Insect Egg Killers

© Copyright Walter Baxter and licensed under CC BY-SA 2.0

Plants and herbivores are engaged in an evolutionary arms race hundreds of millions of years in the making. As plants evolve mechanisms to avoid being eaten, herbivores evolve means of overcoming those defenses. Our understanding of these dynamics is vast but largely focused on the actual act of an organism consuming plant tissues. However, there is growing evidence that plants can take action against herbivores before they are even born.

Taking out herbivores before they even have a chance to munch on a plant seems like a pretty effective means of defense. Indeed, for a growing number of plant species, this starts with the ability to detect insect eggs deposited on or in leaves and stems. As Griese and colleagues put it in their 2020 paper, “Every insect egg being detected and killed, is one less herbivorous larva or adult insect feeding on the plant in the near future.” Amazingly, such early detection and destruction has been found in a variety of plant lineages from conifers to monocots and eudicots.

Gumosis in cherries is a form of defense. Photo by Rosser1954/Public Domain

There are a few different ways plants go about destroying the eggs of herbivores. For instance, upon detecting eggs on their leaves, some mustards will begin to produce volatile compounds that attract parasitoid wasps that lay their eggs on or in the herbivore’s eggs. For other plants, killing herbivore eggs involves the production of special egg-killing compounds. Research on cherry trees (Prunus spp.) has shown that as cicadas push their ovipositor into a twig, the damage induces the production of a sticky gum that floods the egg chamber and prevents the eggs from hatching. Similarly, resin ducts full of insect-killing compounds within the rinds of mangoes will rupture when female flies insert their ovipositor, killing any eggs that are deposited within.

One of the coolest and, dare I say, most badass ways of taking out herbivore eggs can be seen in a variety of plants including mustards, beans, potatoes, and even relatives of the milkweeds and involves a bit of sacrifice on the plant end of things. Upon detecting moth or butterfly eggs, leaf cells situated directly beneath the eggs initiate a defense mechanism called the “hypersensitive response.” Though normally induced by pathogenic microbes, the hypersensitive response appears to work quite well at killing off any eggs that are laid.

“Leaves from B. nigra treated with egg wash of different butterfly species and controls inducing or not a HR-like necrosis. Pieris brassicae (P. b.), P. mannii, (P. m.), P. napi (P. n.), and P. rapae (P. r.) and Anthocharis cardamines (A. c.) induce a strong HR-like necrosis. Egg wash of G. rhamni (G. r.) and Colias sp. (C. sp.) induces a very faint response resembling a chlorosis and does not fit into the established scoring system (faintness indicates 1, but showing up on both sides of the leaf indicates 2).” [SOURCE]

Once eggs are detected, a signalling pathway within the leaf ramps up the production of highly reactive molecules called reactive oxygen species. These compounds effectively kill all of the cells upon which the butterfly eggs sit. The death of those plant cells is thought to change the microclimate directly around the eggs, causing them to either dry up or fall off. These forms of plant defense don’t stop once the eggs have been killed either. There is evidence to suggest that the hypersensitive response to insect eggs also induces these plants to begin producing even more anti-feeding compounds, thus protecting the plants from any herbivores that result from any eggs that weren’t killed.

Plants may be sessile but they are certainly not helpless. Defense mechanisms like these just go to show you how good plants can be at protecting themselves. Certainly, the closer we look at interactions like these, the more we will discover about the amazing world of plant defenses.

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

Further Reading: [1] [2]

A Tree That Makes Poisonous Rats

For many organisms, poisons are an effective means to keep from being eaten. However, making poisons can be costly. Depending on the compounds involved, poison synthesis can require a lot of nutrients that could be directed elsewhere. This is why some animals acquire poisons through their diet. Take, for instance, the monarch butterfly. As its caterpillars feed on milkweed, they sequester the milkweed toxins in their tissues, which makes them unpalatable into adulthood. Cases like this abound in the invertebrate world, but recently scientists have confirmed that at least one mammal has evolved a similar strategy.

Meet the African crested rat (Lophiomys imhausi). Its large size and bold color patterns make it look like the result of a passionate encounter between a porcupine and a skunk. However, it is 100% rat and it has a fascinating defense strategy that begins with a tree native throughout parts of eastern Africa aptly referred to as the poison arrow tree (Acokanthera schimperi).

An African crested rat displaying its crest of toxic hairs and aposematic color pattern. [SOURCE]

The poison arrow tree is a member of the milkweed family (Apocynaceae), and like many of its relatives, this species produces potent toxins that can cause heart failure. The toxic nature of this tree has not been lost on humans. In fact, the particular strain of toxin it produces is referred to as ouabaïne or “arrow poison” as indigenous peoples have been coating their arrows with its sap for millennia. It turns out that humans aren’t the only mammals to find use for this sap either. The African crested rat uses it too.

The African crested rat grows highly specialized crest of hairs along its back. These hairs are thick and porous and when the rat feels threatened, it erects the crest and shows off its stark black and white coloring. It has been noted in the past that predators such as dogs that try to eat the rat run the risk of collapsing into convulsions and dying so the idea was put forth that that crest of hairs was toxic. Only recently has this been confirmed.

By studying a group of these rodents, scientists observed an interesting behavior. Many of the rats in their study would chew and lick twigs and branches of the poison arrow tree and then chew and lick their crest. What this behavior does is transfer the plant toxins onto those specialized hairs. The high surface area of each hair means they can soak up a lot of the toxins. Surprisingly, the rats appear to be resistant to the sap’s toxic effects. Perhaps they possess modified sodium pumps in their heart muscles that counter the effects of the toxin. Or, they may possess a highly specialized gut flora that breaks down the toxins. Either way, the rats do not show any signs of poisoning from this behavior.

A close-up view of the African crested rat’s poison anointed hairs. Photo by Sara B. Weinstein

The rats don’t have to do this very often to remain poisonous. By talking with locals that still use the poison arrow tree sap on their arrows, researchers learned that the compounds are extremely stable. Once coated, arrows will remain toxic for years. As such, the African crested rat likely doesn’t need constant application for this defense mechanism to remain effective.

As far as we know, this is the first example of a mammal sequestering plant toxins as a form of defense. It is amazing to think that a defense strategy evolved by a plant to avoid being eaten can be co-opted by a rat so that it too can avoid being eaten. Sadly, it is feared that this unique relationship between rat and tree is starting to disappear. Though more research is needed to accurately assess their numbers, there is growing evidence that African crested rats are on the decline. Hopefully with a bit more attention, these trends can be properly assessed and conservation measures can be put into place. In the meantime, please avoid putting any and all rats in your mouth.

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

Further Reading: [1]

Why Plant Relationships Matter for Caterpillars

Photo by Judy Gallagher licensed under CC BY 2.0

When it comes to caterpillars, plant diversity matters. By studying nearly 30,000 plant-caterpillar interactions across three continents (Asia, North America, and Europe), scientists have uncovered important insights into lepidopteran biodiversity in temperate broadleaf forests.

Plants and the caterpillars they host are engaged in an evolutionary arms race. As plants evolve different defenses, caterpillars evolve new ways overcoming them. As you can imagine, studying these intricate relationships can be as fascinating as it is challenging. One could easily spend a lifetime trying to understand the relationships among only a handful of species. However, by taking a step back and asking bigger questions related to evolution and herbivory, scientists have found some interesting patterns than help describe the diversity of plant-caterpillar relationships.

As one might expect, they found that as plant diversity increases, so too does the diversity of caterpillars an ecosystem can support. Many caterpillars specialize on one or only a few different host plants and these are often (though not always) within the same plant family. The reason for this has to do with plant defenses. The more closely related plants are, the more likely they are to share similar defense strategies. For instance, most milkweeds (Asclepias spp.) produce toxic compounds called cardiac glycosides and many different members of the nightshade family (Solanaceae) produce similar suites of toxic alkaloids. As a result, insects that munch on their tissues have similar hurdles to overcome in an evolutionary sense.

The more closely related plants there are in an environment, the more likely it is that the caterpillars they host can jump from one plant species to another. As a result, ecosystems that boast relatively few plant lineages support relatively few caterpillar species in part because the caterpillars they do host can more easily jump from plant species to another. The same logic applies in the opposite direction as well. Ecosystems comprised of a diversity of plant lineages limit the likelihood that any given species of caterpillar can find multiple different hosts. Because each clade of plants produces their own brand of herbivore defenses, the caterpillars hosted by each are also more likely to be different. Thus, as plant diversity goes up, so too do the numbers of caterpillar species an ecosystem can support.

Though not tested by this research, this also provides yet another example of why invasive plants harm biodiversity. Plants from other areas of the world are more likely to present novel defenses to native herbivores. If the caterpillars do not have what it takes to overcome these defenses or simply don’t recognize the plant as food, the fewer caterpillars that ecosystem can support.

Of course, none of this should come as a surprise to those interesting in native plants and gardening. The more indigenous plants you grow in and around your landscape, the more insects you can support. I also firmly believe that the results of this research are not limited to caterpillars. The same pattern likely applies to any number of plant eaters, from microbes to mammals, no matter where you look. What this research gives us are some answers to questions like “why does biodiversity matter?”

Photo Credit: [1]

Further Reading: [1]

Corn Lilies, Cyclops Lambs, and Sonic the Hedgehog

Photo by Judy Gallagher licensed by cc-by-2.0

1957 was an alarming year for Idaho ranchers. Some herds of sheep were giving birth to lambs with severe deformities. The lambs simply weren’t developing right. They emerged from the womb sporting limbs from their heads, incomplete brains, and some of them had only a single, malformed eye in the middle of their face. It would take over a decade before the cause of these deformities was identified and another two decades before we knew why it happened. The first line of evidence came from the weather patterns during that fateful year.

In an average year, sheep usually find enough forage at lower elevations. With plenty of rain keeping plants happy and lush, the sheep don’t have to travel far to find food. Things change during severe droughts. As droughts worsen, plants at lower elevations start to disappear. To find enough food, sheep will move up in elevation where plants are not yet affected by drought. However, the move up slope coincides with a change in the presence and abundance of some plant species. Notably, species like the corn lily (Veratrum californicum) are more prevalent at higher elevations.

Photo by Clint Gardner licensed by CC BY-NC-SA 2.0

Now if there is one common thread that winds its way through the genus Veratrum, it’s the fact that all members produce some seriously potent alkaloid compounds. Though toxicity can vary from species to species, it is a safe bet that most Veratrum can harm you if ingested during their active growing period. However, despite the fact that all parts of Veratrum are toxic, it appears that these Idaho sheep were a bit desperate. It was discovered that during the drought of 1957, some sheep were feeding on the flowers of the V. californicum.

A deformed lamb showing the single, malformed eye and the anomalous limbs.

The flowers themselves aren’t the most toxic part of the plant but they produce measurable levels of toxic alkaloids. After 11 years of studying these malformed sheep, scientists realized that although pregnant sheep could feed on the flowers of V. californicum with no ill effects, they would go on to give birth to the deformed lambs. It became readily apparent that the deformities found in these lambs could be traced back to the consumption of V. californicum.

However, this was not case closed. The ranchers learned that they must keep their sheep away from Veratrum but no one had any idea as to how eating these plants led to such horrible birth defects. It took 25 more years before scientists had that answer.

While studying embryonic development in fruit flies, researchers discovered a set of genes that, when deactivated, cause the flies to grow spiny hairs all over their body. They named this gene “Sonic Hedgehog” after the spiky blue video game character. It turns out that the Sonic Hedgehog gene was extremely important in the development of more organisms than just flies. Importantly, these genes control the way in which the body plan of an organism develops. When something goes wrong with the Sonic Hedgehog pathway, a whole slew of deformities follow. Among these is the development of a single, malformed eye on the middle of the mammalian head.

Luckily, researchers studying Sonic Hedgehog remembered the story of the cyclops sheep in Idaho. It didn’t take long to put the puzzle pieces together. It was soon realized that V. californicum produces one alkaloid in particular that interferes with Sonic Hedgehog. The compound was given the name “Cyclopamine” as a reference to the deformities is caused in those sheep back in 1957. Scientists finally had the smoking gun.

The molecular structure of Cyclopamine

When droughts caused sheep to moved into the mountains in search of plants to munch, some of them would nibble on the flowers of V. californicum. If they were pregnant at the time, enough Cyclopamine made it into their system that it would shut down the Sonic Hedgehog gene pathway in their developing offspring. Once that pathway is shut down, the embryo no longer has a sound blueprint for development and all of those horrendous deformities take place.

The story does not end here. Not only was a 30+ year mystery solved, scientists had come away with a far more detailed understanding embryonic development. They also walked away with some new ideas to test. The most exciting of these involves cancer treatments. It turns out, the Sonic Hedgehog pathway is one of the many pathways involved in a couple different kinds of cancer. Normally, Sonic Hedgehog is dormant in adults but certain circumstances can see it reactivate and go into overdrive, leading to cancerous tumors. Some scientists are now using Cyclopamine to turn off the Sonic Hedgehog pathway in those tumors as a form of cancer treatment.

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

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

The Role of Leaf Shape on Insect Herbivory

Plants can defend themselves from herbivores in a variety of ways - thorns, spines, hairs, toxins, etc. - but have you ever considered the role of leaf shape in preventing herbivory? It’s okay if you haven’t because leaf shape rarely, if ever, makes it into conversations of plants defense. A recent experiment from Japan has changed that by demonstrating that leaf shape can actually deter a specialist leaf-rolling weevil.

Meet Apoderus praecellens, a leaf rolling weevil that specializes on a genus of mints called Isodon. To successfully reproduce, female leaf rolling weevils must roll up an Isodon leaf while laying eggs as she goes. The end result is a tiny cigar-shaped, edible nursery chamber in which her larvae will develop. The act of processing a leaf is a complex process.

Isodon trichocarpus Photo by Qwert1234 licensed by CC BY-SA 3.0

The female weevil begins by walking along the margin of the leaf until she reaches the apex. At that point she walks sideways towards the interior of the leaf until she finds the midrib. She then turns around and walks back toward the leaf base again. She repeats these steps several times on both sides of the leaf until she is satisfied. At that point, she will take several bites out of the midrib, which causes the leaf to wilt. The wilted leaf is then much easier to manipulate and thus the rolling process begins.

In the wild, female weevils are well documented on the leaves of I. trichocarpus but not on the leaves of I. umbrosus. This is strange because not only are these plants closely related, they frequently grow in close proximity to one another. Why would the female weevils prefer one over the other? The answer appears to lie in the shape of their leaves.

Isodon trichocarpus produces non-lobed leaves whereas the leaves of I. umbrosus are deeply lobed. When presented with a choice, female weevils did indeed choose to roll I. trichocarpus leaves over those of I. umbrosus. These plants do not differ in their chemical makeup and larvae raised on both species did not differ in their health or development time. Thus, nutritional value or defense compounds don’t explain weevil preference.

Even more amazing is that the preferences seemed to change when I. trichocarpus leaves were cut to resemble the lobed I. umbrosus leaves. It seems that the presence of leaf lobes is the key to whether a weevil decides to lay her eggs or not. The reason for this seems to be the complex leaf inspection behavior outlined above. The deep lobes of I. umbrosus leaves disrupt the female weevils as they carry out their complex inspection process. If the females are interrupted, they rarely progress to the leaf rolling stage.

The researchers are quick to point out that leaf shape in this instance probably didn’t evolve in response to herbivory. Leaf shape is the result of a multitude of selection pressures like light availability, heat, and drought. Still, the fact that leaf shape can also influence herbivore pressure is an interesting piece to add to the puzzle. It is a great reminder that an organism’s niche comprises so much more than simply the abiotic conditions in which it lives. The niche is also the myriad biological interactions each organism undertakes.

Photo Credits: [1] [2]

Further Reading: [1]

The Dual Benefits of Smelling Like Frightened Aphids

Photo by KENPEI licensed under the GNU Free Documentation License

If you garden, you have probably dealt with aphids. These tiny sap-suckers not only drain the plant of valuable sap, they can also serve as vectors for disease. Plants must contend with the ever-present threat of aphid infestation throughout the growing season and have evolved some amazing defenses against these insects. Recently an incredible form of defense against aphids has been described in pyrethrum (Tanacetum cinerariifolium) and it involves smelling like a frightened aphid colony.

Aphids produce their own alarm pheromones when attacked. Because aphids form large, clonal colonies, these pheromones can help warn their kin of impending doom. Other aphids will also eavesdrop on these alarm signals and will avoid settling in on plants where aphids are being attacked. Aphids aren’t the only ones honing in on these scents either. Aphid predators and parasitoids will also use these compounds to locate aphid colonies. As such, these pheromones are helpful to the host plant because it can mean a reduction in aphid numbers.

An alate (winged) green peach aphid (Myzus persicae).

The selection pressured imposed by aphids on plants is so strong that it appears that at least one species of pyrethrum has actually evolved a means of producing these pheromones themselves. Pyrethrum is a member of the aster family (Asteraceae) native to southern portions of Eurasia. Like all flowering plants, its flowers are the most precious organs. They are the key to getting their genes into the next generation and therefore protecting them from herbivore damage is of utmost importance.

It has been discovered that pyrethrums produce an aphid alarm pheromone called ( E )-β-farnesene or EβF for short. The pheromone is not produced in every tissue of the plant but rather it is concentrated near the inflorescence. What’s more, pheromone production is not constant throughout the duration of flowering. Researchers found that production reaches its peak just before the inflorescence opens to reveal the flowers within.

Photo by そらみみ licensed under CC BY-SA 4.0

The production of EβF in pyrethrum appears to serve a dual function. For starters, it actually results in reduced aphid infestation during the early stages of flowering. When the initial aphid attack begins, these insects consume some of the EβF as they feed and release it as they excrete honeydew. Other aphids detect EβF within the honeydew and will actually avoid the plant, likely due to the perception that the aphids feeding there are already under attack.

That does not mean that predators are not to be found. In fact, the other benefit of producing EβF in the inflorescence is that it appears to lure in one of the most voracious aphid predators on the planet - ladybird beetles. The ladybird beetles are able to detect EβF in the air and will come from far and wide to investigate in hopes of finding a tasty aphid meal. The ladybird beetles were most frequently found on plants during the early stages of floral development, which suggests that EβF production in the floral tissues is the main attractant.

A 7-spot ladybird beetle (Coccinella septempunctata). Photo by S. Rae licensed under CC BY 2.0

Interestingly, it has been found that constant production of EβF is less effective at deterring aphids than pulses of EβF. It is thought that just as humans can get used to certain background levels of scent, so too can aphids. If aphids are exposed to high levels of EβF for long periods of time, they simply recognize it as the safe background level and will continue to feed. This may explain why pyrethrum plants only produce EβF for a short period of time during the most crucial stages of floral development. Research like this not only improves our understanding of the myriad ways in which plants defend themselves, it also offers us new avenues for researching more natural ways of defending the plants we rely on from unwanted pests.

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

Further Reading: [1]

An Intruiguing Relationship Between Ants and Cacti

The extrafloral nectaries of Pachycereus gatesii appear as tiny red bumps just below the areole.

It’s hard to think of a group of plants that are better defended than cacti. Frequently and often elaborately adorned with vicious spines, these succulents make any animal think twice about trying to take a bite. And yet, for some cacti, spines don’t seem to cut it. A surprising amount of species appear to have taken their defense system to a whole new level by recruiting nature’s most tenacious bodyguards, ants.

Plants frequently have a friend in ants. Spend some time observing ants at work and it’s east to see why. These social insects have numbers and strength on their side. Give ants a reason to be invested in your survival and they will certainly see to it that nothing threatens this partnership. For cacti, this involves the secretion of nectar from specialized tissues called extrafloral nectaries.

Extrafloral nectaries are not unique to cacti. A multitude of plant species produce them, often for similar reasons. Ants love a sugary food source and the more reliable that source becomes, the more adamant an ant colony will be at defending it. The odd thing about cacti is that they don’t seem to have settled on a single type of extrafloral nectary to do the trick. In fact, as many as four different types of extrafloral nectaries have been described in the cactus family.

Ants visiting the extrafloral nectaries covering the developing flowers of Pilosocereus gounellei.

Some cacti secrete nectar from highly modified spines. A great example of this can be seen in genera such as Coryphantha, Cylindropuntia, Echinocactus, Ferocactus, Opuntia, Sclerocactus, and Thelocactus. Such spines are usually short and blunt, hardly resembling spines at all. Other cacti secrete nectar from regular looking spines. This adaptation is odd as there does not seem to be anything special about the anatomy of such spines. Examples of this can be seen in genera such as Brasiliopuntia, Calymmanthium, Harrisia, Opuntia, Pereskiopsis, and Quiabentia. Still others secrete nectar from highly reduced leaves that are found at the base of where the spines originate (the areole). Such leaves have been described in Acanthocereus, Leptocereus, Myrtillocactus, Pachycereus, and Stenocereus. They aren’t easy to recognize as leaves either. Most look like tiny scales. Finally, the fourth type of extrafloral nectary comes in the form of specialized regions of the stem tissue. This has been described in genera such as Armatocereus, Leptocereus, and Pachycereus.

Highly modified spines functioning as extrafloral nectaries in Ferocactus emoryi.

Seemingly normal spines of Harrisia pomanensis secreting nectar.

Regardless of where they form, their function remains much the same. They secrete a form of nectar which ants find irresistible. The more reliable this food source becomes, the more aggressive ant colonies will be in defending it. This is an especially useful form of defense when it comes to small insect herbivores. Whereas spines deter larger herbivores, they don’t do much to deter organisms that can just slip right through them unharmed. Ants also clean the cacti, potentially removing harmful microbes like fungi and bacteria. Though we are only just beginning to understand the depths of this cactus/ant mutualism, what we have discovered already suggests that the relationship between these types of organisms is far more complex than what I have just outlined above.

For instance, it may not just be sugar that the ants are looking for. In arid desert habitats, water may be the most limiting resource for an ant colony and large, succulent cacti are essentially giant water reservoirs. The key is getting to that water. One study that looked at a species of barrel cactus growing in Arizona called Ferocactus acanthodes found that as spring gives way to summer, the concentration of sugars secreted by the extrafloral nectaries decreases. As a result, the nectar becomes far more watery. Amazingly, ant densities on any given barrel cactus actually increased throughout the summer, despite the fact that the nectar was being watered down. Ants are notoriously prone to desiccation so it stands to reason that water, rather than sugar, is the real prize for colonies hanging out on cacti in such hot desert environments.

The incredible floral display of Ferocactus wislizeni, a species whose reproductive efforts are affected by the types of ants they attract. Photo by Joseph j7uy5 licensed under CC BY-NC-SA 2.0

Another interesting observation about the cactus/ant mutualism is that it appears that the identity of the ants truly matters. Though defense is the main benefit to the cactus, research suggests that there is a tipping point in how much such defenses benefit cacti. It has been found that although cacti initially benefit from anti-herbivore and cleaning services, extra aggressive ant species can actually drive off potential pollinators. At least one study has shown that when less aggressive ant species tend cacti, they produce more fruits and those fruits contain significantly more seeds than cacti that have been tended by extremely aggressive ant species. This is especially concerning when we think about the growing issue of invasive ants. As more and more non-native ant species displace native ants, this could really tip the balance for some cactus species.

Despite all of the interesting things we have learned about extrafloral nectaries in the family Cactaceae, there are so many questions yet to be answered. For starters, we still do not know how many different taxa produce them in one form or another. It is likely that closer inspection, especially of rare or poorly understood groups, will reveal that far more cacti produce some type of extrafloral nectary. Also, we know next to nothing about the anatomy of the different types of nectaries. How do they differ from one another and how do some, especially those derived from ordinary spines, actually function? Finally, do these nectaries function year round or is there some sort of seasonal pattern to their development and utility. How does this affect the types of ants they attract and how does that in turn affect the survival and reproduction of these cacti? For such a charismatic group of plants as cacti, we still have to much to learn.

Photo Credits: Thanks to Dr. Jim Mauseth and Dr. John Rebman and Dr. Silvia Rodriguez Machado for use of their photos [1] [2]

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

Why Are Some Plants Overcompensating?

Photo by CanyonlandsNPS licensed under public domain.

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). Photo by Dcrjsr licensed under CC BY 3.0

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). Photo by Joan Simon licensed under CC BY-SA 2.0

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.

Photo by Hectonichus licensed under CC BY-SA 3.0

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]

Life With Endophytic Fungi

Endophytic fungi living in the cells of a grass leaf. Photo by Nick Hill (Public Domain)

Talk about plants long enough and fungi eventually make their way into the conversation. These two walks of life are inextricably linked and probably have been since the earliest days. At this point we are well aware of beneficial fungal partners like mycorrhizae or pathogens like the cedar apple rust. Another type of relationship we are only starting to fully appreciate is that of plants and endophytic fungi living in their above ground tissues. 

Endophytic fungi have been discovered in many different types of plants, however, it is best studied in grasses. The closer we look at these symbiotic relationships, the more complex the picture becomes. There are many ways in which plants can benefit from the presence of these fungi in their tissues and it appears that some plants even stock their seeds with fungi, which appears to give their offspring a better chance at establishment. 

To start, the benefits to the fungi are rather straight forward. They get a relatively safe place to live within the tissues of a plant. They also gain access to all of the carbohydrates the plants produce via photosynthesis. This is not unlike what we see with mycorrhizae. But what about the plants? What could they gain from letting fungi live in and around their cells?

One amazing benefit endophytic fungi offer plants is protection. Fungi are famous for the chemical cocktails they produce and many of these can harm animals. Such benefits vary from plant to plant and fungi to fungi, however, the overall effect is largely the same. Herbivores feeding on plants like grasses that have been infected with endophytic fungi are deterred from doing so either because the fungi make the plant distasteful or downright toxic. It isn't just big herbivores that are deterred either. Evidence has shown that insects are also affected.

There is even some evidence to suggest that these anti-herbivore compounds might have influences farther up the food chain. It usually takes a lot of toxins to bring down a large herbivore, however, some of these toxins have the potential to build up in the tissues of some herbivores and therefore may influence their appeal to predators. Some have hypothesized that the endophytic fungal toxins may make herbivores more susceptible to predators. Perhaps the toxins make the herbivores less cautious or slow them down, making them more likely targets. Certainly more work is needed before anyone can say for sure.

Italian ryegrass (Lolium multiflorum) is one of the most studied grasses that host endophytic fungi. Photo by Matt Lavin licensed under CC BY-SA 2.0

Another amazing example deals with parasitoids like wasps that lay their eggs in other insects. Researchers found that female parasitoid wasps can discriminate between aphids that have been feeding on plants with endophytic fungi and those without endophytic fungi. Wasp larvae developed more slowly and had a shorter lifespan when raised in aphids that have fed on endophytic fungi plants. As such, the distribution of plants with endophytic symbionts may have serious ramifications for parasitoid abundance in any given habitat.

Another benefit these endophytic fungi offer plants is increased photosynthesis. Amazingly, some grasses appear to photosynthesize better with endophytic fungi living in their tissues than plants without fungi. There are many mechanisms by which this may work but to simplify the matter, it appears that by producing defense compounds, endophytic fungi allow the plant to redistribute their metabolic processes towards photosynthesis and growth. In return, the plants produce more carbohydrates that then feed the fungi living in their tissues. 

One of the most remarkable aspects about the relationship between endophytic fungi and plants is that the plants can pass these fungi on to their offspring. Fungi are able to infect the tissues of the host plants' seeds and therefore can be carried with the seeds wherever they go. As the seedlings grow, so do the fungi. Some evidence suggests this gives infected seedlings a leg up on the competition. Other studies have not found such pronounced effects.

Still other studies have shown that it may not be fungi in the seeds that make a big difference but rather the fungi present in the decaying tissues of plants growing around them. Endophytic fungi have been shown to produce allelopathic compounds that poison neighboring plants. Areas receiving lots of plant litter containing endophytic fungi produced fewer plants but these plants grew larger than areas without endophytic fungi litter. Perhaps this reduces competition in favor of plant species than can host said endophytes. Again, this has potentially huge ramifications for the diversity and abundance of plant species living in a given area.

We are only beginning to understand the role of endophytic fungi in the lives of plants and the communities they make up. To date, it would appear that endophytic fungi are potentially having huge impacts on ecosystems around the globe. It goes without saying that more research is needed.

Photo Credits: [1] [2]

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

Toxic Nectar

I was introduced to the concept of toxic nectar thanks to a species of shrub quite familiar to anyone who has spent time in the Appalachian Mountains. Locals will tell you to never place honeybee hives near a patch of rosebay (Rhododendron maximum) for fear of so-called "mad honey." Needless to say, the concept intrigued me.

A quick internet search revealed that this is not a new phenomenon either. Humans have known about toxic nectar for thousands of years. In fact, honey made from feeding bees on species like Rhododendron luteum and R. ponticum has been used more than once during times of war. Hives containing toxic honey would be placed along known routs of Roman soldiers and, after consuming the seemingly innocuous treat, the soldiers would collapse into a stupor only to be slaughtered by armies lying in wait.

Rhododendron luteum. Photo by Chrumps licensed under CC BY 3.0

The presence of toxic nectar seems quite confusing. The primary function of nectar is to serve as a reward for pollinators after all. Why on Earth would a plant pump potentially harmful substances into its flowers?

It is worth mentioning at this point that the Rhododendrons aren't alone. A multitude of plant species produce toxic nectar. The chemicals that make them toxic, though poorly understood, vary almost as much as the plants that make them. Although there have been repeated investigations into this phenomenon, the exact reason(s) remain elusive to this day. Still, research has drummed up some interesting data and many great hypotheses aimed at explaining the patterns.

Catalpa nectar has been shown to deter some ants and butterflies but not large bees. Photo by Le.Loup.Gris licensed under CC BY-SA 3.0

The earliest investigations into toxic nectar gave birth to the pollinator fidelity hypothesis. Researchers realized that meany bees appear to be less sensitive to alkaloids in nectar than are some Lepidopterans. This led to speculation that perhaps some plants pump toxic compounds into their nectar to deter inefficient pollinators, leading to more specialization among pollinating insects that can handle the toxins.

Another hypothesis is the nectar robber hypothesis. This hypothesis is quite similar to the pollinator fidelity hypothesis except that it extends to all organisms that could potentially rob nectar from a flower without providing any pollination services. As such, it is a matter of plant defense.

The nectar of Cyrilla racemiflora is thought to be toxic to some bees. Photo by Koala:Bear licensed under CC BY-SA 2.0

Others feel that toxic nectar may be less about pollinators or nectar robbers and more about microbial activity. Sugary nectar can be a breeding ground for microbes and it is possible that plants pump toxic compounds into their nectar to keep it "fresh." If this is the case, the antimicrobial benefits could outweigh the cost to pollinators that may be harmed or even deterred by the toxic compounds.

Finally, it could be that toxic nectar may have no benefit to the plant whatsoever. Perhaps toxic nectar is simply the result of selection for defense compounds elsewhere in the plant and therefore is expressed in the nectar as a result of pleiotropy. If this is the case then toxic nectar might not be under as strong selection pressures as is overall defense against herbivores. If so, the plants may not be able to control which compounds eventually end up in their nectar. Provided defense against herbivores outweighs any costs imposed by toxic nectar then plants may not have the ability to evolve away from such traits.

Where Spathodea campanulata is invasive, its nectar causes increased mortality in native bee hives. Photo by mauro halpern licensed under CC BY 2.0

So, where does the science land us with these hypotheses? Do the data support any of these theories? This is where things get cloudy. Despite plenty of interest, evidence in support of the various hypotheses is scant. Some experiments have shown that indeed, when given a choice, some bees prefer non-toxic to toxic nectar. Also, toxic nectar appears to dissuade some ants from visiting flowers, however, just as many experiments have demonstrated no discernible effect on bees or ants. What's more, at least one investigation found that the amount of toxic compounds within the nectar of certain species varies significantly from population to population. What this means for pollination is anyone's' guess.

It is worth noting that most of the pollination-related hypotheses about toxic nectar have been tested using honeybees. Because they are generalist pollinators, there could be something to be said about toxic nectar deterring generalist pollinators in favor of specialist pollinators. Still, these experiments have largely been done in regions where honeybees are not native and therefore do not represent natural conditions.

Simply put, it is still too early to say whether toxic nectar is adaptive or not. It could very well be that it does not impose enough of a negative effect on plant fitness to evolve away from. More work is certainly needed. So, if you are someone looking for an excellent thesis project, here is a great opportunity. In the mean time, do yourself a favor and don't eat any mad honey.

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

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

How Trees Fight Disease

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.

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

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. 

Photo by T.Voekler licensed under CC BY-SA 3.0

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]

Convergent Carnivores

Photo by Natalie McNear licensed under CC BY-NC 2.0

A carnivorous lifestyle has evolved independently in numerous plant lineages. Despite the similarities between genera like Nepenthes, Sarracenia, and Cepholotus they are not closely related. Researchers have wondered how the highly modified leaves of various carnivorous plant species evolved into the insect trapping and digesting organs that we see today. Thanks to a recent article published in Nature, it has been revealed that the mechanisms responsible for carnivory in plants are a case of convergent evolution.

This research all started with the Australian pitcher plant Cepholotus follicularis. More closely related to wood sorrels (Oxalis spp.) than either of the other two pitcher plant families, this species offers a unique window into the genetic controls on pitcher development. Cepholotus produces two different kinds of leaves - normal, photosynthetic leaves and the deadly pitcher leaves that have made it famous the world over.

By observing which genes are activated during the development of these different types of leaves, the research team was able to identify which alleles have been modified. In doing so, they were able to identify genes involved in producing the nectar that attracts their insect prey as well as the genes involved in producing the slippery waxy coating that keeps trapped insects from escaping. But they also found something even more interesting.

[SOURCE]

By examining the digestive fluids produced by Cepholotus as well as many other unrelated carnivorous plant species from around the world, researchers made a startling discovery. They found that the genes involved in synthesizing the deadly digestive cocktails among these disparate lineages have a similar evolutionary origin.

Although they are unrelated, the ability to digest insects seems to have its origins in defending plants against fungi. You have probably heard someone say that fungi are more similar to animals than they are plants. Well, the polymer that makes up the cell walls of fungi is the same polymer that makes up the exoskeleton of insects - chitin. By comparing the carnivorous plant genes to those of the model plant Arabidopsis, the team found that similar genes became active when plants were exposed to fungal pathogens.

It appears that carnivorous plants around the world have all converged on a system in which genes used to defend themselves against fungal infection have been co-opted to digest insect bodies. Taken together, these results show that the path to carnivory in plants is surprisingly narrow. Evolution doesn't always require the appearance of new alleles but rather a retooling of genes that are already in place. 

Photo Credits: [1] [2]

Further Reading: [1]

The Gas Plant

Photo by Jörg Hempel licensed under CC BY-SA 3.0 de

Meet the gas plant, Dictamnus albus. This lovely herbaceous species is native to southern Europe, north Africa, and Asia. The gas plant is a member of the citrus family, Rutaceae, and like many members of this group, it has very showy blossoms. Its affiliation with the citrus fruits on your counter isn't the only interesting thing about this species. As the common name might suggest, this plant does something quite strange. 

During the heat of summer, parts of the gas plant exude an oily substance that smells much like the fruits of its cousin, the lemon. These oils have been known to cause contact dermatitis not unlike the sap of giant hogweed. However, this is not the strangest aspect of the gas plants oily nature. One of the properties of these oils is that they are highly volatile. So volatile in fact that they can ignite. 

Another common name for this species is burning bush (though not the one of biblical lore). If air temperatures get high enough or if someone takes a match to this plant on a hot day, the oils covering its tissues will ignite in a flash. The oils burn off so quickly that it is of no consequence to the plant. It goes on growing like nothing ever happened. If you're like me then you have one burning question after reading this - why?!

Despite how incredible this phenomenon may seem, it doesn't appear that too many people have looked into its function. Research has identified a highly flammable organic compound within the oils called isoprene. In plants, isoprenes are thought to protect against heat stress. This is bolstered by the fact that the gas plant produces these compounds during the heat of summer. 

Another possibility is that spontaneous ignition of these compounds could create small wildfires that clear the surrounding area of competition. I have not seen any evidence suggesting this. Yet another possibility is that this is simply an unrelated side effect of oil production. Since the plant is not hurt by the quick burst of flames, it simply hasn't had any reason to evolve a less flammable alternative. Evolution is funny like that. 

Still don't believe what you are reading? Check out this video:

Photo Credit: Jörg Hempel (Wikimedia Commons)

Further Reading: [1]

Sand Armor

Photo by Franco Folini licensed under CC BY-SA 2.0

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

Photo by Wolfram Burner licensed under CC BY-NC 2.0

The Truth About Coffee

Photo by Ria Tan licensed under CC BY-NC-ND 2.0

Mmm mmm coffee. This wonderful elixir has taken over the world. Though individual tastes and preferences vary, there is no denying that most folks who turn to coffee enjoy its effects as a stimulant. Many an In Defense of Plants post has been written in a coffee-fueled frenzy. Even as I write this piece, I am taking breaks to sip on a warm mug of the stuff. Coffee has plenty of proponents as well as its fair share of nay sayers but the health effects don't really concern me much. Today I would rather talk with you about the shrubs that are behind all of this. 

The coffee we drink comes from a handful of shrubs in the genus Coffea. Native to parts of Africa, these shrubs are distant relatives of plants like buttonbush (Cephalanthus occidentalis) and the bedstraws (Galium sp.). The "beans" that we brew coffee from are not beans at all but rather a type of pit or stone found in the center of a bright red berry. Before they are roasted, the "beans" are actually green. Plants in this genus produce an alkaloid compound known as caffeine. Though it may seem strange, the purpose of caffeine is not to stimulate the human nervous system (though it is a wonderful side effect) but rather it is produced as a defense mechanism for the plant. Making this compound is a complex process that involves many metabolic steps within the tissues of the plant. There are certain factions out there who would like to argue that this is proof against evolution but, as always, evidence seems to be the downfall of their argument. 

Creationists will tell you that the adaptations we see throughout the living world are too complex to have happened by accident. In reality, there is a vast amount of evidence that disputes this. Caffeine is one such example. It has evolved independently multiple times in many different plant lineages. Looking at the genome of coffee, researchers at the University at Buffalo (my alma mater) found that the genes involved in the synthesis of caffeine did not arise all at once. Instead, the genes duplicated multiple times throughout the history of this genus with each duplication coding for another step in the process of producing the caffeine molecule. The interesting part is that each step of this evolutionary process produced a chemical that was itself useful to the plant. The precursor compounds are bitter and toxic to the kinds of animals that like to nibble on the plant. 

As it turns out, the benefits that the plants get from caffeine aren't restricted to defense either. Coffee, as well as other flowering plants such as citrus, produce small amounts of caffeine in their nectar. Researchers at Arizona State University found that bees were 3 times more likely to remember a flowers scent when there was caffeine in the nectar than if there wasn't. This serves a great benefit to the plant producing it because it means that its flowers are much more likely to get pollinated. As it turns out, humans aren't the only species that enjoys a good buzz from caffeine.

Before we get too excited over coffee, we must remember that is definitely has its downside. Worldwide, we humans drink roughly 2.25 billion cups of the stuff every day. In order to produce that much coffee, humans have turned somewhere around 11 million hectares of land into coffee plantations. This has come at an extreme cost to the environment. Also, being a tropical species, the types of habitat used to grow coffee were once lush, tropical rain forests. A majority of coffee consumed around the world is produced in monocultures. Where there once stood towering trees and a lush understory is now an open, chemically-laden field of coffee shrubs. There is hope, however, and it is rising in popularity. 

If you enjoy coffee as much as I do, you should certainly consider switching over to shade grown coffee. I have attached a fair amount of literature at the bottom of this post but the long story short of it is that growing coffee is much less harmful to the environment when it is grown in a forest rather than open plantations. The structural complexity of shade grown coffee farms allows a greater diversity of plant and animal species to coexist with one another. Species diversity and richness are significantly higher on shade grown farms than on open field plantations. 

So, there you have it. Coffee is as complex as it is interesting. We humans are simply lucky to have stumbled across a plant that interacts with our brain chemistry in wonderful ways. Certainly coffee has benefitted in the long run. 

Photo Credit: Ria Tan (http://bit.ly/1pFQD1J)

Further Reading:
http://www.sciencemag.org/content/345/6201/1181.full

https://asunews.asu.edu/20130307_beesandcaffeine

http://s.si.edu/1o6wOaj

http://www.sciencedaily.com/releases/2012/08/120807101357.htm

http://bit.ly/1S6dLVV

Shady Spines

Tephrocactus articulatus. Photo by Frank Vincentz licensed under CC BY-SA 3.0

Fondling cacti with your bare hands is often ill-advised. These spiny plants are icons of plant defense mechanisms. Cactus spines are actually modified leaves/bud scales. They develop from a bundle of cells called "primordia" that are nearly indistinguishable from leaf primordia. Unlike leaves, however, cactus spines are not made up of living tissue. The genes for leaf development are shut off in these cells and instead, genes for wood fibers are ramped up, creating the stiff structures many of us have had to pry out of our skin.

It is easy to assume that spines are simply there for defense. For a lot species they certainly do the trick. However, for many other species, spines serve another important purpose - they provide shade. This is exemplified by the fact that cacti growing in rainforests and cloudy highlands often have reduced or no spines at all.

For cacti living in the sun-baked regions of the world, sunburn is a serious issue to contend with. Full sunlight can damage sensitive photosynthetic machinery and while intense UV rays wreak havoc on the genome. As such, any adaptation that can shelter these sensitive tissues to some degree is advantageous.

Cephalocereus senilis. Photo by Drew Avery licensed under CC BY 2.0

Spines also buffer the cactus from huge temperature swings. Think of fuzzy or papery spines as a sort of blanket covering the cactus. These spines create a boundary between air immediately surrounding the cactus and the cold nighttime air of these arid climates. This insulation can come in handy as desert temperatures can drop quite low when the sun goes down.

Another benefit spines have is to catch and direct water to the base of the plant. Rain is often scarce in these habitats so when it does occur, a cactus needs to be ready. Water collects on the spines and then runs down to the base. They also act as dew catchers, causing water vapor to condense on their surfaces. In this way, cacti are able to take advantage of every last drop available.

Though they certainly offer some protection, many of these shade spines are too thin and flexible to deter a hungry herbivore. That is where secondary compounds come into play. It is no wonder why some cacti are extremely toxic to herbivores. Whether they are for shade, protection, or water harvesting, cacti spines have managed to capture our imagination and knowing a bit more about their function makes these plants even more impressive.

Photo Credit: [1] [2]

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