Are Algae Plants?

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I was nibbling on some nori the other day when a thought suddenly hit me. I don't know squat about algae. I know it comes in many shapes, sizes, and colors. I know it is that stuff that we used to throw at each other on the beach. I know that it photosynthesizes. That's about it. What are algae? Are they even plants?

The shortest answer I can give you is "it depends." The term algae is a bit nebulous in and of itself. In Latin, the word "alga" simply means "seaweed." Algae are paraphyletic, meaning they do not share a recent common ancestor with one another. In fact, without specification, algae may refer to entirely different kingdoms of life including Plantae (which is often divided in the broad sense, Archaeplastida and the narrow sense, Viridiplantae), Chromista, Protista, or Bacteria.

Caulerpa racemosa, a beautiful green algae. Photo by Nhobgood Nick Hobgood licensed under CC BY-SA 3.0

Caulerpa racemosa, a beautiful green algae. Photo by Nhobgood Nick Hobgood licensed under CC BY-SA 3.0

Taxonomy being what it is, these groupings may differ depending on who you ask. The point I am trying to make here is that algae are quite diverse from an evolutionary standpoint. Even calling them seaweed is a bit misleading as many different species of algae can be found in fresh water as well as growing on land.

Take for instance what is referred to as cyanobacteria. Known commonly as blue-green algae, colonies of these photosynthetic bacteria represent some of the earliest evidence of life in the fossil record. Remains of colonial blue-green algae have been found in rocks dating back more than 4 billion years. As a whole, these types of fossils represent nearly 7/8th of the history of life on this planet! However, they are considered bacteria, not plants.

Diatoms (Chromista) are another enormously important group. The single celled, photosynthetic organisms are encased in beautiful glass shells that make up entire layers of geologic strata. They comprise a majority of the phytoplankton in the world's oceans and are important indicators of climate. However, they belong to their own kingdom of life - Chromista or the brown algae.

To bring it back to what constitutes true plants, there is one group of algae that really started it all. It is widely believed that land plants share a close evolutionary history with a branch of green algae known as the stoneworts (order Charales). These aquatic, multicellular algae superficially resemble plants with their stalked appearance and radial leaflets.

A nice example of a stonewort (Chara braunii). Photo by Show_ryu licensed under CC BY-SA 3.0

A nice example of a stonewort (Chara braunii). Photo by Show_ryu licensed under CC BY-SA 3.0

It is likely that land plants evolved from a Chara-like ancestor that may have resembling modern day hornworts that lived in shallow freshwater inlets. Estimates of when this happen go back as far as 500 million years before present. Unfortunately, fossil evidence is sparse for this sort of thing and mostly comes in the form of fossilized spores and molecular clock calculations.

Porphyra umbilicalis  - One of the many species of red algae frequently referred to as nori. Photo by Gabriele Kothe-Heinrich licensed under CC BY-SA 3.0

Porphyra umbilicalis  - One of the many species of red algae frequently referred to as nori. Photo by Gabriele Kothe-Heinrich licensed under CC BY-SA 3.0

Now, to bring it back to what started me down this road in the first place. Nori is made from algae in the genus Porphyra, which is a type of Rhodophyta or red algae. Together with Chlorophyta (the green algae), they make up some of the most familiar groups of algae. They have also been the source of a lot of taxonomic debate. Recent phylogenetic analyses place the red algae as a sister group to all other plants starting with green algae. However, some authors prefer to take a broader look at the tree and thus lump red algae in a member of the plant kingdom. So, depending on the particular paper I am reading, the nori I am currently digesting may or may not be considered a plant in the strictest sense of the word. That being said, the lines are a bit blurry and frankly I don't really care as long as it tastes good.

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

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

 

How Air Plants Drink

 Tillandsia tectorum. Photo by Edu licensed under CC BY-NC-ND 2.0

 Tillandsia tectorum. Photo by Edu licensed under CC BY-NC-ND 2.0

Air plants (genus Tillandsia) are remarkable organisms. All it takes is seeing one in person to understand why they have achieved rock star status in the horticulture trade. Unlike what we think of as a "traditional" plant lifestyle, most species of air plants live a life free of soil. Instead, they attach themselves to the limbs and trunks of trees as well as a plethora of other surfaces. 

Living this way imposes some serious challenges. The biggest of these is the acquisition of water. Although air plants are fully capable of developing roots, these organs don't live very long and they are largely incapable of absorbing anything from the surrounding environment. The sole purpose of air plant roots is to anchor them to whatever they are growing on. How then do these plants function? How do they obtain water and nutrients? The answer to this lies in tiny structures called trichomes. 

Trichomes are what gives most air plants their silvery sheen. To fully appreciate how these marvelous structures work, one needs some serious magnification. A close inspection would reveal hollow, nail-shaped structures attached to the plant by a stem. Instead of absorbing water directly through the leaf tissues, these trichomes mediate the process and, in doing so, prevent the plant from losing more water than it gains. 

The trichomes themselves start off as living tissue. During development, however, they undergo programmed cell death, leaving them hollow. When any amount of moisture comes into contact with these trichomes, they immediately absorb that water, swelling up in the process. As they swell, they are stretched out flat along the surface of the leaf. This creates a tiny film of water between the trichomes and the rest of the leaf, which only facilitates the absorption of more water. 

Trichomes up close.  Photo by Mark Smith1989 licensed under CC BY-SA 4.0

Trichomes up close.  Photo by Mark Smith1989 licensed under CC BY-SA 4.0

Because the trichomes form a sort of conduit to the inside of the leaf, water and any nutrients dissolved within are free to move into the plant until the reach the spongy mesophyll cells inside. In this way, air plants get all of their water needs from precipitation and fog. Not all air plants have the same amount of trichomes either. In fact, trichome density can tell you a lot about the kind of environment a particular air plant calls home. 

Photo by Bernard DUPONT licensed under CC BY-SA 2.0

Photo by Bernard DUPONT licensed under CC BY-SA 2.0

The fuzzier the plant looks, the drier the habitat it can tolerate. Take, for instance, one of the fuzziest air plants - Tillandsia tectorum. This species hails from extremely arid environments in the high elevation regions of Ecuador and Peru. This species mainly relies on passing clouds and fog for its moisture needs and thus requires lots of surface area to collect said water. Now contrast that with a species like Tillandsia bulbosa, which appears to have almost no trichome cover. This smoother looking species is native to humid low-land habitats where high humidity and frequent rain provide plenty of opportunities for a drink. 

Photo by Bocabroms licensed under CC BY-SA 3.0

Photo by Bocabroms licensed under CC BY-SA 3.0

Absorbing water in this way would appear to have opened up a plethora of habitats for the genus Tillandsia. Air plants are tenacious plants and worthy of our admiration. One could learn a lot from their water savvy ways. 

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

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

Ferns Afloat

Photo by Le.Loup.Gris licensed under CC BY-SA 3.0

Photo by Le.Loup.Gris licensed under CC BY-SA 3.0

My introduction to the genus Salvinia was as an oddball aquarium plant floating in a display tank at the local pet store. I knew nothing about plants at the time but I found it to be rather charming nonetheless. Every time the green raft of leaves floated under the filter outlet, water droplets would bead off them like water off of a ducks back. Even more attractive were the upside down forest of "roots" which were actively sheltering a bunch of baby guppies. 

I grew some Salvinia for a few years before my interest in maintaining aquariums faded. I had forgotten about them for quite some time. Much later as I was diving into the wild world of botany, I started revisiting some of the plants that I had grown in various aquariums to learn more about them. It wasn't long before the memory of Salvinia returned. A quick search revealed something astonishing. Salvinia are not flowering plants. They are ferns! 

The genus Salvinia is wide spread. They can be found growing naturally throughout North, Central, and South America, the West Indies, Europe, Africa, and Madagascar. Sadly, because of their popularity as aquarium and pond plants, a few species have become extremely aggressive invaders in many water ways. More on that in a bit. 

Salvinia comprises roughly 12 different species. Of these, at least 4 are suspected to be naturally occurring hybrids. As you have probably already gathered, these ferns live out their entire lives as floating aquatic plants. Their most obvious feature are the pairs of fuzzy green leaves borne on tiny branching stems. These leaves are covered in trichomes that repel water, thus keeping them dry despite their aquatic habit. 

These are not roots! Photo by Carassiuslike licensed under CC BY-SA 4.0

These are not roots! Photo by Carassiuslike licensed under CC BY-SA 4.0

Less obvious are the other types of leaves these ferns produce. What looks like roots dangling below the water's surface are actually highly specialized, finely dissected leaves! I was super shocked to learn this and to be honest, it makes me appreciate these odd little ferns even more. It is on those underwater leaves that the spores are produced. Specialized structures called sporocarps form like tiny nodules on the tips of the leaf hairs.

Sporocarps come in two sizes, each producing its own kind of spore. Large sporocarps produce megaspores while the smaller sporocarps produce microspores. This reproductive strategy is called heterospory. Microspores germinate into gametophytes containing male sex organs or "antheridia," whereas the megaspores develop into gametophytes containing female sex organs or "archegonia." 

As I mentioned above, some species of Salvinia have become aggressive invaders, especially in tropical and sub-tropical water ways. Original introductions were likely via plants released from aquariums and ponds but their small spores and vegetative growth habit means new introductions occur all too easily. Left unchecked, invasive Salvinia can form impenetrable mats that completely cover entire bodies of water and can be upwards of 2 feet thick!

Sporocarps galore! Photo by Kenraiz licensed under CC BY-SA 4.0

Sporocarps galore! Photo by Kenraiz licensed under CC BY-SA 4.0

Lots of work has been done to find a cost effective way to control invasive Salvinia populations. A tiny weevil known scientifically as Cyrtobagous singularis has been used with great success in places like Australia. Still, the best way to fight invasive species is to prevent them from spreading into new areas. Check your boots, check your boats, and never ever dump your aquarium or pond plants into local water ways. Provided you pay attention, Salvinia are rather fascinating plants that really break the mold as far as most ferns are concerned. 

Photo Credits: [1] [2] [3]

Further Reading: [1] [2] [3]

 

How Trees Fight Disease

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Plants do not have immune systems like animals. Instead, they have evolved an entirely different way of dealing with infections. In trees, this process is known as the "compartmentalization of decay in trees" or "CODIT." CODIT is a fascinating process and many of us will recognize its physical manifestations.

In order to understand CODIT, one must know a little something about how trees grow. Trees have an amazing ability to generate new cells. However, they do not have the ability to repair damage. Instead, trees respond to disease and injury  by walling it off from their living tissues. This involves three distinct processes. The first of these has to do with minimizing the spread of damage. Trees accomplish this by strengthening the walls between cells. Essentially this begins the process of isolating whatever may be harming the living tissues.

This is done via chemical means. In the living sapwood, it is the result of changes in chemical environment within each cell. In heartwood, enzymatic changes work on the structure of the already deceased cells. Though the process is still poorly understood, these chemical changes are surprisingly similar to the process of tanning leather. Compounds like tannic and gallic acids are created, which protect tissues from further decay. They also result in a discoloration of the surrounding wood. 

The second step in the CODIT process involves the construction of new walls around the damaged area. This is where the real compartmentalization process begins. The cambium layer changes the types of cells it produces around the area so that it blocks that compartment off from the surrounding vascular tissues. These new cells also exhibit highly altered metabolisms so that they begin to produce even more compounds that help resist and hopefully stave off the spread of whatever microbes may be causing the injury. Many of the defects we see in wood products are the result of these changes.

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The third response the tree undergoes is to keep growing. New tissues grow around the infected compartment and, if the tree is healthy enough, will outpace further infection. You see, whether its bacteria, fungi, or a virus, microbes need living tissues to survive. By walling off the affected area and pumping it full of compounds that kill living tissues, the tree essentially cuts off the food supply to the disease-causing organism. Only if the tree is weakened will the infection outpace its ability to cope.

Of course, CODIT is not 100% effective. Many a tree falls victim to disease. If a tree is not killed outright, it can face years or even decades of repeated infection. This is why we see wounds on trees like perennial cankers. Even if the tree is able to successfully fight these repeat infections over a series of years, the buildup of scar tissues can effectively girdle the tree if they are severe enough.

CODIT is a well appreciated phenomenon. It has set the foundation for better tree management, especially as it relates to pruning. It is even helping us develop better controls against deadly invasive pathogens. Still, many of the underlying processes involved in this response are poorly understood. This is an area begging for deeper understanding.

Photo Credits: kaydubsthehikingscientist & Alex Shigo

Further Reading: [1]

The Strangest Wood Sorrel

Photo by Yastay licensed under CC BY-SA 4.0

Photo by Yastay licensed under CC BY-SA 4.0

For me, wood sorrels are a group of plants I usually have to look down to find. This is certainly not the case for Oxalis gigantea. Native to the coastal mountains of northern Chile, this bizarre Oxalis has forgone the traditional herbaceous habit of its cousins in exchange for a woody shrub-like growth form.

Photo by Jardín Botánico Nacional, Viña del Mar, Chile licensed under CC BY-NC 2.0

Photo by Jardín Botánico Nacional, Viña del Mar, Chile licensed under CC BY-NC 2.0

When I first laid eyes on O. gigantea, I thought I was looking at some strange form of Ocotillo. In front of me was a shrubby plant consisting of multiple upright branches that were covered in a dense layer of shiny green leaves occasionally interrupted by yellow flowers. You would think that at this point in my life, aberrant taxa would not longer surprise me. Think again. 

Photo by Marcelo_Bustamante licensed under CC BY-NC-ND 2.0

Photo by Marcelo_Bustamante licensed under CC BY-NC-ND 2.0

O. gigantea is one of the largest of the roughly 570 Oxalis species known to science. Its woody branches can grow to a height of 2 meters (6 feet)! The branches themselves are quite interesting to look at. They are covered in woody spurs from which clusters of traditional Oxalis-style leaves emerge. Each stem is capable of producing copious amounts of flowers all throughout the winter months. The flowers are said to be pollinated by hummingbirds but I was not able to find any data on this. 

Photo by Claudio Alvarado Solari licensed under CC BY-NC 2.0

Photo by Claudio Alvarado Solari licensed under CC BY-NC 2.0

This shrub is but one part of the Atacama Desert flora. This region of Chile is quite arid,  experiencing a 6 to 10 month dry season every year. What rain does come is often sparse. Any plant living there must be able to cope. And cope O. gigantea does! This oddball shrub is deciduous, dropping its leaves during the dryer months. During that time, these shrubs look pretty ragged. You would never guess just how lush it will become once the rains return. Also, it has a highly developed root system, no doubt for storing water and nutrients to tide them over.  

Photo by Jardín Botánico Nacional, Viña del Mar, Chile licensed under CC BY-NC 2.0

Photo by Jardín Botánico Nacional, Viña del Mar, Chile licensed under CC BY-NC 2.0

O. gigantea has enjoyed popularity as a horticultural oddity over the years. In fact, growing this shrub as a container plant is said to be quite easy. Despite its garden familiarity, O. gigantea is noticeably absent from the scientific literature. In writing this piece, I scoured the internet for any and all research I could find. Sadly, it simply isn't there.

This is all too often the case for unique and interesting plant species like O. gigantea. Like so many other species, it has suffered from the disdain academia has had for organismal research over the last few decades. We humans can and must do better than that. For now, what information does exist has come from horticulturists, gardeners, and avid botanizers from around the world. 

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

Further Reading: [1] [2] 

 

This Isn't Even My Final Form! A Pothos Story

Photo by Forest and Kim Starr licensed under CC BY 2.0

Photo by Forest and Kim Starr licensed under CC BY 2.0

Pothos might be one of the most widely cultivated plants in modern history. These vining aroids are so common that I don't think I can name a single person in my life that hasn't had one in their house at some point or another. Renowned for their hardy disposition and ability to handle extremely low light conditions, they have become famous the world over. They are so common that it is all too easy to forget that they have a wild origin. What's more, few of us ever get to see a mature specimen. The plants living in our homes and offices are mere juveniles, struggling to hang on as they search for a canopy that isn't there.

Trying to find information on the progenitors of these ubiquitous houseplants can be a bit confusing. To do so, one must figure out which species they are talking about. Without a proper scientific name, it is nearly impossible to know which plant to refer to. Common names aside, pothos have also undergone a lot of taxonomic revisions since their introduction to the scientific community. Also, what was thought to be a single species is actually a couple.

Photo by Forest and Kim Starr licensed under CC BY 2.0

Photo by Forest and Kim Starr licensed under CC BY 2.0

To start with, the plants you have growing in your home are no longer considered Pothos. The genus Pothos seemed to be a dumping ground for a lot of nondescript aroid vines throughout the last century. Many species were placed there until proper materials were thoroughly scrutinized. Today, what we know as a "Pothos" has been moved into the genus Epipremnum. This revision did not put all controversies to rest, however, as the morphological changes these plants go through as they age can make things quite tricky.

Photo by Tauʻolunga licensed under CC BY-SA 3.0

Photo by Tauʻolunga licensed under CC BY-SA 3.0

As I mentioned, the plants we keep in our homes are still in their juvenile form. Like all plants, these vines start out small. When they find a solid structure in a decent location, they make their bid for the canopy. Up in a tree in reach of life giving sunlight, these vines really hit their stride. They quickly grow their own version of a canopy that consists of massive leaves nearing 2 feet in length! This is when these plants begin to flower. 

As is typical for the family, the inflorescence consists of a spadix covered by a leafy spathe. The spadix itself is covered in minute flowers and these are the key to properly identifying species. When pothos first made its way into the hands of botanists, all they had to go on were the small, juvenile leaves. This is why their taxonomy had been such a mess for so long. Materials obtained in 1880 were originally named Pothos aureus. It was then moved into the genus Scindapsus in 1908.

Controversy surrounding a proper generic placement continued throughout the 1900's. Then, in the early 1960's, an aroid expert was finally able to get their hands on an inflorescence. By 1964, it was established that these plants did indeed belong in the genus Epipremnum. Sadly, confusion did not end there. The plasticity in forms and colors these vines exhibit left many confusing a handful of species within the group. At various times since the late 1960's, E. aureum and E. pinnatum have been considered two forms of the same species as well as two distinct species. The latest evidence I am aware of is that these two vines are in fact distinct enough to warrant species status. 

Photo by Mokkie licensed under CC BY-SA 3.0

Photo by Mokkie licensed under CC BY-SA 3.0

The plant we most often encounter is E. aureum. Its long history of following humans wherever they go has led to it becoming an aggressive invader throughout many regions of the world. It is considered a noxious weed in places like Australia, Southeast Asia, India, Pakistan, and Hawai'i (just to name a few). It does so well in these places that it has been a little difficult to figure out where these plants originated. Thanks to some solid detective work, E. aureum is now believed to be native to Mo'orea Island off the west coast of French Polynesia. 

Epipremnum pinnatum is similar until you see an adult plant. Photo by Mokkie licensed under CC BY-SA 3.0

Epipremnum pinnatum is similar until you see an adult plant. Photo by Mokkie licensed under CC BY-SA 3.0

It is unlikely that most folks have what it takes to grow this species to its full potential in their home. They are simply too large and require ample sunlight, nutrients, and humidity to hit their stride. Nonetheless there is something to be said for the familiarity we have with these plants. They have managed to enthrall us just enough to be a fixture in so many homes, offices, and shopping centers. It has also helped them conquer far more than the tiny Pacific island on which they evolved. Becoming an invasive species always seems to have a strong human element and this aroid is the perfect example.

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

Further Reading: [1] [2] [3] 

 

Poinsettias Wild Origins

Photo by Dinesh Valke licensed under CC BY-SA 2.0

Photo by Dinesh Valke licensed under CC BY-SA 2.0

Poinsettias are famous the world over for the splash of color they provide indoor spaces during the colder months of the year. The name "poinsettia" is seemingly synonymous with the holiday season. They are so common that it is all too easy to write them off as another disposable houseplant whose only purpose is to dazzle us with a few short weeks of reds and whites. With all of the focus on those colorful bracts, it is also easy to lose sight of the fact that these plants have wild origins. What exactly is a poinsettia and where do they come from?

Poinsettia is the common name given to a species of shrub known scientifically as Euphorbia pulcherrima. No one quite knows the exact origin of our cultivated house guests but the species itself is native to the mountains of the Pacific slope of Mexico. It is a scraggly shrub that lives in seasonally dry tropical forests. Mature specimens can grow to be so large and lanky that they almost resemble vines. As many of you know, the poinsettias we use to decorate our homes never reach the same sizes as their wild counterparts. The reason for this is because all cultivated poinsettias have been purposely infected with a bacteria that stunts their growth, keeping them small and compact.

Photo by Frank Vincentz licensed under CC BY-SA 3.0

Photo by Frank Vincentz licensed under CC BY-SA 3.0

These shrubs flower throughout winter and into spring. What we think of as large, showy, red and white flower petals are not petals at all. They are actually leafy bracts. Like a vast majority of Euphorbia species, E. pulcherrima produces a special type of inflorescence called a cyathium. The flowers themselves are small, yellow, and not much to look at with the naked eye. However, take a hand lens to them and you will reveal rather intriguing little structures. What the flowers lack in showy display is made up for by the colorful bracts, which serve similar functions as petals in that their stunning colors are there to attract potential pollinators. 

Those bracts also caught the attention of horticulturists. Because of their beauty, E. pulcherrima is one of the most widely cultivated plants in human history. As many a poinsettia owner has come to realize, the bracts do not stay colored up all year. In fact, the whole function of these bracts is to save energy on flower production by coloring up leaves that are already in place. If they don’t have to produce pigments, they won’t and for much of the year, the bracts are largely green. The key to the color change lies in Earth’s axial tilt.

Photo by Gavin White licensed under CC BY-NC-ND 2.0

Photo by Gavin White licensed under CC BY-NC-ND 2.0

As the northern hemisphere begins to tilt away from the sun, days grow shorter. In turn, poinsettia plants begin to mature their flowers. At the same time, changes within the leafy bracts cause them to start producing pigments. When the days become shorter than the nights, the plants go into full reproductive mode. Both red- and white-colored bracts have been found in the wild. As soon as the days start to grow longer than the nights, the plants switch out of reproductive mode and the dazzling color fades. In captivity, this change is mimicked by plunging plants into complete darkness for a minimum of 12 hours per day.

Another aspect worth considering about this species is its sap. Whereas most plants hailing from Euphorbiacea or spurge family contain toxic sap, the sap of E. pulcherrima is very mild in its toxicity and an absurd amount of plant material would have to be consumed to suffer any serious side effects. Certainly it serves an anti-herbivore purpose in the wild, however, as long as you're not a tiny insect or a gluttonous deer, you have nothing to worry about from this species at least. So there you have it, some food for thought if you feel the urge to purge some spurge in a post-holiday cleanse. Condsider keeping these wonderful plants in your home for another year. If you follow their natural daylight cycle, you may just coax some color out of them for many winters to come.

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

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

This Is Not The Bamboo You Are Looking For...

Photo by Benoit Giroux licensed under CC BY-NC-SA 2.0

Photo by Benoit Giroux licensed under CC BY-NC-SA 2.0

She has one, he has one, you have one, I have one, the office has one... lets just say "lucky bamboo" has made its way into many a home, office, and waiting room. Popularized by the practice of feng shui and sold for pennies on the dime by New Age stores all over the world, these plants seem to thrive on neglect. It may come as a surprise then that these plants are not a bamboo at all.

These ubiquitous home decorations are actually a species of Dracaena, Dracaena braunii to be exact. It isn't even from the same taxonomical order as bamboo. Whereas bamboo are a type of grass, D. braunii is actually more closely related to lilies. Hailing from Africa, D. braunii grows as an understory shrub in rainforests. This may explain why it does so well in the nutrient poor, low light conditions of most homes. 

In the wild, it can grow upwards of 5 feet tall. In captivity, however, it rarely exceeds 3 feet.. While most people grow theirs in a container of water and pebbles, D. braunii can do equally as well, if not better in potting mix.

Photo Credit: Benoit Giroux

Further Reading: [1]

Why Do Rhododendron Leaves Droop and Curl in the Winter?

Photo by Hanna Sörensson licensed under CC BY-SA 2.0

Photo by Hanna Sörensson licensed under CC BY-SA 2.0

Broad leaved, evergreen plants living in the temperate regions of the world face quite a challenge come winter time. Freezing temperatures, lack of water, and often intense sun can exact quite a toll on living tissues. These are likely just some of the reasons why, relatively speaking, broad leaved evergreens are a rare occurrence in temperate zones. By far the most popular group of plants in this category are the Rhododendrons.

Many a Rhodo lover has said that they can tell how cold it is outside by looking at Rhododendron leaves. Indeed, as temperatures drop, the leaves of these evergreen shrubs frequently droop and curl up like green cigars. These leaf movements do seem to be tied to the weather but their triggers and function have been the source of a lot of debate. Certainly not all Rhododendrons are cold hardy but those that are seem to benefit from reorienting their leaves. Why does this happen?

In the past it has been suggested that leaf reorientation may have something to do with reducing snow load. If the leaves were to remain horizontal, this could cause enough snow buildup to break branches. The fact that a considerable amount of ice and snow can accumulate on branches regardless of leaf position, and largely without harm, seems to suggest that this is not the case. Others have suggested that it could be a way to reduce water loss. As the leaves droop and curl, they are hypothetically increasing the humidity around their leaves and thus reducing their chances of desiccation.

Photo by Nicholas A. Tonelli licensed under CC BY 2.0

Photo by Nicholas A. Tonelli licensed under CC BY 2.0

This seems pretty far fetched for a few reasons. For starters, Rhododendron simply do not open their stomata during the colder months. By keeping them closed, there is no net transfer of water into or out of the leaves. Also, their thick, waxy cuticle keeps water within the leaves from evaporating out as well. Finally, leaf drooping and curling happens long before the ground freezes and therefore doesn't seem to be triggered by a lack of water in the environment.

The leading theories on this phenomenon seem to deal more with issues at the cellular level. The first of these has to do with the sensitive photosynthetic machinery inside the chloroplasts. Leaf drooping may actually be a response to increased light. Though we generally don't think about photosynthesis in the winter months, evergreen plants actually experience the highest light intensities of the year during this time period. Throughout the growing season, they are generally shaded by the overstory. However, once the canopy leaves fall, things change.

Because the plants are, for the most part dormant, the photosystems within the chloroplasts have no way of dissipating the energy from the incoming sunlight. Photosystem II is especially vulnerable under such scenarios. Experiments have shown that leaves that were forced to stay horizontal during the winter experienced permanent sun damage and photosynthesized considerably less than leaves that were allowed to droop once favorable temperatures returned. The thought is that by positioning the leaves vertically, the plants are reducing the amount of direct light hitting them throughout winter and therefore reducing the potential for light damage.

Photo by Lorianne DiSabato licensed under CC BY-NC-ND 2.0

Photo by Lorianne DiSabato licensed under CC BY-NC-ND 2.0

These experiments also revealed something else about the changes in leaf position when it comes to shape. As it turns out, curling made no difference in protecting the leaves from light damage. It would seem that drooping and curling are responses to two different types of environmental stress. So, why do the leaves curl?

The answer to this question is physical and one that has gained a lot of research attention in the field of cryogenics. When living tissues freeze, ice crystals build up to the point that they can rupture cell membranes. This is only exacerbated if the tissues thaw out quickly. Anyone that has ever tried to freeze and then thaw leafy vegetables knows what I am talking about.

To best preserve tissues via freezing, they must freeze quickly, which reduces the size of the ice crystals that can form, and then thaw out slowly. Researchers found that Rhododendron leaves freeze completely at temperatures below -8 degrees Celsius (17.6 degrees Fahrenheit), temperatures that occur regularly throughout the range of temperate Rhodo species. Again, experiments were able to demonstrate that flat leaves thaw much more rapidly than curled leaves. This is because a curled leaf exposes far less surface area to the warming sun than does a flat leaf. As such, curled leaves don't thaw out as fast and thus are able to avoid much of the damaging effects of daily freeze-thaw cycles.

Though these are all components of the Rhodo leaf puzzle, there is still much work to be done. What we do know is that leaf drooping and leaf curling are two separate behaviors responding to different environmental pressures. Indeed, it appears that these two traits seem to be tied to cold hardiness in the genus Rhododendron. What the exact triggers are and how they produce the changes in shape and orientation, as well as the mechanics of winter survival at the cellular level are topics that are going to require further study. Until then, I think its safe to say that we can appreciate and, to some degree, rely on the spot forecasting abilities of these wonderful shrubs.

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

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

 

The Other Balsaminaceae

Have you heard of Hydrocera triflora? I hadn't until just recently. To my surprise, Hydocera is one of only two genera that make up the family Balsaminaceae. What's more, it is a monotypic genus, with this lovely species being the single representative. There is no question that H. triflora has been completely overshadowed by its cousins, the Impatiens. In fact, literature on this species is quite scant across the board.

The first question you may be asking is what differentiates Hydrocera from the Impatiens? The differences are rather subtle. I don't know if I would have considered this plant unique enough to warrant its own genus, however, closer botanical observations tell a more nuanced story. The biggest differences between Hydrocera and Impatiens has to do with flower and fruit morphology.

Photo by Lalithamba licensed under CC BY 2.0

Photo by Lalithamba licensed under CC BY 2.0

For starters, the flowers of Hydrocera consist of a full compliment of 5 sepals and 5 petals. The petals themselves are all free from one another. Contrast this with Impatiens, whose flowers mostly consist of 3 sepals and 4 petals that are fused into pairs. The second major difference lies in the fruits. Many of us will be familiar with the explosive capsules of the various Impatiens species, each of which contains many seeds. Hydrocera on the other hand, produces berries that contain 5 seeds. Such vastly different developmental pathways in reproductive structures appear to be enough to warrant the taxonomic separation between the two genera.

The next question one might asking is why are Impatiens so diverse while Hydrocera contains only a single species? This is anyone's guess, really, but there has been at least a few hypotheses put forward that sound plausible. One has to do with habitat preference. Impatiens are largely plants of upland forests and montane environments. Such habitats may offer more potential for diversification due to high heterogeneity in resources and lots of potential for isolation of various populations. Contrast this with the habitat of H. triflora. Though it occurs throughout a wide swath of lowland Asia and India, it is semi-aquatic and these types of habitats may be more restrictive for diversification.

Another possibility has to do with seed dispersal. As mentioned above, Impatiens produce lots of seeds per capsule and, with their explosive habit, can disperse them over relatively large distances. Contrast this with Hydrocera. When the berries mature, they fall into the water and sink. They remain submerged until rot or various aquatic organisms eat away at the fleshy coating. Once the seeds have been freed, air sacs cause them to float on the currents until seasonal drying brings them back into contact with the mud. Though this is certainly an effective method for dispersal, the lower seed production rate coupled with being at the mercy of the currents means that Hydrocera is probably considerably less likely to find itself in new habitats.

Again, this is largely speculation at this point. We simply don't know enough about this oddball of the balsam world to make any serious conclusions. Luckily H. triflora is not a species under immediate threat. It seems to do quite well throughout its range, frequently occurring in flooded ditches and rice patties. Still, such stories underlie the importance of fostering and funding organism-focused research.

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

Further Reading: [1] [2]

Everlasting or Seven Years Little

Photo by Andrew massyn licensed under CC BY-SA 3.0

Photo by Andrew massyn licensed under CC BY-SA 3.0

Common names are a funny thing. Depending on the region, the use, and the culture, one plant can take on many names. In other situations, many different plants can take on a single name. Though it isn't always obvious to those unfamiliar with them, the use of scientific names alleviates these issues by standardizing the naming of things so that anyone, regardless of where they are, knows what they are referring to. That being said, sometimes common names can be entertaining.

Take for instance, plants in the genus Syncarpha. These stunning members of the family Asteraceae are endemic to the fynbos region of the Eastern and Western Cape of South Africa. In appearance they are impossible to miss. In growth habit they have been described as "woody shrublets," forming dense clusters of woody stems covered in a coat of woolly hairs. Sitting atop their meter-high stems are the flower heads.

Each flower head consists of rings of colorful paper-like bracts surrounding a dense cluster of disk flowers. The flowering period of the various species can last for weeks and spans from October, well into January. Numerous beetles can be observed visiting the flowers and often times mating as they feed on pollen. Some of the beetles can be hard to spot as they camouflage quite well atop the central disk. Some authors feel that such beetles are the main pollinators for many species within this genus.

Photo by JonRichfield licensed under CC BY-SA 3.0

Photo by JonRichfield licensed under CC BY-SA 3.0

Their mesmerizing floral displays are where their English common name of "everlasting" comes from. Due to the fact that they maintain their shape and color for a long time after being cut and dried, various Syncarpha species have been used quite a bit in the cut flower industry. A name that suggests everlasting longevity stands in stark contrast to their other common name. 

These plants are referred to as "sewejaartjie" in Afrikaans, which roughly translates to "seven years little." Why would these plants be referred to as everlasting by some and relatively ephemeral by others? It turns out, sewejaartjie is a name that has more to do with their ecology than it does their use in the floral industry.

As a whole, the 29 described species of Syncarpha are considered fire ephemerals. The fynbos is known for its fire regime and the plants that call this region home have evolved in response to this fact. Syncarpha are no exception. They flower regularly and produce copious amounts of seed but rarely live for more than 7 years after germination. Also, they do not compete well with any vegetation that is capable of shading them out.

Photo by Andrew massyn licensed under  CC BY-SA 3.0

Photo by Andrew massyn licensed under CC BY-SA 3.0

Instead, Syncarpha invest heavily in seed banking. Seeds can lie dormant in the soil for many years until fires clear the landscape of competing vegetation and release valuable nutrients into the soil. Only then will the seeds germinate. As such, the mature plants don't bother trying to survive intense ground fires. They burn up along with their neighbors, leaving plenty of seed to usher in the next generation.

Research has shown that its not the heat so much as the smoke that breaks seed dormancy in these plants. In fact, numerous experiments using liquid smoke have demonstrated that the seeds are likely triggered by some bio-active chemical within the smoke itself.

So, there you have it. Roughly 29 plants with two common names, each referring back to an interesting aspect of the biology of these plants. Despite their familiarity and relative ease of committing to memory, the common names of various species only get us so far. That's not to say we should abolish the use of common names altogether. Learning about any plant should be an all encompassing endeavor provided you know which plant you are referring to.

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

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

 

Appalachia

Welcome to Appalachia. I have fallen in love with this corner of the world in large part because of its wonderfully rich and unique flora. Join In Defense of Plants as we take a sneak peak at a mere fraction of the botanical riches these mountains hold.

Further Readings On Appalachian Flora:

http://www.indefenseofplants.com/blog...

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Producer, Writer, Creator, Host: Matt Candeias

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Of Acorns and Squirrels

I find it fun to watch squirrels frantically scurrying about during the fall. Their usually playful demeanor seems to have been replaced with more serious and directed undertones. If you watch squirrels close enough you may quickly realize that, when it comes to oaks, squirrels seem to have a knack for taxonomy. They quickly bury red oak acorns while immediately set to work on eating white oak acorns. Why is this?

Music by:
Artist: Botanist
Track: Stargazer
https://verdant-realm-botanist.bandcamp.com/

Juicy Citrus

I was enjoying some citrus the other day when I got to thinking about these peculiar fruits. They are some of my favorites yet I know very little about their development. What is a citrus fruit exactly and why are they so juicy?

To start with, citrus fruits are produced by members of the citrus or rue family - Rutaceae. Not all members of this family produce them either. Technically speaking, the oranges, lemons, limes (etc.) we eat are specialized berries called "hesperidia." They are characterized by their tough rind and juicy interior.

Following fertilization, the ovary of each flower begins to swell. On the outside of the swelling fruit you find the rind or “pericarp.” The pericarp itself has a few layers associated with it but this is where the oil-filled pits are located. Anyone that has ever squeezed an orange peel has seen these pits spurt their contents.

Citrus australasica, the Australian finger lime or caviar lime. Photo by Amada44 licensed under CC BY-SA 3.0

Citrus australasica, the Australian finger lime or caviar lime. Photo by Amada44 licensed under CC BY-SA 3.0

Inward from the pericarp are a series of segments, which are the carpels. The individual carpels are the reason why oranges can be so easily segmented. Inside each carpel is a locule. These are small cavities where the seeds are housed. Lining the walls of these loculi are tiny hairs that, as the fruit matures, gradually fill with juices.

These juice-filled hairs makeup the pulp of a citrus fruit. Look closely and you can see that they are indeed individual compartments. This not only provides some nutrients to the developing seeds, it also provides a meal for potential seed dispersers, thus increasing the chances of successful recruitment away from the parent tree.

From a quick snack I spiraled into a world of new information. It is amazing what you can learn from simple questions. As a botanically oriented person, every meal offers a sea of discovery!

Photo Credit: [1] [2]

Further Reading: [1]

Meet the Sweetfern

Photo by Sten Porse licensed under CC BY-SA 3.0

Photo by Sten Porse licensed under CC BY-SA 3.0

I remember the first time I laid my eyes on Comptonia peregrina. I was new to botany at that point in my life so I didn't have a well developed search image for these sorts of things. I was scrambling down a dry ridge with a scattered overstory of gnarly looking chestnut oaks when I saw a streak of green just below me on a sandy outcropping. They were odd looking plants, the likes of which I had never seen before.

I took out my binoculars to get a better look. What were these strange organisms? Were they ferns? No, they seemed to have woody stems. Were they gymnosperms? No, I could make out what appeared to be male catkins. Luckily I never leave home without a field guide or two. Using what little terminology I knew, I was able to narrow my focus to a plant commonly called a "sweetfern."

Photo by Megan Hansen licensed under CC BY-SA 2.0

Photo by Megan Hansen licensed under CC BY-SA 2.0

This was one of the first instances in which I grasped just how troublesome common names can be. C. peregrina is mostly definitely not a fern. It is actually an angiosperm that hails from the bay family (Myricaceae). Comptonia is a monotypic genus, with C. peregrina being the only species. It is a denizen of dry, nutrient poor habitats. As such, it has some wonderful adaptations to deal with these conditions.

To start with, its a nitrogen fixer. Similar to legumes, it forms nodules on its roots that house specialized nitrogen-fixing bacteria called rhizobia. This partnership takes care of its nitrogen needs, but what about others? One study found that not only do the roots form nodules, they also form dense cluster roots. Oddly, closer observation found that these clusters were not associated with mycorrhizal fungi. What's more, they also found that these structures were most prevalent in highly disturbed soils. It is thought that this is one way that the plant can maximize its uptake of phosphorus under the harshest growing conditions. 

Photo by Jomegat licensed under CC BY-SA 3.0

Flowering in this species is not a showy event. C. peregrina can be monoecious or dioecious, producing male and female catkins towards the ends of its shoots. After fertilization, seeds develop inside bristly fruits. Seed banking appears to be an important reproductive strategy for this species. One study found that germinated seeds had lain dormant in the soil for over 70 years until disturbance opened up the canopy above. It is expected that seeds of this species could exhibit dormancy periods of a century or more. 

In total, this is one spectacular species. Not only does it have a unique appearance, it is also extremely hardy and an excellent species to plant in drought-prone soils wherever it is native. I do see it in landscaping from time to time. If you encounter this species in the wild, take the time to observe it in detail. You will be happy you did!

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

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

Understanding the Cocklebur

Photo by Dinesh Valke from Thane, India licensed under CC BY-SA 2.0

Photo by Dinesh Valke from Thane, India licensed under CC BY-SA 2.0

Spend enough time in disturbed areas and you will certainly cross paths with a cocklebur (Xanthium strumarium). As anyone with a dog can tell you, this plant has no problems getting around. It is such a common occurrence in my life that I honestly never stopped long enough to think about its place on the taxonomic tree. I always assumed it was a cousin of the amaranths. You can imagine my surprise then when I recently learned that this hardy species is actually a member of the sunflower family (Asteraceae). 

Cocklebur doesn't seem to fit with most of its composite relatives. For starters, its flowers are not all clustered together into a single flower head. Instead, male and female flowers are borne separately on the same plant. Male flower clusters are produced at the top of the flowering stem. Being wind pollinated, they quickly dump mass quantities of pollen into the air and wither away. The female flowers are clustered lower on the stem and consist of two pistillate florets situated atop a cluster of spiny bracts. 

After fertilization, these bracts swell to form the burs that so many of us have had to dig out of the fur of our loved ones. Inside that bur resides the seeds. Cocklebur is a bit strange in the seed department as well. Instead of producing multiple seeds complete with hairy parachutes, the cocklebur produces two relatively large seeds within each bur. There is a "top" seed, which sits along the curved, convex side of the bur, and a "bottom" seed that sits along the inner flat surface of the bur. Studies performed over a century ago demonstrated that these two seeds are quite important in maintaining cocklebur on the landscape. 

Photo by Dinesh Valke from Thane, India licensed under CC BY-SA 2.0

Photo by Dinesh Valke from Thane, India licensed under CC BY-SA 2.0

You see, cocklebur is an annual. It only has one season to germinate, grow, flower, and produce the next generation. We often think of annual plants as being hardy but in reality, they are often a bit picky about when and where they will grow. For that reason, seed banking is super important. Not every year will produce favorable growing conditions so dormant seeds lying in the soil act as an insurance policy. 

Whereas the bottom seed germinates within a year and maintains the plants presence when times are good, the top seed appears to have a much longer dormancy period. These long-lived seeds can sit in the soil for decades before they decide to germinate. Before humans, when disturbance regimes were a lot less hectic, this strategy likely assured that cocklebur would manage to stick around in any given area for the long term. Whereas fast germinating seeds might have been killed off, the seeds within the seed bank could pop up whenever favorable conditions finally presented themselves. 

Today cocklebur seems to be over-insured. It is a common weed anywhere soil disturbance produces bare soils with poor drainage. The plant seems equally at home growing along scoured stream banks as it does roadsides and farm fields. It is an incredibly plastic species, tuning its growth habit to best fit whatever conditions come its way. As a result, numerous subspecies, varieties, and types have been described over the years but most are not recognized in any serious fashion. 

Sadly, cocklebur can become the villain as its burs get hopelessly tangled in hair and fur. Also, every part of the plant is extremely toxic to mammals. This plant has caused many a death in both livestock and humans. It is an ironic situation to consider that we are so good at creating the exact kind of conditions needed for this species to thrive. Love it or hate it, it is a plant worth some respect. 

Photo Credits: [1] [2] 

Further Reading: [1] [2]

Meet The Compass Plant

Few prairie plants stand out more than the compass plant (Silphium laciniatum). With its uniquely lobed leaves and a flower stalk that rises well above the rest of the vegetation, it is nearly impossible to miss. It is also quite easy to identify. Seeing a population in full bloom is truly a sight to behold but the ecology of this species makes appreciating its splendor all the more enjoyable. Today I would like to introduce you to this wonderful member of the aster family.

Any discussion about this species inevitably turns to its common name. Why compass plant? It all has to do with those lovely lobed leaves. When they first develop, the leaves of the compass plant are arranged randomly. However, within 2 to 3 weeks, the leaves will orient themselves so that their flat surfaces face east and west. They also stand vertically. This is such a reliable feature of the plant that past generations have learned to use it as a reliable way in which to orient themselves.

Photo by peganum licensed under CC BY-SA 2.0

Photo by peganum licensed under CC BY-SA 2.0

Of course, helping humans find their way is not why this feature evolved. The answer to their orientation has to do with surviving in the open habitats in which they grow. Anyone who has ever spent time hiking around in prairie-like habitats will tell you that the sun can be punishing and temperatures get hot. What's more, the range of this species overlaps with much of the rain shadow produced by the Rocky Mountains meaning water can often be in short supply.

By orienting their leaves in a vertical position with the flat surfaces face east and west, the plants are able to maximize their carbon gain as well as their water use efficiency. At the same time, the vertical orientation limits the amount of direct solar radiation hitting the leaf. In essence, compass plant leaf orientation has evolved in response to the stresses of their environment. Research has shown that the sun's position in early morning is the stimulus that the plant cues in on during leaf growth.

Aside from its fascinating biology, the compass plant is also ecologically important. Myriad pollinators visit its large composite flowers and many different species of birds feed on their seeds. However, it is the insect community supported by the compass plant that is most impressive. Surveys have shown that nearly 80 different species of insect can be found living on or in it stems. Many of these are gall making wasps and their respective parasitoids. With individual plants producing up to 12 stems each, these numbers soon become overwhelming. Needless to say, this is one of the cornerstone plant species anywhere it grows naturally.

Photo Credit: [1]

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

 

How Plants Perceive Light

For all but a handful of plants, sunlight is vital to their existence. It provides the energy needed to break molecules of CO2 and water in order to synthesize carbohydrates. It is no wonder then that plants are incredibly attuned to their light environment. They grow towards it, they compete for it, they simply can't live without it. Exactly how plants (as well as many bacteria and even some fungi) perceive light is quite fascinating. It involves a small family of proteins called "phytochromes" whose chemical properties function like an on and off switch. Today I would like to briefly introduce you to this system. 

The activity of phytochrome proteins can be quite complex. In fact, many aspects of this system are still awaiting discovery. Still, we know pretty well how the phytochrome system functions with light and it all comes down to the color red. Pure sunlight is white light. It contains all of the wavelengths in the visible spectrum and then some. Phytochrome responds to two areas of this spectrum: red wavelengths of around 667 nm and far-red wavelengths around 730 nm. 

Photo by byr7 licensed under CC BY 2.0

Photo by byr7 licensed under CC BY 2.0

This range of the spectrum is quite useful when it comes to assessing whether or not there is enough light for photosynthesis. Unfiltered sunlight contains the most red light. As sunlight passes through the leaves of the canopy or as the sun sets, the ratio of far-red light increases. Far-red light is not conducive to photosynthesis. As such, the long-term survival of photosynthetic organisms is tied to figuring out the relative abundance of red and far-red light. 

This is where the phytochrome system comes in. It comes in two forms - an active form and an inactive form. When the inactive form absorbs red wavelengths, it is converted to its active form. This is the form that signals to the plant that there is enough light for physiological activity. When the ratio of wavelengths hitting the active form becomes dominated by far-red wavelengths (as it does when a plant is shaded or when the sun sets), the phytochrome is converted back into its inactive form. This in turn signals the plant to shut down many of the physiological activities within.

The structure of phytochrome in its inactive form (left) and active form (right).

The structure of phytochrome in its inactive form (left) and active form (right).

This on and off switch is how plants regulate everything from growth to flowering. The ratio of active to inactive forms can tell some plants what time of year it is. If there is more inactive form within its tissues, the plant "knows" that the days are growing shorter. Phytochrome is also involved in the number and the size of leaves that a plant will produce. Similarly, it is how plants know when they are being shaded out by their neighbors. The more neighboring plants there are, the more filtered the sunlight becomes and the ratio of far-red light increases. It is even involved in the process of seed germination. Small seeds that don't have enough food reserves (think lettuce seeds) will only germinate once their phytochrome is converted to its active form. In doing so, they ensure that they aren't germinating in an environment with too much shade or deep under the soil. 

Scientists are still working out exactly how the phytochrome system is able to regulate so many functions in plants. In some cases it can directly interact with molecules in the cytoplasm of plant cells. In other cases, it is transported into the nucleus where it can activate or deactivate particular genes. What we do know is that the phytochrome system is vitally important not just for the organisms that produce it, but for life as we know it. Without plants there could be no life on this planet.

Photo Credits: [1] [2]

Further Reading: [1] [2] [3] [4]

Plant Architecture and Its Evolutionary Implications

I make it a point that during my field season I enjoy my breakfast out on the deck. It is situated about halfway up the canopy of the surrounding forest and offers a unique perspective that is hard to come by elsewhere. Instead of looking up at the trees, I am situated in a way that allows for a better understanding of the overall structure of the forest. Its this perspective that generates a lot of different questions about what it takes to survive in a forested ecosystem, especially as it relates to sessile organisms like plants.

Quite possibly my favorite plants to observe from the deck are the pagoda dogwoods (Cornus alternifolia). Hinted at by its common name, this wonderful small tree takes on a pagoda-like growth form with its stacked, horizontal branching pattern. It is unmistakable against the backdrop of other small trees and shrubs in the mid canopy. The fact that it, as well as many other plant species, can be readily recognized and identified on shape alone will not be lost on most plant enthusiasts.

The fact that diagrams like these exist in tree guides is proof of the utility of this concept.

The fact that diagrams like these exist in tree guides is proof of the utility of this concept.

Even without the proper vocabulary to describe their forms, anyone with a keen search image understands there is a gestalt to most species and that there is more to this than simply fodder for dichotomous keys. The overall form of plants has garnered attention from a variety of disciplines. Such investigations involve fields of study like theoretical and quantitative biology to engineering and biomechanics. It has even been used to understand how life may evolve on other planets. It is a fascinating field of investigation and one worth spending time in the literature. 

Some of the pioneering work on this subject started with two European botanists: Dr. Francis Hallé and Dr. Roelof Oldeman. Together they worked on conceptual models of tree architecture. Using a plethora of empirical studies on whether a tree branches or doesn't, where branches occurs, how shoots extend, how branches differentiate, and whether reproductive structures are terminal or lateral, they were able to reduce the total number of tree forms down to 23 basic architectural models (pictured above). Each model describes the overall pattern with which plants grow, branch, and produce reproductive structures. At the core of these models is the concept of reiteration or the repitition of form in repeatable sub-units. The models themselves were given neutral names that reflect the botanists that provided the groundwork necessary to understand them.  

Despite the fact that these models are based on investigations of tropical tree species, they are largely applicable to all plant types whether they are woody or herbaceous and whether they occur in the temperate zone or the tropics. The models themselves do not represent precise categories but rather points on a spectrum of architectural possibilities. Some plants may be intermediate between two forms or share features of more than one model. It should also be noted that most trees conform to a specific model for only a limited time period during their early years of development. Random or stochastic events throughout a trees life greatly influence its overall structure as it continues to grow. The authors are careful to point out that a trees crown is the result of all the deterministic, opportunistic, and chance events in its lifetime.  

Despite these exceptions, the adherence of most plants to these 23 basic models is quite astounding. Although many of the 23 models are only found in the tropics (likely an artifact of the higher number of species in the tropics than in the temperate zones), they provide accurate reference points for further study. For instance, the restriction of some growth forms to the tropics raises intriguing questions. What is it about tropical habitats that restricts models such as Nozeran's (represented by chocolate - Theobroma cacao) and Aubréville's (represented by the sea almond - Terminalia catappa) to these tropical environments? It likely has to do with the way in which lateral buds develop. In these models, buds develop without a dormancy stage, a characteristic that is not possible in the seasonal climates of the temperate zones. 

Reiteration is an important process in plant architectural development in which plants repeat their basic model. This is especially important in repairing damage. [SOURCE]

Another interesting finding borne from these models is that there doesn't seem to be strong correlations between architecture and phylogeny. Although species within a specific genus often share similar architecture, there are plenty of exceptions. What's more, the same form can occur in unrelated species. For instance, Aubréville's model occurs in at least 19 different families. Similarly, the family Icacinaceae, which contains somewhere between 300 and 400 species, exhibits at least 7 of the different models. Alternatively, some families are architecturally quite simple. For instance the gymnosperms are considered architecturally poor, exhibiting only 4 of the different models. Even large families of flowering plants can be architecturally simplistic. Take the Fabaceae, which is largely comprised of plants exhibiting Troll's model. 

So, at this point the question of what is governing these models becomes apparent. If most plants can be reduced to these growth forms at some point in their life then there must be some aspect of the physical world that has shaped their evolution through time. Additionally, how does plant architecture at the physical level scale up to the level of a forest? Questions such as this are fundamental to our understanding of not only plants as organisms, but the role they play in shaping the world around us. 

Although many scientists have attempted to tackle these sorts of questions, I want to highlight the work of one individual in particular - Dr. Karl Niklas. His work utilizes mathematics to explain plant growth and form in relation to four basic physical constraints:

1) Plants have to capture sunlight and avoid shading their own leaves.

2) Plants have to support themselves structurally.

3) Plants have to conduct water to their various tissues.

4) Plants must be able to reproduce effectively.

Using these basic constraints, Dr. Niklas built a mathematical simulation of plant evolution. His model starts out as a "universe" containing billions of possible plant architectures. The model then assesses each of these forms on how well they are able to grow, survive, and reproduce through time. The model is then allowed to change environmental conditions to assess how these various forms perform and how they evolve. 

An example of Niklas' model showing how simple branching pattern (bottom) can evolve over time into more complex, yet familiar, forms (top).

An example of Niklas' model showing how simple branching pattern (bottom) can evolve over time into more complex, yet familiar, forms (top).

The most remarkable part of this model is that it inevitably produces all sorts of familiar plant forms, such as those we see in lycophytes, ferns, as well as many of the tree architectural models mentioned above. What's more, later iterations of the model do an amazingly accurate job at predicting forest structure dynamics such as self-thinning, mortality, and realistic size/frequency distributions of various species. 

It would appear that the rules governing what we know as a plant are to some degree universal. Because constraints such as light capture and the passive movement of water are firmly grounded in the laws of physics, it makes sense that the successful plant architectures we know and love today (as well as those present through the long history of plant evolution on this planet) are in large part a result of these physical constraints. It also begs the question of what photosynthetic life would look like on other planets. It is likely that if life arose and made its living in a similar way, familiar "plant" architecture could very well exist on other planets.

Listen to my interview with Dr. Karl Niklas here.


Photo Credits: [1] [2] [3]

Further Reading: [1] [2] [3] [4] [5]

Meet Jones' Columbine

Photo by Steve licensed under CC BY-NC-SA 2.0

Photo by Steve licensed under CC BY-NC-SA 2.0

Meet Aquilegia jonesii. This interesting little columbine can be found growing in a narrow range along the northern Rockies. It only grows in alpine and sub-alpine zones, making it quite rare. It has a cushion-like growth form to shield it from the elements but disproportionately large flowers. It is a lucky day if one stumbles across this species! 

Fun Fact: Both the common name and generic name of the flowers referred to collectively as "columbines" have their origins in ornithology? 

That's right, the genus to which they belong, Aquilegia, can trace its origin to the word "aquila," which is Latin for "eagle." When the genus was being described, it was felt that the flower resembled the claw of an eagle. 

The word "columbine" has it's origins in the word "columba," which is Latin for "pigeon" or "dove." Early botanical enthusiasts felt that the nectar spurs resembled the heads of a group of doves. 

More and more I am coming on board with the idea that etymology can be quite fun.

Photo Credit: Steve (http://bit.ly/NbGbmz)

Further Reading: [1] [2] [3]