Raphides: A Gnarly Form of Plant Defense

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Take a bite out of a dumbcane (Dieffenbachia spp.) or a pothos (Philodendron spp.) and it won’t be long before your mouth and throat start to burn (please don’t actually do that). Eat enough of it and your symptoms may also include intense numbing, oral irritation, excessive drooling, localized swelling, and possibly even kidney and liver failure (again, please don’t). What you are experiencing is a brutal form of plant defense caused by tiny crystals called raphides.

Raphides are tiny, needle-shaped crystals made up of calcium oxalate. A lot of plants accumulate calcium oxalate. Research has shown that in doing so, plants are able to sequester excess calcium in their cells. Many plant lineages then use that calcium oxalate to make raphides. Not all raphides come in the form of needle-like crystals. Often they are ‘H’ shaped or even twinned. Others are blunt, kind of like tiny crystalline cigars.

 Cigar-shaped raphides found in the tissues of the polka dot plant ( Hypoestes phyllostachya ).

Cigar-shaped raphides found in the tissues of the polka dot plant (Hypoestes phyllostachya).

How raphides form within the plant is rather fascinating. As far as we can discern, raphide crystals form in vacuoles of specialized cells called “idioblasts.” It is thought that an exquisitely controlled scaffolding or matrix shapes the biomineralization process. To the best of my knowledge, no one has been able to reproduce this process in a laboratory setting. For now, plants are the undeniable masters of raphide manufacturing.

Within the cells, raphides are often associated with acrid and toxic proteins. Together, they comprise one hell of a defense against herbivory. Raphides are only the first part of the defensive equation. When plant tissues containing raphides are damaged, usually by chewing, the raphides shoot out of the idioblasts and into the oral cavity of the herbivore. This is where their needle shape comes in.

 Needle-like raphides extracted from the leaves of an  Epipremnum  species.

Needle-like raphides extracted from the leaves of an Epipremnum species.

Raphides wreak havoc on sensitive tissues. They literally act like tiny needles, cutting into and tearing the lining of the mouth, esophagus, and gut. This is only half of the story though. As mentioned, raphides are often packed in with acrid and toxic proteins. The laceration caused by the raphides allows these compounds to enter into the wounds. This is where things can get especially nasty. If the proteins are toxic enough, the herbivore now has far more to worry about than simply the burning sensation.

Raphides are not produced in equal amounts in all tissues. Stems tend to have more than leaves, but raphide content in leaves has also shown to be a function of leaf size. Raphides also differ from species to species. Not all plants that produce raphides produce them in the same shape and quantity. Still, more than 200 plant families contain species that have evolved this form of defense and many of our most prized houseplants fall into this category. However, this should not scare you away from these plants. Provided you or your loved ones don’t go nibbling on the leaves or stems, all will be fine. If anything, this remarkable form of plant defense should earn these plants even more respect than they already get.

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

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

Why Are Some Plants Overcompensating?

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Gardeners are all too familiar with herbivory. Countless times I have been awaiting a bloom to burst only to have the buds nipped off the night before they opened. While this can be devastating for many plant species (not to mention my sanity), for certain plant species, an encounter with a hungry herbivore may actually lead to an increase in reproductive fitness.

Overcompensation theory is the idea that, under certain conditions, plants can respond to herbivore damage by producing more shoots, flowers, and seeds. It goes without saying that when this idea was originally proposed in the late 80's, it was met with its fair share of skepticism. Why would a plant capable of producing more shoots and flowers wait to be damaged to do so? The answer may lie in in the realm of biological trade-offs.

Overcompensation may evolve in lineages that tend to grow in habitats where there is a "predictable" amount of herbivory in any given growing season, perhaps a region where large herbivores migrate through annually. Plants in these habitats may conserve dormant growing tips and valuable resources to be used once herbivory has occurred. Perhaps this also serves as a cue to upregulate antiherbivory compounds in new tissues. The trade-off is that the plants incur a cost in the form of fewer flowers and thus reduced reproduction when herbivory is low or absent.

  Scarlet gilia ( Ipomopsis aggregata )

Scarlet gilia (Ipomopsis aggregata)

It could also be that plants are exhibiting two different strategies - one to deal with competition and one to deal with herbivory. If herbivory is low, plants may become more competitive, thus favoring rapid vertical growth of one or a couple shoots. When herbivory is high, rapid vertical growth becomes disadventageous and overcompensatory branching and flowering can provide the higher fitness benefits.

These possibilities are not mutually exclusive. In fact, since the late 80's, experts now believe that overcompensation is not an "either/or" phenomenon but rather a spectrum of possibilities that are dictated by the conditions in which the plants are growing. Certainly overcompensation exists but which conditions favor it and which do not?

Research on scarlet gilia (Ipomopsis aggregata), a biennial native to western North America, suggests that overcompensation comes into play only when environmental conditions are most favorable. Soil nutrients seem to play a role in how well a plant can bounce back following herbivore damage. When resources are high, the results can be quite astounding. Early work on this species showed that under proper conditions, plants that were browsed by upwards of 95% produced 2.4 times as much seed as uneaten control plants. What's more, the resulting seedlings were twice as likely to survive than their uneaten counterparts.

Things change for scarlet gilia growing in poor conditions. Low resource availability appears to place limits on how much any given plant may respond to browsing. Also, herbivory can really hamper flowering time. Because scarlet gilia is pollen limited, anything that can cause a disruption in pollinator visits can have serious consequences for seed set. In at least one study, browsed plants flowered later and received fewer pollinator visits as a result.

More recent work has been able to add more nuance to the overcompensation story. For instance, experiments done on two subspecies of field gentian (Gentianella campestris), add further support to the idea that overcompensation is a matter of trade-offs. They showed that, whereas competition with neighboring plants alone could not explain the benefits of overcompensation, browsing certainly can.

  Field gentian ( Gentianella campestris )

Field gentian (Gentianella campestris)

Plants growing in environments where herbivory was higher overcompensated by producing more branching, more flowers, and thus more seed, all despite soil nutrients. It appears that herbivory is the strongest predictor of overcompensation for this gentian. What's more, when these data were fed into population models, only the plants that responded to herbivory by overcompensation were predicted to show any sort of population growth in the long term.

Despite all of the interest overcompensation has recieved in the botanical literature, we are only just beginning to understand the biological mechanisms that make it possible. For starters, we know that when a dominant shoot or stem gets damaged or removed, it causes a reduction in the amount of the plant hormone auxin being produced. When auxin is removed, tiny auxiliary buds at the base of the plant are able to break dormancy and begin growing.

Removal of the dominant shoot or stem can also have major impact on the number of chromosomes present in regrowing tissues. Work on Arabidopsis thaliana revealed that when the apical meristem (main growing tip of a vertical stem) was removed, the plant underwent a process called "endoreduplication" in which the cells of the growing tissues actually duplicate their entire genome without undergoing mitosis.

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Endoreduplication is a complex process with lots of biological significance but in plants it is often associated with stress responses. By duplicating the genomes of these new cells, the plants may be able to adjust more rapidly to their environment. This often manifests in changes to leaf size and shape and an uptick in plant defenses. Thus, plants may be able to fine tune the development of new tissues to overcompensate for browsing. Certainly far more work is needed to understand these mechanisms and their functions in more detail.

Overcompensation is not universal. Nonetheless, it is expected to occur in certain plants, especially those with short life cycles, and under certain environmental conditions, mainly when herbivore pressure and nutrient availability are relatively high. That being said, we still have plenty more to learn about this spectrum of strategies. When does it occur and when does it not? How common is it? What are the biological underpinnings of plants capable of overcompensation? Are some lineages more prone to overcompensation than others? Only more research can say for sure!

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

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

 

 

Giant Hogweed And Other Toxic Plants

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Everybody run, giant hogweed is coming! I am sure by now, many of you reading this will have picked up a story or two about a nasty invasive plant that will render you blind and nursing third degree burns. Indeed, giant hogweed (Heracleum mantegazzianum) is a plant worth learning how to identify. However, the tone of these articles is often one of hysterics, leaving the reader feeling like this plant is more like a Triffid, actively uprooting itself to hunt down unwary humans. Is giant hogweed worth all of this anxiety?

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Let's start with the plant itself. Giant hogweed is a member of the carrot family (Apiaceae). Its native range encompasses much of the Caucasus region and into parts of central Asia. It was (and probably still is in some areas) considered a wonderfully large and unique addition to a temperate garden. And large it is. Individual plants regularly reach heights of 6 feet (2 m) or more and some records indicate that individuals over 10 feet (3 m) in height are not unheard of.

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Because it was once a popular garden plant, this species has been introduced far outside of its native range. For many decades, giant hogweed probably lurked in the background unnoticed, its seeds finding favorable spots for germination among other weedy plants along roadsides, fallow fields, and abandoned lots. In the last few years it has grown harder to ignore. More and more plants are showing up where they shouldn't. Indeed, it seems that giant hogweed is yet another invasive species we need to get on top of. But what about all of that panic? Certainly its invasive status alone isn't what all the hype is about.

Well, like all members of the carrot family, giant hogweed produces an impressive array of chemical compounds. Many of these compounds serve to protect the plants from hungry herbivores and a plethora of microbial infections. Some of the compounds in the giant hogweed arsenal are a group known as the furocoumarins. These compounds defend the plant in a rather alarming way. These furocoumarins are phototoxic, which means when the sap gets on the body of an animal and is exposed to sunlight, they cause severe chemical burns.

 Giant hogweed when not in bloom.

Giant hogweed when not in bloom.

Stories of people being hospitalized due to an unfortunate run in with this plant make headlines wherever it pops up. That being said, simply touching the plant isn't going to hurt you. The chemicals are sloshing around in the sap of giant hogweed and the plant needs to be injured in some way before they will leak out onto whatever is hurting it. For humans, this usually occurs while mowing or weed whacking, or if a child mistakenly uses the hollow stem as a pea shooter.

With stories like this floating around, it is no wonder then why people get so upset when this plant shows up. However, I can't help but feel that this is being fed on a bit by media fear-mongering. It is worth putting giant hogweed into some practical context. It may actually alarm you to know just how many plants on the landscape have the ability to cause you harm if handled the wrong way.

 Wild parsnip ( Pastinaca sativa )

Wild parsnip (Pastinaca sativa)

Even hogweeds less robust relatives are capable of causing phototoxic reactions. I once weed whacked a large patch of Queen Anne's lace (Daucus carota) and wild parsnip (Pastinaca sativa) and ended up covered in nasty blisters the next day. I recovered but I sure did learn to give those two species more respect whenever I encountered them. Plants like poison ivy, oak, and sumac certainly cause their fair share of misery but even these do not get the sort of media attention that giant hogweed does.

Even more interesting are some of the species we actively plant in our gardens. For instance, castor bean (Ricinus communis) is quite popular among gardeners and it is responsible for producing ricin, a protein with enough killing power to bring down an adult human many times over. Take a bite out of the castor bean in your garden and it will be the last thing you ever eat. Even more potent than ricin is aconitine, an alkaloid produced by beloved garden plants like the monkshoods (Aconitum) and the larkspurs (Delphinium). This powerful alkaloid causes your nervous system to endlessly fire, leading to convulsions and death.

 Castor bean ( Ricinus communis )

Castor bean (Ricinus communis)

Similarly, a few different species of Datura are commonly grown around the world. Datura posioning is nothing to mess with and symptoms include "a complete inability to differentiate reality from fantasy; hyperthermia; tachycardia; bizarre, and possibly violent behavior; and severe mydriasis (dilated pupils) with resultant painful photophobia that can last several days." Even plants we grow for food can hurt us in bad ways. Most members of the tomato family produce a multitude of toxic alkaloids like solanine. That is why only ripe tomatoes and eggplants should ever be consumed.

 Jimsonweed ( Datura stramonium )

Jimsonweed (Datura stramonium)

In reality, I could devote an entire blog and podcast series to the chemical warfare plants have taken up during their long and complicated evolutionary history. Long story short, plants are sessile organisms that must defend themselves in order to survive and toxic chemicals are really great means to do just that. The reality is that we welcome many toxic and potentially harmful plants (both knowingly and unknowingly) into our lives and it seems slightly odd that species like giant hogweed warrant such fervor from media outlets. That being said, it is important to treat these plants with the respect they deserve. Don't bother them and they won't bother you.

So, is giant hogweed coming to attack you and your family? No. Is giant hogweed a plant worth learning to identify? Yes. Is giant hogweed dangerous to humans? Yes, but only under certain conditions.

Plants like giant hogweed are the perfect reminder as to why we must give plants more respect in our society. Teaching friends and family which plants can feed them and which plants can hurt them is something everyone should invest some time in doing. If you find giant hogweed in your area and you do not live in the Caucasus or central Asia, don't be a hero. Call a professional to come and deal with it. Otherwise, stay calm and keep on botanizing. Giant hogweed is not out to get you.

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

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

The Wild World of the Creosote Bush

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Apart from the cacti, the real rockstar of my Sonoran experience was the creosote bush (Larrea tridentata). Despite having been quite familiar with creosote as an ingredient, I admit to complete ignorance of the plant from which it originates. Having no familiarity with the Sonoran Desert ecosystem, I was walking into completely new territory in regard to the flora. It didn’t take long to notice creosote though. Once we hit the outskirts of town, it seemed to be everywhere.

If you are in the Mojave, Sonoran, and Chihuahuan Deserts of western North America, you are never far from a creosote bush. They are medium sized, slow growing shrubs with sprays of compact green leaves, tiny yellow flowers, and fuzzy seeds. Apparently what is thought of as one single species is actually made up of three different genetic populations. The differences between these has everything to do with chromosome counts. Populations in the Mojave Desert have 78 chromosomes, Sonoran populations have 52 chromosomes, and Chihuahuan have 26. This may have to do with the way in which these populations have adapted to the relative amounts of rainfall each of these deserts receive throughout the year, however, it is hard to say for sure.

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Regardless, creosote is supremely adapted to these xeric ecosystems. For starters, their branching architecture coupled with their tiny leaves are arranged so as to make the most out of favorable conditions. If you stare at these shrubs long enough, you may notice that their branches largely orient towards the southeast. Also, their leaves tend to be highly clustered along the branches. It is thought that this branching architecture allows the creosote to minimize water loss while maximizing photosynthesis.

Deserts aren’t hot 24 hours per day. Night and mornings are actually quite cool. By taking advantage of the morning sun as it rises in the east, creosote are able to open their stomata and commence photosynthesis during those few hours when evapotranspiration would be at its lowest. In doing so, they are able to minimize water loss to a large degree. Although their southeast orientation causes them to miss out on afternoon and evening sun to a large degree, the benefits of saving precious water far outweigh the loss to photosynthesis. The clustering of the leaves along the branches may also reduce overheating by providing their own shade. Coupled with their small size, this further reduces heat stress and water loss during the hottest parts of the day.

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Creosote also secrets lots of waxy, resinous compounds. These coat the leaves and to some extent the stems, making them appear lacquered. It is thought that this also helps save water by reducing water loss through the leaf cuticle. However, the sheer diversity of compounds extracted from these shrubs suggests other functions as well. It is likely that at least some of these compounds are used in defense. One study showed that when desert woodrats eat creosote leaves, the compounds within caused the rats to lose more water through their urine and feces. They also caused a reduction in the amount of energy the rats were able to absorb from food. In other words, any mammal that regularly feeds on creosote runs the risk of both dehydration and starvation. This isn’t the only interesting interaction that creosote as with rodents either. Before we get to that, however, we first need to discuss roots.

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Creosote shrubs have deep root systems that are capable of accessing soil water that more shallowly rooted plants cannot. This brings them into competition with neighboring plants in intriguing ways. When rainfall is limited, shallowly rooted species like Opuntia gain access to water before it has a chance to reach deeper creosote roots. Surprisingly this happens more often than you would think. The branching architecture of creosote is such that it tends to accumulate debris as winds blow dust around the desert landscape. As a result, the soils directly beneath creosote often contain elevated nutrients. This coupled with the added shade of the creosote canopy means that seedlings that find themselves under creosote bushes tend to do better than seedlings that germinated elsewhere. As such, creosote are considered nurse plants that actually facilitate the growth and survival of surrounding vegetation. So, if recruitment and resulting competition from vegetation can become such an issue for long term creosote survival, why then do we still so much creosote on the landscape?

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The answer may lie in rodents and other burrowing animals in these desert ecosystems. Take a look at the base of a large creosote and you will often find the ground littered with burrows. Indeed, many a mammal finds the rooting zone of the creosote shrub to be a good place to dig a den. When these animals burrow under shallowly rooted desert plants, many of them nibble on and disturb the rooting zones. Over the long-term, this can be extremely detrimental for the survival of shallow rooted species. This is not the case for creosote. Its roots run so deep that most burrowing animals cannot reach them. As such, they avoid most of the damage that other plants experience. This lends to a slight survival advantage for creosote at the expense of neighboring vegetation. In this way, rodents and other burrowing animals may actually help reduce competition for the creosote.

Barring major disturbances, creosote can live a long, long time. In fact, one particular patch of creosote growing in the Mojave Desert is thought to be one of the oldest living organisms on Earth. As creosote shrubs grow, they eventually get to a point in which their main stems break and split. From there, they begin producing new stems that radiate out in a circle from the original plant. These clones can go on growing for centuries. By calculating the average growth rate of these shrubs, experts have estimated that the Mojave specimen, affectionately referred to as the “King Clone,” is somewhere around 11,700 years old!

 The ring of creosote that is King Clone.

The ring of creosote that is King Clone.

For creosote, its slow and steady wins the race. They are a backbone of North American desert ecosystems. Their structure offers shelter, their seeds offer food, and their flowers support myriad pollinators. Creosote is one shrub worthy of our respect and admiration.

Photo Credit: [1] [2]

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

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]

Delayed Greening

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It goes without saying that leaves are vital to the existence of any photosynthetic plant. They are, after all, the food making organs. This is why plants go to great lengths to protect them. Losing leaves can be extremely costly. One of the most intriguing methods of anti-herbivory in plants is known as delayed greening. Flushes of new growth bathed in reds, whites, and light greens can color forests from top to bottom. 

Delayed greening is a matter of resource conservation and herbivore protection. The cellular machinery that makes photosynthesis possible is costly to produce. It requires large amounts of nutrients, such as nitrogen and phosphorus, that are often in short supply. If a plant can help it, its best to avoid losing a leaf chock full of these precious materials. Delayed greening does just that. 

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Essentially, the process proceeds exactly as it sounds. Young shoots and leaves gradually expand over time, becoming more green as they grow tougher and better defended. When a plant packs its leaves full of photosynthetic machinery right out of the gates, when leaves are small and tender, it runs the risk of loosing all of its investment to a hungry herbivore. In contrast, non-photosynthetic leaves are thought to be less palatable to herbivores because they simply do not have the nutritional content of photosynthetic leaves.

By delaying the development of chlorophyll until the leaf is fully expanded and a bit tougher, some plants are maximizing the chances of successfully increasing their photosynthetic capacity over time. Research has shown that plants that exhibit the delayed greening strategy experience significant reductions in the amount of herbivory over time. What they lose with the lack of photosynthesis early on they make up for in the fact that such leaves last longer.  

Photo Credits: [1] [2]

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

 

Got Herbivores? Turn Them Into Cannibals!

Plants have to deal with quite a lot in their day to day lives. They can't get up and move like animals can. Due to their sessile nature, plants rely on a suite of physical and chemical traits for defense. The world of plant chemistry is quite amazing and thanks to new research published in Nature Ecology & Evolution, it has gotten even more interesting. Under attack by herbivorous insects, some plant species are able to turn their vegetarian predators into cannibals. 

Cannibalism in insects is not unheard of, even among the herbivorous species. When the going gets tough, why not eat your sibling or your neighbor? Well, research using tomatoes and the army beetworm (Spodoptera exigua) suggests that plants might be able to induce this behavior in caterpillars long before it would happen naturally. It makes sense too. Plants that are able to induce cannibalistic behavior via chemical means not only reduce grazing pressures on their own tissues, they also reduce the number of herbivores in the system.

The chemical in question here is called methyl jasmonate. It is a volatile organic compound produced by a plethora of plant species and is thought to play in role in a diverse array of biological functions such as germination, root growth, fruit ripening, and defense. It is often released when a plant becomes damaged. Neighboring plants are able to pick up on this compound and will begin to beef up their own defenses in response. After all, if your neighbor is being attacked, there is a decent chance you will be too. 

Researchers investigating the effects of this chemical on the beetworm (a common aggricultural pest) found that plants that were treated with methyl jasmonate induced beetworms to turn on one another through cannibalism. Caterpillars hanging out on plants that were not treated with methyl jasmonate only turned to cannibalism after they had consumed all of the leaves available, if at all. 

The researchers are now gearing up to figure out whether inducing cannibalism also helps to spread disease among caterpillars. This exciting new form of plant defenses opens up doors to many new questions and potentially safer forms of pest control. Considering the near ubiquity of methyl jasmonate in the botanical world, it begs the question as to how common this form of defense really is. 

Photo Credits: [1] [2] 

Further Reading: [1] [2]

The Curious Case of a Dancing Plant

Plants aren't generally known for their speed. They tend to move at rates we simply can't perceive. The few species that exhibit rapid movements such as the sensitive plant (Mimosa pudica) and the Venus fly trap (Dionaea muscipula) have become quite famous as a result. Such movements happen in fits and bursts. These plants certainly cannot maintain such activity. However, there is another plant out there whose activity puts these other plants to shame.

Meet the telegraph plant. It has gone by a handful of scientific names since its discovery (Desmodium motorium, D. gyrans, Hedysarum gyrans, Codariocalyx motorius) but that's not why its famous. This Asian legume is renown for its maneuvers. Its compound leaves are surprisingly active organs. The larger terminal leaflets move up and down throughout the course of a day but its smaller lateral leaflets exhibit rhythmic movements on the scale of minutes.

Perhaps most famously, the leaflets show an increase in movement when exposed to music. Search the web and you will find lots of videos of the telegraph plant "dancing" to a variety of musical styles. Though entertaining, music is not why this plant moves. Having evolved long before music was ever invented, its movement must have its roots in something a bit more natural. However, despite how popular such motion has made this species over the past few centuries, their its function has remained a bit of a mystery.

Before we get into the theories, let's take a closer look at exactly how this plant moves. At the base of its leaflets there sits a ring of cells called the "pulvinus." They act a bit like water balloons and thanks to some dedicated work, it has been found that, when stimulated, these cells can quickly move water in and out via osmosis. This causes the cells to either swell or deflate and this is where the movement originates. Now, onto the why...

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A relatively recent opinion piece puts forth some of the most interesting theories on telegraph plant movement yet. The author suggests that leaflet movements are defensive in nature. They believe that the leaves could be mimicking butterfly (or some other winged arthropod movements). In doing so, it may convince gravid female insects that this individual plant is already occupied. Such strategies do indeed exist in some plant species, though via physical adornments rather than movements. Another theory this author puts forth is that their movements could also attract potential predators. By mimicking the movement of a tasty insect, it could entice birds to come in to take a closer look. Once there, they could easily find other herbivores hiding on the plant.

Another possibility related to defense is that the movements are meant to deter herbivory altogether. Studies on other plants have shown that some species can actually detect the vibrations of an insect chewing on leaves, which signals to the plant to uptick the production of defense compounds. Perhaps when sensing vibration, the telegraph plant increases its movements to knock away a hungry insect. Certainly a moving meal is less appealing than a stationary one. This is also thought to be the reason for rapid leaflet closure in sensitive plants. Hungry insects have a hard time hanging on to a plant when the leaf suddenly collapses from underneath it.

Another hypothesis is that these movements are meant to increase sun exposure. It has been discovered that far from only responding to music, the leaflets move throughout the day depending on temperature. When temperatures are low, leaflet movements are more vigorous. They eventually slow down if temperatures are high enough. This hypothesis is bolstered by the fact that movements cease once the sun goes down. In a sense, the leaflets seem to be using temperature as a means of detecting whether or not they are getting as much sun on them as possible.

In reality, it very well could be a mix of these ideas. Natural selection works like that. In the end, movement of the leaflets has certainly benefited the telegraph plant whether it be fore defense or just to take advantage of as much sun as possible. Despite centuries of popularity, this awesome little legume still has some secrets tucked away and I kind of like that about it.

NOTE: The image at the top of this page is of a time lapse and does not represent actual speed.

Photo Credit: [1]

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

Trees Know Who Nibbles

Not being able to escape form your predators makes for an interesting challenge. Plants, being the static organisms that they are, have risen to this challenge in some amazing ways. From thorns, to hairs, and even chemical warfare, there is no end to the strategies plants have evolved to discourage herbivores.

These adaptations come at a cost. Whether its physical or chemical, defenses require ample resources to produce. That is why so many plant species have evolved induced defense mechanisms. They don't bother producing their chemical cocktails until they sense that herbivores are munching on their tissues. Whereas the literature is rife with examples of insect-induced defenses in plants, few studies have ever investigated whether plants can sense mammalian herbivory. 

A 2016 paper published in Functional Ecology suggests that some trees certainly can. By looking at the responses of two different tree species, sycamore maple (Acer pseudoplantanus) and European beech (Fagus sylvatica), the research team found that trees seem to be able to distinguish between being clipped and being browsed, in this case, by deer. Deer populations are at historically high densities. This has had severe ramifications on forest health and composition. Tree saplings are especially vulnerable, so much so that we are witnessing an alarming decrease in tree replacement over time. 

The research team tested if trees could sense herbivory in a pretty ingenious way. They set out into the forest with pruners. Saplings were subjected to two different treatments - simple pruning and pruning followed by the addition of deer saliva. The team then took a look at how each tree responded on a molecular level. What they found was quite startling. Trees that were subjected to pruning alone began producing a class of hormones called jasmonates. This was not surprising as jasmonates are involved in some generic plant defenses. The most interesting results came from the treatments in which deer saliva was added.

With the addition of deer saliva to pruned beech twigs, it was discovered that the trees increased their production of metabolites related to growth of buds and leaves. They also found that the addition of deer saliva caused an increase in the production of defense compounds, specifically tannins. Tannins bind to proteins in animal guts, making them harder to digest. In maples specifically, they also found an increase in certain types of flavanols, which have shown to have anti-herbivory properties in insects and humans, however, more work is needed to see if they do in fact deter other mammalian herbivores.

Although we still don't know what exactly the trees are responding to in deer saliva, these results nonetheless offer the first evidence of trees not only being able to perceive mammalian herbivores but also responding with an increase in defense compounds. Although they only looked at two tree species, it stands to reason that such responses are wide spread throughout many plant lineages. It also calls into question previous research that used simple pruning as a proxy for herbivory. Taken together, the picture of plants being unresponsive backdrops to more charismatic fauna is entirely erroneous. Plants are proving to be quite "aware" of their environment.

Photo Credit: [1]

Further Reading: [1]

 

 

Sand Armor

Plants go through a lot to protect themselves from the hungry jaws of herbivores. They have evolved a multitude of ways in which to do this - toxins, stinging hairs, thorns, and even camouflage. And now, thanks to research by a team from UC Davis, we can add sand to this list. 

At this point you may be asking "sand?!" Stick with me here. Undoubtedly you have noticed that sticky plants often have bits of whatever substrate they are growing in stuck to their stems and leaves. You wouldn't be the first to notice this. Back in 1996 a term was coined for this very phenomenon. It has been called “psammophory,” which translates to "sand-carrying."

Over 200 species of plants hailing from 88 genera in 34 families have been identified as psammorphorous. The nature of this habit has been an object of inquiry for at least a handful of researchers over the last few decades. Hypotheses have ranged from protection from physical abrasion, reduction of water loss, reduced surface temperature, reduced solar radiation, and protection from herbivory. 

It was this last hypothesis that seemed to stick. Indeed, many plants produce crystalline structures in their tissues (phytoliths, raphides, etc., which are often silica or calcium based) to deter herbivores. Sand, being silica based, is known to cause tooth wear in humans, ungulates, and rodents. Perhaps a coating of sand is enough to drive away insects and other hungry critters looking to snack on a plant. 

By controlling the amount and color of the sand stuck to plants, the researchers were able to demonstrate that plants covered in sand were less palatable to both mammalian and insect herbivores. In total, sand-covered individuals received significantly less damage to their leaves than individuals that had their sand coat removed. By altering the color of the sand, the researchers were able to demonstrate that this was not a function of camouflage. In total, the presence of sand led to an overall increase in fitness due to a decrease in damage over time. These results are the first conclusive evidence in support of psammophory as yet another fantastic plant defense mechanism. 

Photo Credit: Franco Folini (bit.ly/1RApG1R) and Wolfram Burner (
http://bit.ly/1RMNR9V)

Further Reading:
http://onlinelibrary.wiley.com/wol1/doi/10.1890/15-1696/abstract