An Iris With Multiple Parents

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The Abbeville iris (Iris nelsonii) is a very special plant. It is the rarest of the so-called “Louisiana Irises” and can only be found growing naturally in one small swamp in southern Louisiana. If you are lucky, you can catch it in flower during a few short weeks in spring. The blooms come in a range of colors from reddish-purple to nearly brown, an impressive sight to see siting atop tall, slender stems. However, the most incredible aspect of the biology of this species is its origin. The Abbeville iris is the result of hybridization between not two but three different iris species.

When I found out I would be heading to Louisiana in the spring of 2019, I made sure that seeing the Abbeville iris in person was near the top of my to-do list. How could a botany nut not want to see something so special? Iris nelsonii was only officially described as a species in 1966. Prior to that, many believed hybridization played a role in its origin. Multiple aspects of its anatomy appear intermediate between other native irises. It was not until proper molecular tests were done that the picture became clear.

The Abbeville iris genome contains bits and pieces of three other irises native to Louisiana. The most obvious parent was yet another red-flowering species - the copper iris (Iris fulva). It also contained DNA from the Dixie iris (Iris hexagona) and the zig-zag iris (Iris brevicaulis). If you had a similar childhood as I did, then you may have learned in grade school biology class that hybrids are usually biological dead ends. They may exhibit lots of beneficial traits but, like mules, they are often sterile. Certainly this is frequently the case, especially for hybrid animals, however, more and more we are finding that hybridization has resulted in multiple legitimate speciation events, especially in plants.

How exactly three species of iris managed to “come together” and produce a functional species like I. nelsonii is interesting to ponder. Its three parent species each prefers a different sort of habitat than the others. For instance, the copper iris is most often found in seasonally wet, shady bottomland hardwood forests as well as the occasional roadside ditch, whereas the Dixie iris is said to prefer more open habitats like wet prairies. In a few very specific locations, however, these types of habitats can be found within relatively short distances of eachother.

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Apparently at some point in the past, a few populations swapped pollen and the eventual result was a stable hybrid that would some day be named Iris nelsonii. As mentioned, this is a rare plant. Until it was introduced to other sites to ensure its ongoing existence in the wild, the Abbeville iris was only know to occur in any significant numbers at one single locality. This necessitates the question as to whether or not this “species” is truly unique in its ecology to warrant that status. It could very well be that that single locality just happens to produce a lot of one off hybrids.

In reality, the Abbeville iris does seem to “behave” differently from any of its parental stock. For starters, it seems to perform best in habitats that are intermediate of its parental species. This alone has managed to isolate it enough to keep the Abbeville from being reabsorbed genetically by subsequent back-crossing with its parents. Another mechanism of isolation has to do with its pollinators. The Abbeville iris is intermediate in its floral morphology as well, which means that pollen placement may not readily occur when pollinators visit different iris species in succession. Also, being largely red in coloration, the Abbeville iris receives a lot of attention from hummingbirds.

Although hummingbirds do not appear to show an initial preference when given the option to visit copper and Abbeville irises at a given location, research has found that once hummingbirds visit an Abbeville iris flower, they tend to stick to that species provided enough flowers are available. As such, the Abbeville iris likely gets the bulk of the attention from local hummingbirds while it is in bloom, ensuring that its pollen is being delivered to members of its own species and not any of its progenitors. For all intents and purposes, it would appear that this hybrid iris is behaving much like a true species.

As with any rare plant, its ongoing survival in the wild is always cause for concern. Certainly Louisiana is no stranger to habitat loss and an ever-increasing human population coupled with climate change are ongoing threats to the Abbeville iris. Changes in the natural hydrologic cycle of its swampy habitat appears to have already caused a shift in its distribution. Whereas it historically could be found in abundance in the interior of the swamp, reductions in water levels have seen it move out of the swamp and into ditches where water levels remain a bit more stable year round. Also, if its habitat were to become more fragmented, the reproductive barriers that have maintained this unique species may degrade to the point in which it is absorbed back into an unstable hybrid mix with one or a couple of its parent species. Luckily for the Abbeville, offspring have been planted into at least one other location, which helps to reduce the likelihood of extinction due to a single isolated event.


Photo Credit: [1]

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

The Drought Alert System of Terrestrial Plants has an Underwater Origin

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For plants, the transition from water to land was a monumental achievement that changed our world forever. Such a transition was fraught with unique challenges, not the least of which being the ever present threat of desiccation. A new study now suggests that those early land plants already had the the tools to deal with drought and they have their aquatic algal ancestors to thank.

One of the keys to being able to survive drought is being able to detect it in the first place. Without some sort of signalling pathway, plants would not be able to close up stomata and channel vital water and nutrients to more important tissues and organs. As such, elucidating the origins and function of drought signalling pathways in plants has been of great interest to science.

One key set of pathways involved in plant drought response is collectively referred to as the “chloroplast retrograde signaling network.” I’m not even going to pretend that I understand how these pathways operate in any detail but there is one aspect of this network that is the key to this recent discovery. It involves the means by which drought and high-light conditions are sensed in one part of the plant and how that information is then communicated to the rest of the plant. When this signalling pathway is activated, the plant can then begin to produce enzymes that go on to activate defense strategies such as stomatal closure.

Chara braunii  - a modern day example of a streptophyte alga

Chara braunii - a modern day example of a streptophyte alga

The surprise came when researchers at the Australian National University, in collaboration with researchers at the University of Florida, decided to study the chloroplast retrograde signaling network in more detail. They were interested in the inner workings of this process in relation to stomata. Stomata are tiny pores on the leaves and stems of terrestrial plants that regulate the exchange of gases like CO2 and oxygen as well as water vapor. To add some controls to their experiment, the team added a few species of aquatic algae into the mix. Algae do not produce stomata and therefore they reasoned that no traces of chloroplast retrograde signaling network enzymes should be present.

This is not what happened. Instead, the team discovered that the enzymes in question also showed up in a group of algae known as the streptophytes. This was exciting because streptophyte algae hail from the lineage thought to be ancestral to all land plants. It appears that the tools necessary for terrestrial plants to survive drought were already in place before their ancestors ever left the water.

Why this is the case could have something to do with the streptophyte lifestyle. Today, these algae are known to tolerate very tough conditions. Though outright drought is rarely a threat for these aquatic algae, they nonetheless have to deal with scenarios that resemble drought such as high salinity. Streptophyte algae found growing in ephemeral pools must cope with ever increasing concentrations of salinity as the water around them evaporates. It is possible that this drought signalling pathway may have evolved as a response to hyper-saline conditions such as these. Regardless of what was going on during those early days of plant evolution, this research indicates that the ability for terrestrial plants to deal with drought evolved before their ancestors ever left the water.

The closer we look, the more we can appreciate that evolution of important traits isn’t always de novo. More often it appears that new innovations result from a retooling of of older genetic equipment. In the case of land plants, a signalling pathway that allowed their aquatic ancestors to deal with water loss was coopted later on by organs such as leaves and stems to deal with the stresses of life on land. As the old saying goes, “life uhhh… finds a way.”

Photo Credits: [1] [2]

Further Reading: [1] [2]

Rein In Those Seeds

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Plants living on islands face a bit of a conundrum. In order to get to said islands, the ancestors of those plants had to exhibit extreme seed or spore dispersal strategies. However, if plants are to persist after arriving to an island, long-distance dispersal becomes rather risky. In the case of oceanic islands, seeds or spores that travel too far end up in the water. As such, we often observe an evolutionary reduction in dispersal ability for island residents. 

Islands, however, are not always surrounded by water. You can have "islands" on land as well. The easiest example for most to picture would be the alpine zone of a mountain. Species adapted to these high-elevation habitats find it hard to compete with species native to low-elevation habitats and are therefore stuck on these "islands in the sky." Less obvious are islands created by a specific soil type. 

Take, for instance, gypseous soils. Such soils are the result of large amounts of gypsum deposits at or near the soil surface. Gypseous soils are found in large quantities throughout parts of western North America, North and South Africa, western Asia, Australia, and eastern Spain. They are largely the result of a massive climatic shift that occurred during the Eocene, some 50 million years ago. 

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Massive mountain building events during that time were causing large reductions in atmospheric CO2 concentrations. The removal of this greenhouse gas via chemical weathering caused a gradual decline in average temperatures around the world. Earth was also becoming a much drier place and throughout the areas mentioned above, hyper-saline lakes began to dry up. As they did, copious amount of minerals, including gypsum, were left behind. 

These mineral-rich soils differ from the surrounding soils in that they contain a lot of salts. Salt makes life incredibly difficult for most terrestrial plants. Life finds a way, however, and a handful of plant species inevitably adapted to these mineral-rich soils, becoming specialists in the process. They are so specialized on these types of soils that they simply cannot compete with other plant species when growing in more "normal" soils. 

Essentially, these gypseous soils function like soil or edaphic islands. Plants specialized in growing there really don't have the option to disperse far and wide. They have to rein it in or risk extirpation. For a group of plants growing in gypseous soils in western North America, this equates to changes in seed morphology. 

Mentzelia is a genus of flowering plants in the family Loasaceae. There are somewhere around 60 to 70 different species, ranging from annuals to perennials, and forbs to shrubs (they are often referred to as blazing stars but since that would lead to too much confusion with Liatris, I will continue to refer to them as Mentzelia).

For most species in this genus, seed dispersal is accomplished by wind. Plants growing on "normal" soils produce seeds with a distinct wing surrounding the seed. A decent breeze will dislodge them from their capsule, causing them blow around. With any luck some of those seeds will land in a suitable spot for germination, far from their parents. Such is not the case for all Mentzelia though. When researchers took a closer look at species that have specialized on gypseous soils, they found something intriguing. 

Mentzelia  phylogeny showing reduction in seed wings.

Mentzelia phylogeny showing reduction in seed wings.

The wings surrounding the seeds of gypseous Mentzelia were either extremely reduced in size or had disappeared altogether. Just as it makes no sense for a plant living on an oceanic island to disperse its seeds far out into the ocean, it too makes no sense for gypseous Mentzelia to disperse their seeds into soils in which they cannot compete. It is thought that limited dispersal may help reinforce the types of habitat specialization that we see in species like these Mentzelia. The next question that must be answered is whether or not such specialization and limited dispersal comes at the cost of genetic diversity. More work will be needed to understand such dynamics. 

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

Further Reading: [1] [2]

 

Hydatellaceae: The Other Basal Angiosperms

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Though rather obscure to most of the world, the genus Trithuria has enjoyed somewhat of a celebrity status in recent years. A paper published in 2007 lifted this tiny group of minuscule aquatic plants out of their spot in Poales and granted them a place among the basal angiosperm lineage Nymphaeales. This was a huge move for such little plants. 

The genus Trithuria contains 12 species, the majority of which reside in Australia, however, two species, T. inconspicua and T. konkanensis, are native to New Zealand and India. They are all aquatic herbs and their diminutive size and inconspicuous appearance make them easy to miss. For quite some time these odd plants were considered to be a group of highly reduced monocots. Their original placement was in the family Centrolepidaceae. All of that changed in 2007.

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Close inspection of Trithuria DNA told a much different story. These were not highly reduced monocots after all. Instead, multiple analyses revealed that Trithuria were actually members of the basal angiosperm lineage Nymphaeales. Together with the water lilies (Nymphaeaceae) and the fanworts (Cabombaceae), these plants are living representatives of some of the early days in flowering plant evolution. 

Of course, DNA analysis cannot stand on its own. The results of the new phylogeny had to be corroborated with anatomical evidence. Indeed, closer inspection of the anatomy of Trithuria revealed that these plants are truly distinct from members of Poales based on a series of features including furrowed pollen grains, inverted ovules, and abundant starchy seed storage tissues. Taken together, all of these lines of evidence warranted the construction of a new family - Hydatellaceae.

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The 12 species of Trithuria are rather similar in their habits. Many live a largely submerged aquatic lifestyle in shallow estuarine habitats. As you may have guessed, individual plants look like tiny grass-like rosettes. Their small flower size has lent to some of their taxonomic confusion over the years. What was once thought of as individual flowers were revealed to be clusters or heads of highly reduced individual flowers. 

Reproduction for these plants seems like a tricky affair. Some have speculated that water plays a role but close inspections of at least one species revealed that very little pollen transfer takes place in this way. Wind is probably the most common way in which pollen from one plant finds its way to another, however, the reduced size of these flowers and their annual nature means there isn't much time and pollen to go around. It is likely that most of the 12 species of Trithuria are self-pollinated. This is probably quite useful considering the unpredictable nature of their aquatic habitats. It doesn't take much for these tiny aquatic herbs to establish new populations. In total, Trithuria stands as living proof that big things often come in small packages. 

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

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

 

Gnetum Are Neat!

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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.

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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.

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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]

 

Closed on Account of Weather

Alpine and tundra zones are harsh habitats for any organism. Favorable conditions are fleeting and nasty weather can crop up in the blink of an eye. Whereas animals in these habitats can take cover, plants don't have that luxury. They are stuck in place and have to deal with whatever comes their way. Despite these challenges, myriad plant species have adapted to these conditions and thrive where other plants would perish. The intense selection pressures of these habitats have led to some fascinating evolutionary adaptations, especially when it comes to reproduction.

Take, for instance, the Arctic gentian (Gentianodes algida). This lovely plant can be found growing in alpine and tundra habitats in both North America and Asia. Like most plants of these habitats, the Arctic gentian has a low growth habit, forming a dense cluster of fleshy, narrow leaves that hug the ground. This protects the plant from blustering winds and extreme cold. From late July until early September, when the short growing season is nearly over, this wonderful plant comes into bloom. 

Clusters of white and blue speckled flowers are borne on short stems and, unlike other angiosperms that readily self-pollinate under harsh conditions, the Arctic gentian requires outcrossing to set seed. This can be troublesome. As you can imagine, pollinators can be in short supply in these habitats. What's more, with conditions changing on a dime, the flowers must be able to cope with whatever comes their way. The Arctic gentian is not helpless though. It has an interesting adaptation to these habitats and it involves movement.

Only a handful of plant species are known for their ability to move their various organs with relative rapidity. This gentian probably doesn't make that list very often. However, it probably should as its flowers are capable of responding to changes in weather by closing up shop. It is not alone in this behavior. Plenty of plant species will close their flowers on cold, dreary days. What is so special about the Arctic gentian is that it seems especially attuned to the weather. Within minutes of an incoming thunderstorm (a daily occurrence in the Rockies, for example) the Arctic gentian will close up its flowers. This is done via changes in turgor pressure within the cells. But what is the signal that cues this gentian in that a storm is fast approaching?

Researchers have investigated multiple stimuli in search of the answer. Plants don't seem to respond to changes in sunlight, wind, or humidity. Instead, temperature seemed to be the only signal capable of eliciting this response. When temperatures suddenly drop, the flowers will begin to close. Only when the temperature begins to rise will the flowers reopen. These movements are quite rapid too. Flowers will close completely within 6 - 10 minutes of a rapid decease in temperature. The reverse takes a bit longer, with most flowers needing 25 - 40 minutes to reopen.

So, why does the plant go through the trouble of closing up shop? It all has to do with sexual reproduction in these harsh conditions. Because this species doesn't self, pollen is at a premium. The plant simply can't afford the risk of rain washing it all away. The tightly closed flowers prevent that from happening. Also, wet flowers have been shown to discourage pollinators, even when favorable weather returns. Aside from interfering with pollen, rain also dilutes nectar, reducing its energy content and thus reducing the reward for any bee that would potentially visit the flower.

Being able to rapidly respond in changes in weather is important in these volatile habitats. Plants must be able to cope otherwise they risk extirpation. By closing up its flowers during inclement weather, the Arctic gentian is able to protect its vital reproductive resources.

Photo Credits: [1]

Further Reading: [1]

 

A Fern Unchanged

Ferns are old. Arising during the late Devonian period, some 360 million years ago, ferns once dominated the land. These ancient ferns were a bit different than the ferns we know today. It wasn't until roughly 145 million years ago, during the late Cretaceous period, that many extant fern families started to appear. However, a recent fossil discovery shows that at least one familiar fern was hanging out with dinosaurs as far back as 180 million years ago!

A team of scientists in Sweden recently unearthed an exquisitely preserve fossil of a fern from some early Jurassic deposits. Usually the fossilization process does not preserve very fine details, especially not at the cellular level, but that is not the case for this fossil. Falling into volcanic hydrothermal brine, the fern quickly mineralized. The speed at which the tissues of the fern were replaced by minerals preserved details that scientists usually only dream about. Clearly visible in the fossilized stem are subcellular structures like nuclei and even chromosomes in various stages of cell division!

 

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Using sophisticated microscopy techniques, the team was able to analyze the properties of the nuclei undergoing division. What they discovered is simply amazing. The number of chromosomes as well as other properties of the DNA matched a fern that is quite common in eastern North America and Asia today. This fossilized fern, as far as the team can tell, is a cinnamon fern (Osmundastrum cinnamomeum). Based on the fossil evidence, cinnamon ferns were not only around during the early Jurassic, they have remained virtually unchanged for 180 million years. Talk about a living fossil!

Further Reading: [1]

On Soil and Speciation

Lord Howe Island

Many of you will undoubtedly be familiar with some variation of this evolutionary story: A population of one species becomes geographically isolated from another population of the same species. Over time, these two separate populations gradually evolve in response to environmental pressures in their respective habitats. After enough time has elapsed, gradual genetic changes result in reproductive isolation and eventually the formation of two new species. This is called allopatric speciation and countless examples of this exist in the real world.

At the opposite end of this speciation spectrum is sympatric speciation. Under this scenario, physical isolation does not occur. Instead, through some other form of isolation, perhaps reproductive or phenological, a species gives rise to two new species despite still having contact. Examples of this in nature are far less common but various investigations have shown it is indeed possible. Despite its rarity, examples of sympatric speciation have nonetheless been found and one incredible example has occurred on a small oceanic island off the coast of Australia called Lord Howe Island.

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Lord Howe Island is relatively small, volcanic island that formed approximately 6.4–6.9 million years ago. It is home to four distinct species of palm trees from three different genera, all of which are endemic. Of these four different palms, two species, Howea belmoreana and Howea forsteriana, are quite common. Interestingly enough, H. forsteriana, commonly known as the kentia palm, is one of the most commonly grown houseplants in the entire world. However, their horticultural value is not the most interesting thing about these palms. What is most remarkable is how these two species arose. 

Multiple genetic analyses have reveled that both species originated on Lord Howe Island. This is kind of odd considering how small the island actually is. Both palms can regularly be found growing in the vicinity of one another so the big question here is what exactly drove the evolution of their common ancestor? How does a single species growing on a small, isolated island become two? The answer is quite surprising.

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When researchers took a closer look at the natural histories of these two species, they found that they were in a sense isolated from one another. The isolation is due to major phenological or timing differences in their reproductive efforts. H. forsteriana flowers roughly six weeks before H. belmoreana. Flowering time is certainly enough to drive a wedge between populations but the question that still needed answering was how do such phenological asynchronies occur, especially on an island with a land area less than 12 square kilometers? 

As it turns out, the answer all comes down to soil. Individuals of H. belmoreana are restricted to growing in neutral to acidic soils whereas H. forsteriana seems to prefer to grow in soils rich in calcarenite. These soils have a more basic pH and dominate the low lying areas of the island. Growing in calcarenite soils is stressful as they are poor in nutrients. This physiological stress has caused a shift in the way in which the flowers of H. forsteriana mature. When found growing on richer volcanic soils, the researchers noted that the flowers mature in a way that is more synchronous, not unlike the flowers of H. belmoreana.

Thanks to their attention to detailed life history events and conditions, researchers were able to show that soil preferences caused a phenological shift in the flowering of these two related species. Because they flower at completely different times when growing on their respective soil types, enough reproductive isolation was introduced to disrupt the random mating process of these wind pollinated palms. As soon as such reproductive biases are introduced, speciation can and will occur.

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

Further Reading: [1]

Mighty Magnolias

Magnolias are one of those trees that even the non-botanically minded among us will easily recognize. They are one of the more popular plant groups grown as ornamentals and their symbolism throughout human history is quite interesting. But, for all this attention, few may realize how special magnolias really are. Did you know they they are one of the most ancient flowering plant lineages in existence?

Magnolias first came on to the scene somewhere around 95 million years ago. Although they are not representative of what the earliest flowering plants may have looked like, they do offer us some interesting insights into the evolution of flowers. To start with, the flower bud is enclosed in bracts (modified leaves) instead of more differentiated sepals. The "petals" themselves are not actually petals but tepals, which are also undifferentiated. The most striking aspect of magnolia flower morphology is in the actual reproductive structures themselves.

Magnolias evolved before there were bees. Because of this, the basic structure that makes them unique was in place long before bees could work as a selective pressure in pollination. Beetles are the real pollinators of magnolia flowers. The flowers have a hardened carpel to avoid damage by their gnawing mandibles as the feed. The beetles are after the protein-rich pollen. Because the beetles are interesting in pollen and pollen alone, the flowers mature in a way that ensures cross pollination. The male parts mature first and offer said pollen. The female parts of the flower are second to mature. They produce no reward for the beetles but are instead believed to mimic the male parts, ensuring that the beetles will spend some time exploring and thus effectively pollinating the flowers.

It is pretty neat to think that you don't necessarily have to track down a dawn redwood or a gingko to see a plant that has survived major extinction events. You can find magnolias very close to home with a keen eye. Looking at one, knowing that this is a piece of biology that has worked for millennia, is quite astounding in my opinion.

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

Evolving For City Life

Urban environments pose unique challenges to any plant. Cities are generally warmer, have significantly higher CO2 levels, and experience altered levels of disturbance and precipitation patterns than do rural areas nearby. Still, many plants have taken to these concrete jungles, popping up wherever they can eke out an existence. Although we are not reinventing ecological principals in urban areas, they nonetheless present distinct selective pressures on every living thing within their jurisdiction. Evidence now suggests that urban environments are actually shaping the evolution of at least some plant species. 

Motivated by a desire to better understand how urban conditions are influencing evolution, a team of researchers based out of the University of Minnesota decided to take a closer look at a common mustard called Virginia pepperweed (Lepidium virginicum). This hardy little annual is at home wherever disturbance occurs. As such, it can be found throughout most of North America and beyond. Because it self pollinates readily, researchers were able to quantify phenotypic differences between populations growing in dense urban centers and compare them to those growing in more rural areas.

They collected seeds from numerous urban and rural populations and grew them together in a greenhouse experiment. By exposing each population to the same conditions in the greenhouse, the team were able to tease out the true phenotypic differences between these populations. 

What their data revealed were distinct differences between urban and rural populations. For starters, urban plants had larger rosettes but fewer leaves. They also bolted sooner than rural plants but then exhibited a much longer period of time between bolting and flowers. Previous studies have shown that the inflorescence of related species "accounted for 55% of a plants photosynthetic activity but only 25% of water loss." Coupled with the reduction in the number of leaves, these results suggest that urban plants are maximizing photosynthesis under drier conditions. 

Another interesting difference is that urban plants produced far more seed than their rural counterparts. This very well may be due to the fact that urban plants tended to be larger. This could also be due to reduced herbivory in urban environments, though such pressures may vary from city to city. Due to the urban heat island effect, it is likely that this could be a result of more stable temperature conditions than those experienced by their rural counterparts. Taken together, these results show that there is indeed selection for traits that allow plants to not only survive but thrive in urban environments.

Photo Credit: Wikimedia Commons

Further Reading:
[1]

Yeast in Lichens

Quite possibly one of the oldest symbiotic relationships on Earth has been hiding in plain sight all this time. Lichens have long been regarded as the poster child for symbiotic relationships. Certain species of fungi team up with specific algae and/or cyanobacteria in a sort of "you scratch my back and I'll scratch yours" type of relationship. In return for room and board the photosynthetic partner feeds the fungus. There are many variations on this theme which translates into the myriad shapes and colors of lichen species around the globe. For 150 years we have been operating under the assumption that there is only ever one species of fungus (in the phylum Ascomycota) for any given lichen. We were wrong. 

Originally thought to be contamination, researchers at the University of Montana and Perdue found gene expression belonging to the other major fungal phyla, Basidiomycota. The research team soon realized that they had uncovered something quite monumental. Lichens were harboring a partner we never knew existed. These newly discovered fungi are an entirely new lineage of yeast. What's more, this relationship has been documented in upwards of 52 other lichen genera worldwide! 

This discovery has led to another major breakthrough in lichen biology, their bizarre variety. The exact same species of fungus and alga can produce completely different lichens with wildly different attributes. Take the example of Bryoria torturosa and B. fremontii. They were thought to share the same partners and yet one is yellow and toxic whereas the other is brown and innocuous. Knowing what to look for, however, has revealed that their yeast partners are entirely different. The yeast is thought to be a sort of shield for the lichen, producing noxious acids that deter infections and predation. 

Almost overnight a new light has been shown on our lichen neighbors. These newly discovered partners aren't a recent evolutionary development. This trifecta likely stems back to the early days when little else lived on land. It just goes to show you how much we still do not know about our planet. It's nice to be reminded of this. 

Further Reading:

http://bit.ly/29WWZ2z 

Plant Plasticity

One of the central tenets of evolutionary science is that individuals within a species vary, however slightly, in their form, physiology, and behavior. Without variability, life would languish, remaining static in a soupy ooze somewhere in the oceans. Perhaps it may not have evolved in the first place. Regardless, observation and experimentation has taught us a lot about how variation among individuals or populations can drive evolution. Today I would like to introduce you to a tiny plant native to northern and western North America that is teaching us a lot about how mating systems develop in plants.

Meet Collinsia parviflora, the maiden blue eyed Mary. Few plants are as iconic to my time living out west than this wonderful little plant. Indeed, C. parviflora is highly variable. It ranges in size from 5 for 40 centimeters in height and produces lovely little flowers that range from 4 to 7 millimeters in length. The size range of these flowers is key to investigating variations in pollination strategies. 

C. parviflora has evolved what researchers refer to as a mixed mating strategy. Populations differ in that some plants self pollinate whereas others fully outcross with the help of a variety of bees. Exactly why these plants would maintain both strategies can tell us a lot about how mating systems develop in plants. What researchers have found is that there seems to be a tradeoff. 

Populations that frequently self are often located in the harshest environments. Cold temperatures and a short growing season make investing in complex floral development a risky strategy. Indeed, plants growing where environmental conditions are harshest produce smaller flowers. These small flowers pack all of their reproductive bits close together, thus increasing the chances of self fertilization. It has been found that despite the risk of inbreeding, these plants produce far more seeds than plants that produce larger flowers and experience high rates of insect pollination. 

The reasons for this are quite complex and more work is needed to be certain but it would seem that this is all an evolutionary adaptation to dealing with varied climates. With wide ranging species like C. parviflora, populations can experience highly varied environmental conditions. It would seem maladaptive to focus in on one particular reproductive strategy. As such, C. parviflora has evolved a range of possible anatomies as a way of adapting to many unique local conditions. If times are good and pollinators are abundant, it makes more sense to hedge bets on sexual reproduction whereas when conditions are poor and pollinators are scarce, it makes sense to produce offspring with a genome identical to that of the parents. If they can exist in a harsh location then so can the cloned offspring. 

Investigations into the mating system of this tiny plant has revealed that big things can really come in small packages. I miss seeing this species. Its amazing how these tiny little flowers can be so numerous as to turn wide swaths of its habitat a pleasing shade of blue. 

Further Reading:
http://www.amjbot.org/content/90/6/888.full

http://plants.usda.gov/core/profile?symbol=COPA3

The Evolution of Bulbs

Spring time is bulb time. As the winter gives way to warmer, longer days, some of our most beloved botanical neighbors begin their race for the sun. Functionally speaking, bulbs are storage organs. They are made up of a short stem surrounded by layers of fleshy leaves, which contain plenty of energy to fuel rapid growth. Their ability to maintain dormancy is something most of us will be familiar with.

They are incredibly hardy at this stage. As you might expect, bulbs are an adaptation for short growing seasons. Their ability to rapidly grow shoots gives them an advantage during short periods of time when conditions improve. Despite the energetic costs associated with supplying and maintaining such a relatively large storage organ, the ability to rapidly deploy leaves when conditions become favorable is nonetheless quite advantageous.

Contrast this with rhizomatous species, which are often associated with a life in the understory. Their ambling subterranean habit allows them to vegetatively "explore" for light and nutrients. What's more, the connected rhizomes allow the parent plant to provide nutrients to the developing clones until they grow large enough to support themselves. Under such conditions, bulbs would be at a disadvantage.

Bulbs have evolved independently throughout the angiosperm tree. Many instances of a switch from rhizomatous to bulbous growth habit occurred during the Miocene (23.03 to 5.332 million years ago) and has been associated with a global decrease in temperature and an increase in seasonality at higher latitudes. The decrease in growing season may have favored the evolution of bulbous plants such as those in the lily family. Today, we take advantage of this hardy habit, making bulbous species some of the most common plants used in gardens.

Photo Credit: Pixel Addict (Wikimedia Commons)

Further Reading: [1]
 

The Accidental Grain - How Rye Evolved Its Way Into Our Diet

Humans have been altering the genomes of plants for a very long time. Nowhere is this more apparent than in the crops we grow. These botanical mutants are pampered beasts compared to their wild congeners. It is easy to see why some traits have been selected over others, whether it be larger leaves or fruit to munch on, smaller seeds to keep them out of our way, or tough rinds to make shipping easier. However, not all of our crops have been consciously bred for our consumption. Just as many weed species are adapting to herbicides today, some species of plant were able to adapt to the more archaic methods of early farming, which allowed them to avoid the ever watchful eye of the farmer.

This concept is known as Vavilovian mimicry (sometimes referred to as crop mimicry) and it is named after the Soviet botanist and geneticist Nikolai Vavilov (who was later imprisoned and starved to death by Stalin because of his firm stance on basic genetic principles). The idea is rather simple. At its core it involves artificial selection, albeit unintentional. A wild plant species finds certain forms of agriculture appealing. It becomes an apparent weed and the farmer begins to deal with it. Perhaps this plant is a close relative and thus looks quite similar to the crop in question. As the farmer weeds out plants that look different from the crop, they may be unintentionally selecting for individual weeds that more closely resemble the crop species. Over enough seasons, only those weeds that look enough like the crop survive and reproduce, sometimes to the point in which the two are almost indistinguishable.

Rye is an interesting example of this idea. Wild rye (Secale montanum) was not intentionally grown for food. It was a weed in the fields of other crops like wheat and barley. Both wheat and barley are annual plants, producing their edible seeds at the end of their first growing season. Wild rye, however, is a perennial and does not produce seed until at least its second season. Therefore, most wild rye plants growing in wheat or barely fields are killed at the end of the season when the field gets tilled. However, there are some mutant rye plants that occasionally pop up and produce seeds in their first year.

It is believed that these mutant annual rye were harvested unintentionally and reseeded season after season. Over time, other traits likely developed to help push rye into the spotlight for these early farmers. Like many wild grasses, wild rye has weak spindles (the part that holds the seed to the plant). In the wild, this allows for efficient seed dispersal. On the farm, this is not a desirable trait as you end up quickly losing the seeds you want to harvest. Again, by accidentally selecting for mutants that also had thicker spindles and thus held on to their seeds, farmers were unintentionally domesticating rye to parallel other cereal crops. It is believed that oats (Avena sterilis) also originated in this manner.

Photo Credit: Lotte Grønkjær (http://bit.ly/1xMEfVw)

Further Reading: [1] [2]

Of Pines and Crossbills

I would like to tell you a tale that involves three major players, the pine squirrel, crossbills, and lodgepole pines. The tale I am about to tell is an interesting one about speciation due to coevolution. Few examples in nature seem to play out so wonderfully apparent as this one.

Throughout most of the Rockies where lodgepole pines grow, the main seed predator is the pine squirrel. Crossbills are out-competed for this resource wherever the squirrels are present. Because of this, lodgepole pine cones have evolved methods to deter squirrel seed predation as much as possible. In Idaho, there are a few isolated mountain tops that have lodgepole pines but no pine squirrels. On these mountains, crossbills are the main seed predator and the pines have responded accordingly.

Crossbills are named because their bills are crisscrossed so that they can pry open the cone scales to get at the seeds. The trees in these isolated mountains have developed stouter cones with thicker scales that are harder to pry apart. The birds have responded by evolving shorter, deeper bills.

The most interesting thing about this response is the effect the new bill morphology has had on mating dynamics for the crossbills on these isolated mountains. Crossbills utilize calls to select mates. Also, crossbills prefer to flock with other crossbills that have similar calls. This is because beak morphology directly affects calls so by flocking with birds with the same call, crossbills can guarantee that the flock will find food sources that they can all utilize.

As beak morphology changes, so does the sound of the calls. It has been shown that the crossbills on these isolated mountains are showing preference for mating only with crossbills with the same bill morphology. Also, because they are the only crossbills that can utilize these lodgepole pines, they are one of the few crossbill populations that can stay in an area instead of migrating large distances to find other food sources.

These pre-mating isolation rituals are so strong on these mountains that the resident crossbills rarely mate with crossbills migrating through in search of food. In essence, the coevolution between these isolated lodgepole pine populations and the resident crossbills is driving these birds towards speciation. Whats more, this coevultion is estimated to have arisen only about 5,000 to 7,000 years ago.

On a side note, this may be a world record for how many times the word "crossbill" can find its way into a story.

Photo Credit: Perry van Munster and
nebirdsplus (http://bit.ly/1N42Ytf)

Further Reading:

http://www.jstor.org/stable/10.1086/511961

http://onlinelibrary.wiley.com/doi/10.1111/j.0014-3820.2001.tb01293.x/abstract

Carnivorous Bromeliads

Brocchinia reducta

Brocchinia reducta

I would like to introduce you to quite possibly the strangest members of the Bromeliad family - those of the genus Brocchinia. Aside from the odd appearance and habits of this particular group, researchers have learned quite a bit about the Bromeliad family as a whole from studying this group. From their origins to their impressive radiation, Brocchinia offers us a window into the history of this charismatic family.

Brocchinia are considered sister to all other bromeliads. They were the first genus to diverge some 70 million years ago. Their center of origin can be traced back to the Guayana Shield, a region in the northeast corner of what is now South America. The earliest members of this group were likely terrestrial plants growing in nutrient poor areas. Surprisingly, the epiphytic nature of many bromeliad species we know and love today evolved more recently.

Brocchinia reducta

Brocchinia reducta

Since this time, Brocchinia has undergone an impressive adaptive radiation. Because they have remained specialists on nutrient poor soils, much of this radiation has to do with the evolution of nutrient acquisiton. Like its cousins, Brocchinia utilize a "tank" formed by their tightly clasped leaves.

Interestingly, at least two species of Brocchinia, (B. reducta and B. hechtioides) have taken this to the extreme and have adopted a carnivorous lifestyle. Their tall, columnar growth form coupled with slick, waxy leaves means insects can't keep a foothold and fall down into the tank. The plants sweeten the deal by luring them in with sweet secretions.

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Whether or not this was a case of true carnivory was highly debated until 2005 when a group of researchers analyzed the chemical makeup of the liquid inside the tank. They discovered that they plant was secreting an enzyme called phosphatase, which actively digest hapless insects that fall in. A true carnivore indeed!

Others have even more peculiar evolutionary adaptations for nutrient acquisition. B. tatei, for example, was discovered to house nitrogen-fixing cyanobacteria within its tank. Another species, B. acuminata, produces hallow chambers at the base of its leaves that house ant colonies. The ants pay rent via their nutrient-rich waste and voracious defense of their bromeliad home.

Brocchinnia acuminata

Brocchinnia acuminata

In total, this group is quite amazing. The amount of information we have been able to glean from studying Brocchinia has allowed us to shine a light on the whole bromeliad family. As we have also seen, the species within this group have quite the evolutionary history to tell in their own way. Brocchinia serves as a reminder to researchers blind to organismal study. We shouldn't have to study ecology at the expense of individual organisms. There is plenty to learn from both avenues of research.

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

Further Reading: [1] [2]

Nature's Radar

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Are you sitting down? You may want to before you read this. The relationship I am about to tell you about is pretty amazing. Coevolution is never a dull topic and the following example may be one of the coolest in the living world. 

Meet Marcgravia evenia. This vining plant species is native to Cuba and, like other members of its genus, relies on bats for pollination. This is nothing new. Many plant species utilize bats as pollen vectors. Bat pollinated flowers are often quite fragrant, using powerful odors to tap into the bats keen sense of smell. Marcgravia evenia is different though. This tropical vine taps into another batty sense, echolocation. 

Right above the flowers is a dish-shaped leaf. This leaf functions as a reflector for the bats sonar! Indeed, when tested, bats were twice as likely to find plants with these dish-shaped leaves than they were if the leaves were removed. This is an incredible coevolutionary adaptation! Because the vines are rare in the wild, anything that would increase the likelihood of a bat visitation would incur a considerable selective advantage. The dish-shaped leaves do just that. According to the authors of the paper, "the leaf's echoes fulfilled requirements for an effective beacon, that is, they were strong, multidirectional, and had a recognizable invariant echo signature." Nature never fails to amaze!

Photo Credit: Ralph Simon

Further Reading:

http://www.sciencemag.org/content/333/6042/631

Noble Rhubarb

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The Himalayas. If there was ever a natural wonder worthy of the title "epic" it would certainly be these towering peaks. Home to some of the tallest points on our planet, these ragged peaks are best known for the near insurmountable challenges faced by adventurers from all around the world. Considering their elevation, it would seem that permanent life simply isn't possible on these mountains. However, this could not be further from the truth. Among sprawling shrubs and diminutive herbs towers one of the most peculiar plants known to the world. To make things more interesting, it is a relative of rhubarb, a denizen of gardens and pies throughout much more hospitable climates. 

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Meet the noble rhubarb, Rheum nobile. Growing at elevations between 13,000 and 15,000 feet (4000–4800 m), this species is quite deserving of its noble status. Plants growing at such elevations face some serious challenges. Temperatures regularly drop well below freezing and there is no shortage of damaging UV radiation. As with most alpine zones, a majority of plants cope with these conditions by growing prostrate over the ground and taking what little refuge they can find behind rocks. Not Rheum nobile. This member of the buckwheat family can grow to heights of 6 feet, making it easily the tallest plant around for miles. 

The most striking feature of this plant is the large spire of translucent bracts. These modified leaves contain no chlorophyll and thus do not serve as centers for photosynthesis. Instead, these structures are there to protect and warm the plant. Tucked behind the bracts are the flowers. If they were to be exposed to the elements, they would either freeze or be fried by UV radiation. Instead, these ghostly bracts contain specialized pigments that filter out damaging UV wavelengths while at the same time creating a favorable microclimate for the flowers and seeds to develop. In essence, the plant grows its own greenhouse.

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As a result, temperatures within the plant can be as much as 10 degrees warmer than the ambient temperatures outside. At such elevations, this is a real boost to its reproductive efforts. Even more of a challenge is the fact that at this elevation, pollinators are often in short supply. Plants have to do what they can to get their attention. Not only does Rheum nobile offer a visual cue that is in stark contrast to its bleak surroundings, it also goes about attracting pollinators chemically as well.

Rheum nobile has struck up a mutualistic relationship with fungus gnats living at these altitudes. The plant produces a single chemical compound that attracts the female fungus gnats. The females lay their eggs in the developing seeds of the plant but, in return, pollinate far more flowers than they can parasitize. These organisms have managed to strike a balance in these mountains. In return for pollination, the fungus gnats have a warm place to raise their young that is sheltered from the damaging UV radiation outside. 

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

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