Gymnosperms and Fleshy "Fruits"

Fleshy red aril surrounding the seeds of  Taxus baccata.

Fleshy red aril surrounding the seeds of Taxus baccata.

Many of us were taught in school that one of the key distinguishing features between gymnosperms and angiosperms is the production of fruit. Fruit, by definition, is a structure formed from the ovary of a flowering plant. Gymnosperms, on the other hand, do not enclose their ovules in ovaries. Instead, their unfertilized ovules are exposed (to one degree or another) to the environment. The word “gymnosperm” reflects this as it is Greek for “naked seed.” However, as is the case with all things biological, there are exceptions to nearly every rule. There are gymnosperms on this planet that produce structures that function quite similar to fruits.

Cross section of a  Ginkgo  ovule with red arrow showing the integument.  Photo copyright Bruce Kirchoff, Licensed under CC-BY

Cross section of a Ginkgo ovule with red arrow showing the integument.

Photo copyright Bruce Kirchoff, Licensed under CC-BY

The key to understanding this evolutionary convergence lies in understanding the benefits of fruits in the first place. Fruits are all about packing seeds into structures that appeal to the palates of various types of animals who then eat said fruits. Once consumed, the animals digest the fruity bits and will often deposit the seeds elsewhere in their feces. Propagule dispersal is key to the success of plants as it allows them to not only to complete their reproductive cycle but also conquer new territory in the process. With a basic introduction out of the way, let’s get back to gymnosperms.

“Fruits” of  Cephalotaxus fortunei  (Cephalotaxaceae)

“Fruits” of Cephalotaxus fortunei (Cephalotaxaceae)

There are 4 major gymnosperm lineages on this planet - the Ginkgo, cycads, gnetophytes, and conifers. Each one of these groups contains members that produce fleshy structures around their seeds. However, their “fruits” do not all develop in the same way. The most remarkable thing to me is that, from a developmental standpoint, each lineage has evolved its own pathway for “fruit” production.

Ginkgo  “fruits” are full of butyric acid and smell like rotting butter or vomit.

Ginkgo “fruits” are full of butyric acid and smell like rotting butter or vomit.

For instance, consider ginkgos and cycads. Both of these groups can trace their evolutionary history back to the early Permian, some 270 - 280 million years ago, long before flowering plants came onto the scene. Both surround their developing seed with a layer of protective tissue called the integument. As the seed develops, the integument swells and becomes quite fleshy. In the case of Ginkgo, the integument is rich in a compound called butyric acid, which give them their characteristic rotten butter smell. No one can say for sure who this nasty odor originally evolved to attract but it likely has something to do with seed dispersal. Modern day carnivores seem to be especially fond of Ginkgo “fruits,” which would suggest that some bygone carnivore may have been the main seed disperser for these trees.

“Fruits” contained within the female cone of a cycad ( Lepidozamia peroffskyana ).

“Fruits” contained within the female cone of a cycad (Lepidozamia peroffskyana).

The Gnetophytes are represented by three extant lineages (Gnetaceae, Welwitschiaceae, and Ephedraceae), but only two of them - Gnetaceae and Ephedraceae - produce fruit-like structures. As if the overall appearance of the various Gnetum species didn’t make you question your assumptions of what a gymnosperm should look like, its seeds certainly will. They are downright berry-like!

Berry-like seeds of  Gnetum gnemon .

Berry-like seeds of Gnetum gnemon.

The formation of the fruit-like structure surrounding each seed can be traced back to tiny bracts at the base of the ovule. After fertilization, these bracts grow up and around the seed and swell to become red and fleshy. As you can imagine, Gnetum “fruits” are a real hit with animals. In the case of some Ephedra, the “fruit” is also derived from much larger bracts that surround the ovule. These bracts are more leaf-like at the start than those of their Gnetum cousins but their development and function is much the same.

Red, fleshy bracts of  Ephedra distachya .

Red, fleshy bracts of Ephedra distachya.

Whereas we usually think of woody cones when we think of conifers, there are many species within this lineage that also have converged on fleshy structures surrounding their seeds. Probably the most famous and widely recognized example of this can be seen in the yews (Taxus spp.). Ovules are presented singly and each is subtended by a small stalk called a peduncle. Once fertilized, a group of cells on the peduncle begin to grow and differentiate. They gradually swell and engulf the seed, forming a bright red, fleshy structure called an “aril.” Arils are magnificent seed dispersal devices as birds absolutely relish them. The seed within is quite toxic so it usually escapes the process unharmed and with any luck is deposited far away from the parent plant.

The berry-like cones of  Juniperus communis .

The berry-like cones of Juniperus communis.

Another great example of fleshy conifer “fruits” can be seen in the junipers (Juniperus spp.). Unlike the other gymnosperms mentioned here, the junipers do produce cones. However, unlike pine cones, the scales of juniper cones do not open to release the seeds inside. Instead, they swell shut and each scale becomes quite fleshy. Juniper cones aren’t red like we have seen in other lineages but they certainly garnish the attention of many a small animal looking for food.

I have only begun to scratch the surface of the fruit-like structures in gymnosperms. There is plenty of literary fodder out there for those of you who love to read about developmental biology and evolution. It is a fascinating world to uncover. More importantly, I think the fleshy “fruits” of the various gymnosperm lineages stand as a testament to the power of natural selection as a driving force for evolution on our planet. It is amazing that such distantly related plants have converged on similar seed dispersal mechanisms by so many different means.

Photo Credits: [1] [2] [3] [4] [5] [6] [7] [8]

Further Reading: [1] [2] [3] [4] [5] [6] [7]

Cycad Pollinators


When it comes to insect pollination, flowering plants get all of the attention. However, flowers aren't the only game in town. More and more we are beginning to appreciate the role insects play in the pollination of some gymnosperm lineages. For instance, did you know that many cycad species utilize insects as pollen vectors? The ways in which these charismatic gymnosperms entice insects is absolutely fascinating and well worth understanding in more detail.

Cycads or cycad-like plants were some of the earliest gymnosperm lineages to arise on this planet. They did so long before familiar insects like bees, wasps, and butterflies came onto the scene. It had long been assumed that, like a vast majority of extant gymnosperms, cycads relied on the wind to get pollen from male cones to female cones. Indeed, many species certainly utilize to wind to one degree or another. However, subsequent work on a few cycad genera revealed that wind might not cut it in most cases.

White-haired cycad ( Encephalartos friderici-guilielm i)

White-haired cycad (Encephalartos friderici-guilielmi)

It took placing living cycads into wind tunnels to obtain the first evidence that something strange might be going on with cycad pollination. The small gaps on the female cones were simply too tight for wind-blown pollen to make it to the ovules. Around the same time, researchers began noting the production of volatile odors and heat in cycad cones, providing further incentives for closer examination.

Subsequent research into cycad pollination has really started to pay off. By excluding insects from the cones, researchers have been able to demonstrate that insects are an essential factor in the pollination of many cycad species. What's more, often these relationships appear to be rather species specific.

Cycadophila yunnanensis ,  C. nigra , and other beetles on a cone of  Cycas  sp.

Cycadophila yunnanensis, C. nigra, and other beetles on a cone of Cycas sp.

By far, the bulk of cycad pollination services are being performed by beetles. This makes a lot of sense because, like cycads, beetles evolved long before bees or butterflies. Most of these belong to the superfamily Cucujoidea as well as the true weevils (Curculionidae). In some cases, beetles utilize cycad cones as places to mate and lay eggs. For instance, male and female cones of the South African cycad Encephalartos friderici-guilielmi were found to be quite attractive to at least two beetle genera. 

Beetles and their larvae were found on male cones only after they had opened and pollen was available. Researchers were even able to observe adult beetles emerging from pupae within the cones, suggesting that male cones of E. friderici-guilielmi function as brood sites. Adult beetles carrying pollen were seen leaving the male cones and visiting the female cones. The beetles would crawl all over the fuzzy outer surface of the female cones until they became receptive. At that point, the beetles wriggle inside and deposit pollen. Seed set was significantly lower when beetles were excluded.

Male cone of  Zamia furfuracea  with a mating (lek) assembly of  Rhopalotria mollis  weevils.

Male cone of Zamia furfuracea with a mating (lek) assembly of Rhopalotria mollis weevils.

For the Mexican cycad Zamia furfuracea, weevils also utilize cones as brood sites, however, the female cones go to great lengths to protect themselves from failed reproductive efforts. The adult weevils are attracted to male cones by volatile odors where they pick up pollen. The female cones are thought to also emit similar odors, however, larvae are not able to develop within the female cones. Researchers attribute this to higher levels of toxins found in female cone tissues. This kills off the beetle larvae before they can do too much damage with their feeding. This way, the cycad gets pollinated and potentially harmful herbivores are eliminated. 

Beetles also share the cycad pollination spotlight with another surprising group of insects - thrips. Thrips belong to an ancient order of insects whose origin dates back to the Permian, some 298 million years ago. Because they are plant feeders, thrips are often considered pests. However, for Australian cycads in the genus Macrozamia, they are important pollinators.

Macrozamia macleayi  female cone.

Macrozamia macleayi female cone.

Thrip pollination was studied in detail in at least two Macrozamia species, M. lucida and M. macleayi. It was noted that the male cones of these species are thermogenic, reaching peak temperatures of around   104 °F (40 °C). They also produce volatile compounds like monoterpenes as well as lots of CO2 and water vapor during this time. This spike in male cone activity also coincides with a mass exodus of thrips living within the cones.

Thrips ( Cycadothrips chadwicki ) leaving a thermogenic pollen cone of  Macrozamia lucida.

Thrips (Cycadothrips chadwicki) leaving a thermogenic pollen cone of Macrozamia lucida.

Thrips apparently enjoy cool, dry, and dark places to feed and breed. That is why they love male Macrozamia cones. However, if the thrips were to remain in the male cones only, pollination wouldn't occur. This is where all of that male cone metabolic activity comes in handy. Researchers found that the combination of rising heat and humidity, and the production of monoterpenes aggravated thrips living within the male cones, causing them to leave the cones in search of another home.

Inevitably many of these pollen-covered thrips find themselves on female Macrozamia cones. They crawl inside and find things much more to their liking. It turns out that female Macrozamia cones do not produce heat or volatile compounds. In this way, Macrozamia are insuring pollen transfer between male and female plants.

Thrips up close.

Thrips up close.

Pollination in cycads is a fascinating subject. It is a reminder that flowering plants aren't the only game in town and that insects have been providing such services for eons. Additionally, with cycads facing extinction threats on a global scale, understanding pollination is vital to preserving them into the future. Without reproduction, species will inevitably fail. Many cycads have yet to have their pollinators identified. Some cycad pollinators may even be extinct. Without boots on the ground, we may never know the full story. In truth, we have only begun to scratch the surface of cycads and their pollinators.

Photo Credits: [1] [2] [3] [4] [5] [6] [7]

Further Reading: [1] [2] [3] [4] [5] [6]

Gnetum Are Neat!


As much as I hate to admit it, when I think of gymnosperms my mind autopilots to conifers and ginkgos. I too easily forget about some of the other extant gymnosperm lineages with which we share space on this planet. Whereas one can easily pick out a conifer or a ginkgo from a lineup, some of the other gymnosperms aren't readily recognized as such. One group in particular challenges my gymnosperm search image to the extreme. I am, of course, talking about a family of gymnosperms known as Gnetaceae.

Gnetaceae is home to a single genus, Gnetum, of which there are about 40 species. They can be found growing in tropical forests throughout South America, Africa, and Southeast Asia. Gnetum essentially come in two forms, small trees and larger, scrambling vines. To most passersby, the various Gnetum species appear to be yet another tropical angiosperm with elliptical evergreen leaves. Indeed, the various species of Gnetum exhibit features that suggest a close link with flowering plants. This has led some to hypothesize that they represent a sort of living "link" between gymnosperms and angiosperms. We will get to that in a bit. First, we must taker a closer look at these odd plants.


We will start with their leaves. They are quite strange by gymnosperm standards. Gnetum produce elliptical leaves with reticulate or web-like venation. Also, their vascular tissues contain vessel elements. Such traits are usually associated with dicotyledonous angiosperms. Characteristics such as these explain why the taxonomic position of Gnetaceae has floundered a bit over the years. What about reproduction? Surely that can help gain a better understanding of where this groups stands taxonomically.

Gnetum reproductive bits require a bit of scrutiny. They are certainly not what we would call flowers. They aren't quite cones either. The technical term for gymnosperm reproductive structures are stobili. In Gnetum, these arise from the axils of the leaves. They are strange looking structures to say the least. Male strobili are long and cylindrical. They, of course, produce pollen. They also contain infertile ovules whose function I will get to in a minute. Female strobili, on the other hand, are larger and consist of ovules enclosed in a thin tissue or integument.


Pollination in Gnetum is largely accomplished via insects, though wind plays a significant role for some species as well. In insect pollinated species, the female strobili emit a strong odor and secret tiny beads of liquid called "pollination droplets." Pollination droplets are also secreted from the sterile ovules on the male strobili. It was observed that moths were the main visitors for at least two species of Gnetum.  The reason both sexes produce pollination droplets is to ensure that moths will visit multiple individuals in their search for food.

Following pollen transfer, even more angiosperm-like activity takes place. Some Gentum undergo a type of double fertilization that is quite unique among gymnosperms. Double fertilization is largely considered a defining feature of flowering plants. It is a process by which two sperm cells unite with an egg and become the embryo and the nutritive endosperm that will fuel seedling growth. Along with its cousin Ephedra, Gnetum double fertilization also involves two sperm cells, though the end result is a bit different. Instead of forming an embryo and an endosperm, double fertilization in Gentum (and Ephedra) results in the formation of two viable zygotes and no endosperm.

Fertilized seeds gradually swell into large drupe-like structures. Integument tissues develop with the seed, covering it in a fruit-like substance that turns from green to red as it matures. As far as anyone knows, birds are the main seed dispersal agents for most Gnetum species. 

Taken together, their peculiar anatomy and intriguing pollination have led many to suggest that Gnetum are more closely allied to flowering plants than they are gymnosperms. Certainly it is easy to draw lines from one dot to another in this case but the real test lies in DNA. Are they highly derived gymnosperms or possibly a so-called missing link? 

No. Recent work by the Angiosperm Phylogeny Group found that Gnetaceae are more closely related to the family Pinaceae than they are any of the sister angiosperm lineages. Their work also revealed that, although this lineage arose some 250 million years ago, much of the diversity we see today is the result of rapid speciation events during the Oligocene and Miocene. It would appear that these derived gymnosperms are not the missing link they we once thought to be. In fact, the whole concept of an evolutionary missing link is flawed to begin with. 

Still, this should not take away from fully appreciating the bizarre nature of this family. The uniqueness of the genus Gnetum is certainly worth celebrating. They serve as a reminder of just how diverse gymnosperms once were. Today they are a mere shadow of their former glory, overshadowed by the bewildering diversity of angiosperms. If you encounter a Gnetum, take the time to appreciate it as a representative of just how strange gymnosperms can be. 

Photo Credits: [1] [2] [3] [4]

Further Reading: [1] [2] [3] [4] [5]


The World's Only Parasitic Gymnosperm

When we talk about parasitic plants, 99.9% of the time we are talking about angiosperms. However, deep in the mysterious forests of New Caledonia grows the single exception to the rule. Parasitaxus usta is the only parasitic gymnosperm known to exist. The sole member of its genus, P. usta is as strange and beautiful as it is mysterious.

P. usta hails from a strange family of gymnosperms known scientifically as Podocarpaceae. Its purple coloration is absolutely stunning and is the result of high concentrations of anthocyanin pigments in the vacuoles of its cells. Although this strange gymnosperm does in fact produce chloroplasts, they are quite small and the electron transport mechanisms that make photosynthesis possible no longer function.

The true nature of its parasitic lifestyle has remained quite a mystery over the last few decades. A handful of investigations have shown it to be rather unlike any other type of parasitic plant currently known. One of the most bizarre aspects of its morphology is that P. usta does not form any roots. This provided botanists the first clues that it may be a parasite. Further investigation has suggested that, similar to parasitic ericads and orchids, P. usta utilized a fungal intermediary to parasitize the roots of its only known host, another member of the Podocarpaceae family, Falcatifolium taxoides.

Transfer of carbohydrates has been shown to occur through this fungal connection, however, P. usta also seems to obtain nitrogen and water via a direct connection to its host's xylem tissues. In this way it is similar to some mistletoes. As such, it not only can maintain a very high rate of stomatal conductance and a very low water potential, it can also produce cone crops year round. To the best of my knowledge, no other parasitic plant on Earth adopts such a strange combination of strategies.

Despite its unique status, much of the ecology of P. usta remains a complete mystery. For instance, despite being a root parasite, stems of P. usta have been found sprouting from its host tree over 3 feet above the ground. This suggests that P. usta may actually be a strange type of holoparasite. Also, it is entirely unknown how this parasitic gymnosperm becomes established on its host. To date no seed dispersal mechanisms have been described, nor are the seeds sticky. Perhaps its all a matter of chance, which would explain why so few individuals have been found. At the end of the day, the fact that it occurs on a remote island in very few locations means that this bizarrely unique gymnosperm will hold on to its mysteries for many years to come.

Photo Credit: [1] [2]

Further Reading:

The Crazy World of Cycad Sex

When we think about plants, swimming ability generally doesn't come to mind. As kids we learn that one of the major differences between plants and animals is that plants generally can't move on their own volition. Certainly there are exceptions to this rule - sensitive plants and Venus flytraps to name a few. However, there are plants out there in which swimming is a crucial component of their life cycle. Though it isn't the plant itself that does the swimming, some of the ancestral plant lineages alive today have motile sperm!

Swimming sperm is a throwback to the early days of plant evolution. Because they arose from aquatic algae, a sperm's ability to swim to an ovule helped increase the chances of reproduction. Today we see this adaptation in plants like liverworts, mosses, and ferns, which still require water to complete their life cycle. However, swimming sperm are not restricted to the cryptograms. This adaptation also can be found in cycads (as well as ginkgoes). Their sperm are super strange too. They look like little seeds covered in concentric rings of beating flagella. Unlike cryptograms, however, their swimming ability doesn't come into play until pollen comes into contact with the ovule.

Cycads are either male or female. Each produces cone-like structures called strobili. This is where the magic happens. When pollen from a male plant finds its way onto the ovule of a female, it does something quite strange. It fuses with the ovule and begins to grow. In essence it acts almost like a parasite, sucking up nutrients from the ovule tissue and destroying it in the process. This is okay because once this happens, these tissues soon become obsolete. What matters is the female gametophyte, which is embedded inside the ovule.

The pollen begins to grow a tube down into the ovule. Once it has gained enough energy, the pollen will then burst and release its sperm. This is where the flagella come in. Each sperm is like a tiny submarine, capable of swimming around inside the ovule until it locates the female gametophyte. Then and only then is fertilization accomplished. Pretty wild for an otherwise sessile organism, wouldn't you say?

Photo Credit:…/lab-5-origin-of-plants

Further Reading: