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]

 

 

Light Pollution and Plants

I love walking around my town at night. Things really seem to slow down when the sun sets. Growing up in the country, my evening walks were lit only by the moon. Now that I live in civilization, however, street lights punctuate the darkness on every block. Walking around I can't help but wonder what all of this artificial light is doing to our photosynthetic neighbors. 

The vast majority of plants need light to make food. It doesn't matter if this light comes from the sun or a high powered electric light, as long as it is strong enough for photosynthesis. Even weaker wavelengths of light serve a purpose for our botanical friends. Plants can sense the relative length of uninterrupted darkness in their environment and they use that information for myriad internal processes. Its this dependence on light that makes many plant species vulnerable to our addiction to artificial lighting.

Just because a light isn't strong enough for photosynthesis doesn't mean it isn't affecting nearby plants. This is especially true for plants that use day length for timing events like bud break, flowering, and dormancy. The type of lighting favored by most municipalities emit wavelengths that peak especially high in the red to far-red ratio of the electromagnetic spectrum, which makes them particularly adept at disrupting plant photoperiods.

One of the most obvious effects of artificial lighting on plants can readily be seen in street trees growing in temperate regions. Though light sensitivity varies from species to species, trees growing near street lights tend to hold onto their leaves much longer in the fall than trees farther away. Because artificial lighting is enough to trick the red to far-red receptors in plants, it can "convince" trees that the days are longer than they actually are. Additional photosynthesis may not seem that bad but holding onto leaves longer makes trees more susceptible to ice damage. 

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The effects of artificial lighting continues into spring as well. Trees growing near lights tend to break buds and flower earlier in the spring. This too makes them susceptible to frost damage. Early flowering plants run the risk of losing their entire reproductive effort by blooming before the threat of frost is gone. This can really mess up their relationship with pollinators. 

The effects of artificial lighting can even influence the way in which plants grow. Research has found that plants growing near street lights had larger leaves with more stomatal pores and these pores remained open for considerably longer than plants growing under unlit night conditions. This made them more susceptible to pollution and drought, two stressors that are all too common in urban environments. This issues is made much worse if the artificial lighting never turns off throughout the night. 

Artificial lighting affects more than just plant physiology too. Scaling up, the effects of night lights can have whole ecosystem consequences. For instance, researchers found that artificial lighting was enough to change the entire composition of grassland communities. Some plants responded well to artificial lights, producing more biomass and vegetative offshoots to the point that they pushed out other species. This was compounded by the change in reproductive output, with certain species showing higher seed production than others.

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Changes in plant physiology, phenology, and composition also affect myriad other organisms in the environment. Changes in the timing of flowering or bud break can disrupt things like insects and birds that rely on these events for food and shelter. Research even suggests that forest regeneration is being altered by artificial lighting. Seed dispersers such as bats often will not fly into well-lit areas at night, therefore reducing the amount of seeds falling in those areas. Such research is still in its infancy meaning we have a lot more to learn about how artificial lighting is disrupting natural events.

Light pollution is so much more than an aesthetic issue. Artificial lighting is clearly having pronounced effects on plant life. Disrupt plants and you disrupt life as we know it. Certainly more work is needed to tease out all the ways in which lights influence plants, however, it is clear that we must work hard on reducing light pollution around the globe.

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

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

Resisting the Wind

Have you ever wondered how some plants can withstand heavy winds? At lease one group, the cattails, produce specialized support structures within their cells to cope with winds. This is great, especially when growing near a large, windy water source.

A team of researchers recently took a much closer look at the leaf cells of a variety of cattail species (genus Typha). For decades, there has been knowledge of fibers that traverse the air chambers within the cells. These have largely been ignored but as it turns out, they indeed serve a purpose.

 (A) Longitudinal section showing the fibre cables anchored in diaphragms composed of aerenchyma tissue. (B) Longitudinal section showing the fibre cables anchored in diaphragms composed of aerenchyma tissue. (C) Cross section. The thick ventral (v)and dorsal (d) surfaces of the leaf are separated by thick partitions (P) that run the length of the leaf. Thin diaphragms (D) connected perpendicular to the thick partitionsare traversed by very fine fibre cables (FC), which are anchored to them. This construction has often been compared to sandwich-type construction, giving a low-density structure of high stiffness and strength (Rowlatt and Morshead, 1992)

(A) Longitudinal section showing the fibre cables anchored in diaphragms composed of aerenchyma tissue. (B) Longitudinal section showing the fibre cables anchored in diaphragms composed of aerenchyma tissue. (C) Cross section. The thick ventral (v)and dorsal (d) surfaces of the leaf are separated by thick partitions (P) that run the length of the leaf. Thin diaphragms (D) connected perpendicular to the thick partitionsare traversed by very fine fibre cables (FC), which are anchored to them. This construction has often been compared to sandwich-type construction, giving a low-density structure of high stiffness and strength (Rowlatt and Morshead, 1992)

As any good engineer will tell you, if a structure is to remain sound, it needs multiple avenues in which stress can be redistributed. The same goes for living structures like leaves. The fibers are arranged within the cells makes them quite strong under tension. In this way, multiple load paths are created to distribute the stress of high winds on the leaves. We like to take credit for most of our ideas but, time and again, nature beat us to it first.

Photo Credit: [1] [2]

Further Reading: [1]

What Are Plants Made Of?

Have you ever thought about what plants are made of? I mean, really thought about it. Strip away all the splendor and glory of all the different plant species on this planet and really take a close look at how plants grow and make more plants. It is a fascinating realm and it all has to do with photosynthesis. To go from photons given off by our nearest star to a full grown plant is quite the journey and, at the end of that journey, you may be surprised to learn what plants are all about.

It starts with photons. Leaving the sun they travel out into the universe. Some eventually collide with Earth and make their way to the surface. Plants position their leaves to absorb these photons. Energy from the photons is used to split water molecules inside the chloroplasts. In the process of splitting water, oxygen is released as a byproduct (thanks plants!). Splitting water also releases electrons and hydrogen ions.

These electrons and hydrogen ions are used to make energy in the form of ATP. Along with some electrons, ATP is then used in another cycle known as the Calvin cycle. The point of the Calvin cycle is to take in CO2 and use the energy created prior to reduce carbon molecules into chains of organic molecules. Most of the carbon in a plant comes from the intake of CO2. Through a series of steps (I will spare you the details) plants piece together carbon atoms into long chains. Some of these chains form glucose and some of that glucose gets linked together into cellulose.

Cellulose is the main structural component of plant cells. From the smallest plants in the world (genus Wolffia) all the way up to the largest and tallest redwoods and sequoias (incidentally some of the largest organisms to have ever existed on this planet) , all of them are built out of cellulose. So, in essence, all the plant life you see out there is literally built from the ground up by carbon originating from CO2 gas. Pretty incredible stuff, wouldn't you agree?

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 Power of Leaves

When we think of the dominance of flowering plants on the landscape, we usually invoke the evolution of flowers and seed characteristics like endosperm and fruit. However, evolutionary adaptations in the structure of the angiosperm leaf may have been one of the most critical factors in the massive diversification that elevated them to their dominant position on the landscape today. 

Leaves are the primary organs used in water and gas exchange. They are the centers of photosynthesis, allowing plants to take energy from our closest star and turn it into food. To optimize this system, plants must balance water loss with transpiration in order to maximize their energy gain. This requires a complex plumbing system that can deliver water where it needs to be. It makes sense that plant physiology should maximize vein production, however, there are tradeoffs in doing so. Veins are not only costly to construct, they also displace valuable photosynthetic machinery. 

It appears that this is something that flowering plants do quite well. Because leaves fossilize with magnificent detail, researchers are able to look back in time through 400 million years of leaf evolution. What they found is quite incredible. There appears to be a consistent pattern in the vein densities between flowering and non-flowering plants. The densities found in angiosperm leaves both past and present are orders of magnitude higher than all non-flowering plants. These high densities are unique to flowering plants alone. 

This innovation in leaf physiology allowed flowering plants to maintain transpiration and carbon assimilation rates that are three and four times higher than those of non-flowering plants. This gives them a competitive edge across a multitude of different environments. The evolution of such dense vein structure also had major ramifications on the environment. 

The massive change in transpiration rates among the angiosperm lineage is likely to have completely changed the way water moved through the environment. These effects would be most extreme in tropical regions. Today, transpiration from tropical forests account for 30-50% of precipitation. A lot of this has to do with patterns in the intertropical convergence zone, which ensures that such humid conditions can be maintained. However, in areas outside of this zone such as in the Amazon, a high abundance of flowering plants with their increased rates of transpiration enhances the amount of rainfall and thus forms a sort of positive feedback.

Because precipitation is the single greatest factor in maintaining plant diversity in these regions, increases in rainfall due to angiosperm transpiration effectively helps to maintain such diversity. As angiosperms rose to dominance, this effect would have propagated throughout the ecosystems of the world. Plants really are the ultimate ecosystem engineers. 

Photo Credit: Bourassamr (Wikimedia Commons)

Further Reading: [1]

Plants May Be Piping Light to Their Roots

Plants just might be piping more than just carbohydrates down to their roots. A study published in Science Signalling offers the first evidence that plants may actually be piping light down underground. No this isn't a metaphor either.

The presence of photoreceptors in the roots has been a bit of a puzzle ever since they were identified. A handful of hypotheses have been put forth in attempt to explain their function. It has been suggested that these photoreceptors are able to sense minuscule amounts of light penetrating through the soil. However, this research suggests there is another mechanism.

A team of researchers based out of South Korea found that certain stem tissues efficiently conducted wavelengths of red light down to the roots. Now before we get too ahead of ourselves, it should be noted that these are minuscule amounts of light. It certainly isn't enough for photosynthesis. However, it is light. Detectors placed under the soil at the ends of roots confirmed that light was indeed being transmitted.

Light is conducted through the tissues in much the same way as fiber optic cables. It is likely that the affinity for red wavelengths in particular has to do with the fact that it can travel farther than other, more intense wavelengths.

By experimenting with gene expression and light exposure, the team was able to demonstrate that light being piped to the roots activates a transcription factor involved in root growth and response to gravity. When the researchers blocked the ability to transmit light they found that root growth was severely stunted. Taken together, these results suggest that not only do roots receive information regarding light conditions above ground, they also directly perceive it.

It should be noted that all of this research was done on a single species, Arabidopsis thaliana. The question remains how common this phenomenon is throughout the plant kingdom. Most plants have photoreceptors in their roots, suggesting this light-piping ability is widespread.

Photo Credit: Dr John Runions/Science Photo Library

Further Reading: [1]