndrome,” while all other characteristics are highly variable among species. The family Fabaceae contains the highest number of bat-pollinated genera. Cactaceae, Malvaceae, and Bignoniaceae follow closely behind.

The characteristics of bat pollinated flowers vary widely partly because the bats that visit them are so diverse. Between the two bat families there are similarities in their nectar-feeding species, including an elongated rostrum, teeth that are smaller in number and size, and a long tongue with hair-like projections on the tip. Apart from that, New World bats are much smaller than Old World bats, and their rostrums and tongues are much longer relative to the size of their bodies. New World bats have the ability to hover in front of flowers, while Old World bats land on flowers to feed. Old World bats do not have the ability to use echolocation to spot flowers, while New World bats do. Fleming, et al. conclude, “because of these differences, we might expect plants visited by specialized nectar-feeding [New World bats] to produce smaller flowers with smaller nectar volumes per flower than those visited by their [Old World bat] counterparts.”

Pollination by bats is a relatively new phenomenon, evolutionarily speaking. Flowers that are currently pollinated by bats most likely evolved from flowers that were once pollinated by insects. Some may have evolved from flowers that were previously bird pollinated. The question is, why adopt this strategy? Flowers that are bat pollinated are “expensive” to make. They are typically much bigger than insect pollinated flowers, and they contain large amounts of pollen and abundant, nutrient-rich nectar. Due to resource constraints, many plants are restricted from developing such flowers, but those that do may find themselves at an advantage with bats as their pollinator. For one, hairy bat bodies collect profuse numbers of pollen grains, which are widely distributed as they visit numerous flowers throughout the night. In this way, bats can be excellent outcrossers. They also travel long distances, which means they can move pollen from one population of plants to an otherwise isolated neighboring population. This serves to maintain healthy genetic diversity among populations, something that is increasingly important as plant populations become fragmented due to human activity.

Pollinating bats are also economically important to humans, as several plants that are harvested for their fruits, fibers, or timber rely on bats for pollination. For example, bat pollinated Eucalyptus species are felled for timber in Australia, and the fruits of Durio zibethinus in Southeast Asia form after flowers are first pollinated by bats. Also, the wild relatives of bananas (Musa spp.) are bat pollinated, as is the plant used for making tequila (Agave tequilana).

Durio sp. (photo credit: wikimedia commons)

The flowers of durian (Durio sp.), trees native to Southeast Asia, are pollinated by bats (photo credit: wikimedia commons)

There is still much to learn about nectarivorous bats and the flowers they visit. It is clear that hundreds of species are using bats to move their pollen, but the process of adopting this strategy and the advantages of doing so remain ripe for discovery. Each bat-plant relationship has its own story to tell. For now, here is a fun video about the bat that pollinates Agave tequilana:

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


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.

Year of Pollination: Botanical Terms for Pollination, part one

When I began this series of posts, I didn’t have a clear vision of what it would be. I had a budding interest in pollination biology and was anxious to learn all that I could. I figured that calling 2015 the “Year of Pollination” and writing a bunch of pollination-themed posts would help me do that. And it has. However, now that the year is coming to a close, I realize that I neglected to start at the beginning. Typical me.

What is pollination? Why does it matter? The answers to these questions seemed pretty obvious; so obvious, in fact, that I didn’t even think to ask them. That being said, for these last two “Year of Pollination” posts (and the final posts of the year), I am going back to the basics by defining pollination and exploring some of the terms associated with it. One thing is certain, there is still much to be discovered in the field of pollination biology. Making those discoveries starts with a solid understanding of the basics.

Pollination simply defined is the transfer of pollen from an anther to a stigma or – in gymnosperms – from a male cone to a female cone. Essentially, it is one aspect of plant sex, albeit a very important one. Sexual reproduction is one way that plants multiply. Many plants can also reproduce asexually. Asexual reproduction typically requires less energy and resources – no need for flowers, pollen, nectar, seeds, fruit, etc. – and can be accomplished by a single individual without any outside help; however, there is no gene mixing (asexually reproduced offspring are clones) and dispersal is limited (consider the “runners” on a strawberry plant producing plantlets adjacent to the mother plant).

To simplify things, we will consider only pollination that occurs among angiosperms (flowering plants); pollination/plant sex in gymnosperms will be discussed at another time. Despite angiosperms being the youngest group of plants evolutionarily speaking, it is the largest group and thus the type we encounter most.

A flower is a modified shoot and the reproductive structure of a flowering plant. Flowers are made up of a number of parts, the two most important being the reproductive organs. The androecium is a collective term for the stamens (what we consider the male sex organs). A stamen is composed of a filament (or stalk) topped with an anther – where pollen (plant sperm) is produced. The gynoecium is the collective term for the pistil (what we consider the female sex organ). This organ is also referred to as a carpel or carpels; this quick guide helps sort that out. A pistil consists of the ovary (which contains the ovules), and a style (or stalk) topped with a stigma – where pollen is deposited. In some cases, flowers have both male and female reproductive organs. In other cases, they have one or the other.

photo credit: wikimedia commons

photo credit: wikimedia commons

When pollen is moved from an anther of one plant to a stigma of another plant, cross-pollination has occurred. When pollen is moved from an anther of one plant to a stigma of the same plant, self-pollination has occurred. Cross-pollination allows for gene transfer, and thus novel genotypes. Self-pollination is akin to asexual production in that offspring are practically identical to the parent. However, where pollinators are limited or where plant populations are small and there is little chance for cross-pollination, self-pollination enables reproduction.

Many species of plants are unable to self-pollinate. In fact, plants have evolved strategies to ensure cross-pollination. In some cases, the stamens and pistils mature at different times so that when pollen is released the stigmas are not ready to receive it or, conversely, the stigmas are receptive before the pollen has been released. In other cases, stigmas are able to recognize their own pollen and will reject it or inhibit it from germinating. Other strategies include producing flowers with stamens and pistils that differ dramatically in size so as to discourage pollen transfer, producing separate male and female flowers on the same plant (monoecy), and producing separate male and female flowers on different plants (dioecy).

As stated earlier, the essence of pollination is getting the pollen from the anthers to the stigmas. Reproduction is an expensive process, so ensuring that this sex act takes place is vital. This is the reason why flowers are often showy, colorful, and fragrant. However, many plants rely on the wind to aid them in pollination (anemophily), and so their flowers are small, inconspicuous, and lack certain parts. They produce massive amounts of tiny, light-weight pollen grains, many of which never reach their intended destination. Grasses, rushes, sedges, and reeds are pollinated this way, as well as many trees (elms, oaks, birches, etc.) Some aquatic plants transport their pollen from anther to stigma via water (hydrophily), and their flowers are also simple, diminutive, and produce loads of pollen.

Inforescence of big bluestem (Andropogon gerardii), a wind pollinated plant - pohto credit: wikimedia commons

Inflorescence of big bluestem (Andropogon gerardii), a wind pollinated plant – photo credit: wikimedia commons

Plants that employ animals as pollinators tend to have flowers that we find the most attractive and interesting. They come in all shapes, sizes, and colors and are anywhere from odorless to highly fragrant. Odors vary from sweet to bitter to foul. Many flowers offer nectar as a reward for a pollinator’s service. The nectar is produced in special glands called nectaries deep within the flowers, inviting pollinators to enter the flower where they can be dusted with pollen. The reward is often advertised using nectar guides – patterns of darker colors inside the corolla that direct pollinators towards the nectar. Some of these nectar guides are composed of pigments that reflect the sun’s ultraviolet light – they are invisible to humans but are a sight to behold for many insects.

In part two, we will learn what happens once the pollen has reached the stigma – post-pollination, in other words. But first, a little more about pollen. The term pollen actually refers to a collection of pollen grains. Here is how Michael Allaby defines “pollen grain” in his book The Dictionary of Science for Gardeners: “In seed plants, a structure produced in a microsporangium that contains one tube nucleus and two sperm nuclei, all of them haploid, enclosed by an inner wall rich in cellulose and a very tough outer wall made mainly from sporopollenin. A pollen grain is a gametophyte.”

A pollen grain’s tough outer wall is called exine, and this is what Allaby has to say about that: “It resists decay, and the overall shape of the grain and its surface markings are characteristic for a plant family, sometimes for a genus or even a species. Study of pollen grains preserved in sedimentary deposits, called palynology or pollen analysis, makes it possible to reconstruct past plant communities and, therefore, environments.”

Scanning electron microscope image of pollen grains from narrowleaf evening primrose (Oenothera fruticosa) - photo credit: wikimedia commons

Scanning electron microscope image of pollen grains from narrowleaf evening primrose (Oenothera fruticosa) – photo credit: wikimedia commons