Plant Defenses – myriad strategies

Plants, as you might imagine, devote a great deal of energy to defending themselves from predators.  We humans have a natural bias towards animals, creatures that are like us in that they are mobile and respond to stimulus on a timescale similar to ours.  Plants operate, with a few notable exceptions, on a slower timescale, but this in no way should be taken to imply that they are any less interactive vis-a-vis their surroundings.

Plants react to light and dark, sense gravity, moisture, nutrients, and toxins; some can “feel” other organisms (Venus fly traps for example) or “hear” sounds (sensitive plants).  One thing all organisms must cope with is predators and competitors, and all organisms need defenses against these threats.  Plants are no different in their needs, but they are largely immobile, so some of their defenses tend to take a different form than they do in animals.  Surprisingly, their defenses are not as different from animal defenses as one might expect.  I would break plant defenses into three broad categories: chemical, physical, and co-optive.

Chemical defenses often involve toxins of one sort or another or pungent aromas.  Some of these we assiduously avoid, such as certain members of the Sumac (Anacardiaceae) family like poison oak (Toxicodendron diversilobum) and poison ivy (Toxicodendron radicans) due to the allergen urushiol found in the sap.  Other plants using chemical defenses we consume with great relish, many of our foods and spices, for example, derive their strong flavors from the defenses the plant manufactures to deter herbivorous predators.  Mints (Lamiaceae), rosemary, cinnamon, peppers, and onions are good example of common foods we consume that utilize strong chemical defenses.  Other chemical defenses we find recreational and/or medical uses for; ephedrine from plants in the Ephedra family, THC from Cannibus, and cocaine refined from alkaloids found in the coca family (Erythroxylaceae) all have enormous economies reliant on them.

Coca cultivation in Bolivia near Coroico

Coca cultivation in Bolivia near Coroico

Chemical defenses are enormously interesting and extremely sophisticated, but they are largely hidden from us until we are affected by them.  This is part of the reason why eating unfamiliar plants is so dangerous, there are few good ways to determine if a plant is edible upon first encounter.

Physical defenses are the most obvious to us, especially when they come in the form of thorns and barbs, but those defenses barely scratch the surface of the types of physical defenses plants can employ.

An impressive but unsubtle defense - Ceiba speciosa in the Bolivian Amazon

An impressive but unsubtle defense – Ceiba speciosa in the Bolivian Amazon

Sharp pokey bits may defend plants against larger herbivores and chemicals help to protect them from insects or pathogens, but other plants themselves can be, if not predatory, at least detrimental to large trees.  Lianas and other climbing plants, epiphytes, parasitic plants, and even other large trees may need to be defended against.

Strangler fig (Ficus spp.) overwhelming a palm tree's defenses - Bolivia

Strangler fig (Ficus spp.) overwhelming a palm tree’s defenses – Bolivia

The photo above I find particular interesting as the palm tree being overwhelmed by the strangler fig usually has an effective counter to this sort of attack.  Palm trees and tree ferns both allow their old fronds to droop as they age, sheathing the trunk and providing a structure for climbing plants to adhere to.  Eventually these canny plants shed their dead fronds, and with them the uninvited plant guests that have taken up residence.  Many trees employ a similar strategy, eucalyptus and madrone have smooth bark that regularly sloughs off in strips.  The combination of smoothness and shedding makes it difficult for other plants to gain purchase.

Strangler fig is a generic term for a wide variety of tropical fig trees sharing a similar life strategy.  These are the “matapalo” or killer trees.  Rather than growing from the ground and climbing up these trees co-opt animals to carry their tiny seeds through the canopy.  A small portion of these seeds wind up in a place like the crotch of a branch or a broken limb where organic material has built up.  The young fig sends dangling roots down from the canopy in search of nutrients, eventually reaching the ground and transitioning from a vine-like life style to a more tree-like life style.  More and more ground-seeking tendrils make their way downwards, eventually ringing the host tree and strangling it.  As this happens the strangler fig uses the original host as a scaffold and sends its own canopy high enough to overshadow the unfortunate host.  The palm tree in the photo above was underneath a tree the strangler fig took root in and had the misfortune to be attacked from above rather than from below.

Color is an oft overlooked plant defense, the role of which is still being debated.  I don’t mean fruit color, that is blatant advertising and animal bribery.  The color and pattern of the leaves and trunk of plants may serve as defense against predators.

The most familiar example of this is variegation in leaves, that is the white or colored mottling seen most often in ornamental plants, but also occasionally found in the wild.

Variegated hibiscus leaf. Source

Color mottling in leaves is often a symptom of nutrient deficiency, insect predation, viral infection, or genetic chimerism (expression of more than one genetic sequence in a single organism).  In most of the above cases this indicates poor health in the plant, and a plant in poor health makes for an unappetizing meal.  Some plants seem to capitalize on this and mimic the effects of various types of poor health (eg. false leaf damage and variegation) to trick predators into avoiding what appears to a be an un-nutritious meal(1)(2).  Bark color, whether natural or as a result of mutualistic lichen growth may be a predator deterrent as well, as lighter colors may make predators more visible to other predators higher up on the food chain.

Before moving on to animal co-option I should mention one other strategy employed by some plants.  Outgrow your predators.  In this case a plant allocates few resources to defense and focuses on growth and/or reproduction.  Balsa trees follow this strategy, they grow astoundingly rapidly and produce copious numbers of seeds.  They are not long lived and have few toxins, as a result they are subject to immense amounts of predation from a wide range of species.  Some of these, such as tapirs, they avoid by growing out of their reach.  Others are more problematic.  I saw a 30 foot tall young balsa tree completely stripped of leaves by leaf cutter ants in less than two days.

Basla saplings - Bolivia

Basla saplings – Bolivia

The most interesting of the plant defenses, to my mind, is animal co-option for defense.  Ants are probably the animal most often co-opted by plants.  We don’t often think of plants as being the ones to manipulate animals, but that is more a reflection of our animal bias than of the true nature of things.  Plants are highly manipulative, in their slow manner.  Like many effective manipulators, they accomplish their ends via bribes (and in a few cases by outright lies – orchids tricking bees into trying to mate with the flowers is a classic example of vegetative duplicity).  Ants are employed as guards by a great number of plant species.

When I first arrived in the Amazon I recall thinking to myself, “Cool, I hope I get to see some of the ant/plant mutualism.”  The first plant I looked at closely was a common understory shrub in the widespread and diverse Melastomataceae family.

Melastomataceae with ant sheltering nodes at the base of the leaves - Peru

Melastomataceae with ant sheltering nodes at the base of the leaves – Peru

At the base of each leaf there was a hollow, swollen node with two small openings on the underside.  Tiny ants occupied these nodes and would rush out to defend the plant when it was bumped.  This is a surprisingly effective defense against herbivores of all sorts, insect and mammalian.  All through the Amazon (and elsewhere in the world) ants and plants have banded together.  I found arboreal ferns with hollow rootballs harboring and colonies, tall cecropia and smoke trees (“palo diablo” – devil trees) with hollow trunks harboring vast numbers of fierce and painful fire ants, and evocative single species plant stands called Chullachaqui gardens.

According to legend the Chullachaqui is a forest guardian spirit that keeps small monoculture garden plots scattered here and there in the forest.  One should be wary entering these areas, ask permission first, and be careful not to damage any of the plants the Chullachaqui grows.  This is good advice as the Chullachaqui gardens are home to a species of ant that lives underground and kills all the plants growing on the surface except for one species.  Animals that interlope are attacked also attacked and the bite and sting of the ant is painful as it contains formic acid.

One of the neatest ant-plant interactions comes in the form of hanging “ant gardens”.

Ant garden in Peru with Monstera spp.

Ant garden in Peru with Monstera spp.

The dense cluster of plants in the above photo is growing from an arboreal ant hive.  Certain plants produce seeds with fatty nodules on them that ants eat.  Ants collect the seeds and store them in their hives, clipping the edible portions off as needed.  Some of the sees sprout and send their roots into the rich material of the ant hive, reinforcing it and protecting it from rain and predators as they grow.  These hanging ant gardens are found throughout the tropics.

Ant-plant interactions are not limited to the tropics.  Some trillium species bribe ants with food to carry their seeds to good growing locations and elderberries in certain portions of North America keep a protective coterie of ants nearby by bribing them with sugar produces from nectaries grown specifically for the ants.

Co-opting an animal may well be the most sophisticated of plant strategies for its subtly, specificity, and efficiency.  Energy cost is at the root of all these strategies.  A plant only has as much energy as it can collect from the sun and soil nutrients.  It must balance its energy use amongst growth, reproduction, and defense.  Every defense a plant employs lessens the amount of energy it can devote to growth and reproduction.  Combining forces with other species can provide a relatively low-cost way for a plant to gain an aggressive, mobile, multi-pronged defense force.  Ants, for example defend their plant hosts with both physically damaging attacks and with chemicals.  That’s a two-for-one defense with an added bonus of rapid deployment for the relatively minimal investment of a home or some food.

Humans may well be mammal plants have trained best.

***

A note about the images and film vs digital in the field:

These photos (with the exception of the hibiscus leaf which is not mine) were taken in 2005 with a well used and abused Canon AE1 with a 50mm 1.4 lens and scanned from the negatives by the developing kiosks in Peru and Bolivia.  The quality of images reflects both the environmental stresses put on the camera and film and the irregular scanning quality.

For long periods of time in the field I still think that film is the better option.  I was in the jungle for months at a time, sometimes in places with no power (20+ days hiking and on a raft in Bolivia for example) and in hot, humid conditions with frequent thundering downpours and rampant mildew growth.  I love the digital camera I use now, but it would have been completely unusable for the majority of the time I spent in South America.

Film cameras do still have their place.

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Segovia: sandstones and granite

I have always loved things made of stone, especially ancient constructions.  The stone-masonry I have done has only increased my respect for the strength, vision, and talent of past masons.

Vermont garden wall

Small garden wall of Panton Shale for a friend in Vermont

Most of my stone projects have been small in scale.  The largest project was a 180 foot long retaining wall standing between 2 and 6 feet high, using 30 or 40 tons of stone.  That seems large when you’re doing it by yourself, but that’s a tiny project, barely larger than the little garden wall in the photo above.

In Peru there were some truly astounding pieces of megalithic engineering, many of them little known like Lanche and Kuelap, others well known like Saqsaywaman.

Walls at Saqsaywaman.  For scale zoom into the center of the full-size image to see the person.

Walls at Saqsaywaman. For scale zoom into the center of the full-size image to see the person.

Two days ago I went to the small Spanish city of Segovia and got to see several astounding pieces of stone-based architecture.  The first of these is the ancient Roman aqueduct.

The aqueduct in Segovia

The aqueduct in Segovia

The aqueduct runs about 15 km from the mountains into Segovia, with a 683 meter long raised section running through town.  The tall double arch of granite blocks is impressive enough by modern standards, even more so when you consider that it was built in the 1st or 2nd century, that the granite had to be carried in from the mountains, and that it is a dry-laid structure (no morter holding the blocks together) that has been standing for 1800 or 1900 years.  Clearly, this is a place with few earthquakes.

Granite is a favorite building material for many people.  It is an igneous rock that bubbles up in volcanic flows and cools in place.  The size of the crystals in the rock give an estimation of how long it took for the rock to cool and how much water there was in the melt.  The colors tell of the mineral content.  This granite is pale, with moderately large crystals weathering out, leaving the exposed stone extremely rough to the touch.

Due to the way it forms granite has no preferential cleavage plane, meaning that, given the right tools, it is easy to shape into whatever form is needed.  It is a dense and strong rock as well, another reason it is often used as a foundational material.

The blocks of stone making up the aqueduct are large, not enormous, but large, hundreds of pounds each.  At its highest point the aqueduct is 29 meters tall (that’s about as tall as a 4 or 5 story building).  Nearly 2 thousand years ago those blocks had to be hoisted up and set in place.  Clues as to how the Romans did so are carved into the blocks.

Lifting divots on the granite blocks

Lifting divots on the granite blocks

Each block was lifted into place with a pair of metal pincers, like those people used to carry ice-blocks with.  Divots were carved into the stone to prevent the pincers from losing their grip.  Presumably the divots were carved at the balance point of the block as well, a calculation I would be very curious to know how was done.

Supposedly Segovia was a “small outpost” when the Romans ran things in the area, though the effort and cost of building the aqueduct makes me question that assessment.  Small outpost or no, very little happened in the area for a long while, then in the 1200s the town began to grow and with that growth came the buildings that Europe is so well known for.

Castles and Cathedrals.  Segovia has impressive examples of both, the castle being the inspiration for Walt Disney’s version of Sleeping Beauty, and the cathedral being on the of the last of built of the great Gothic cathedrals.

Segovia cathedral

Segovia cathedral

Construction of the cathedral began in the 1500s, but took more than a century to complete.  The massive building looms over the city, glowing golden in the sunlight.

The first thing that struck me was neither the size nor the the tremendous amount of fine detail.  It was the color.  A warm, yellow/orange, not the color one associates with Gothic architecture, or with goths in general.  The castle, cathedral, and much of the rest of Segovia is made from this stone, not from the granite the aqueduct is made from.

The town of Segovia rests upon an outcrop of calcareous sandstone (sandstone with the grains cemented together by calcium rather than silica) and the land around rises and falls, exposing the bedrock in numerous small cliffs.  Sandstone is a sedimentary rock, a class of rock at the opposite end of the formation spectrum as granites and other igneous rocks.

Sandstone tends to have horizontal cleavage planes, refection the initial depositional patterns, and is often soft and easy to carve.  The sandstone in Segovia seems made for carving and the cathedral  builders took full advantage of this.

Cathedral detail carved from sandstone

Cathedral detail carved from sandstone

Sandstone weathers and erodes easily, especially in the presence of water.  Segovia, despite being a dry region by my standards (about a half meter of rain per year) is considered a wet place in comparison with nearby areas.  As such the builders took pains to protect the soft sandstone, making their waterspout gargoyles of the more resistant granite.

Cathedral gargoyle rain-spout

Cathedral gargoyle rain-spout

Statues of sandstone have not weathered as well as those of granite.

A royal lion slowly weathering away

A royal lion slowly weathering away

The the level of fine detail in the cathedral architecture is reflected elsewhere in the town.  The older buildings and the castle are covered with patterned façades.  In the past these patterns seem to have indicated which family owned the building and in a few cases older patterns could be seen under the more recent ones.

Old wall pattern, the material looks and feels like reconstituted sandstone.

Old wall pattern, the material looks and feels like reconstituted sandstone.

The castle, the Alcázar de Segovia, has a more simple pattern, but each intersection is studded with fragments of volcanic rock.

Looking up the castle wall to the battlements.  the small black studs are fig sized pieces of vesicular volcanic rock brought in from far away.

Looking up the castle wall to the battlements. the small black studs are fig sized pieces of vesicular volcanic rock brought in from far away.

Like many European castles the one at Segovia has gone through a number of iterations; fort, castle, palace, prison, artillery college, and museum.  It still serves the latter two roles.

The castle commands a wonderful view of the countryside in all directions.  One of the most magnificent views is of the cathedral:

Segovia cathedral from atop the Alcázar de Segovia battlements

Segovia cathedral from atop the Alcázar de Segovia battlements

In the opposite direction an old Templar keep and the sandstone cliffs much of the stone was quarried from to make the city are visible.

Templar keep and sandstone cliffs above the river below the castle

Templar keep and sandstone cliffs above the river below the castle

This has been a less science based post than most, but the trip to Segovia was far too interesting to keep all to myself.

The castle, aqueduct, and cathedral are the largest of the attractions, but not the only ones by far.  The food is delicious, mockingbirds flit about the city, interesting small plants grow from the old walls and on the red tile roofs, and great architecture abounds.

Small church in Segovia

Small church in Segovia

Club-Mosses on the mountain

A few days ago I had the opportunity to be a guest speaker on an alpine botany field trip for a class a friend of mine is teaching.  The highest and largest alpine environment in Vermont is atop Mt. Mansfield, two hundred acres of exposed rock, lichens, and a delicate assortment of tiny plants bordered by dense krummholz forest housing several rare bird species.

Mt Mansfield ridge trail

This areas is one of only three tiny regions of Vermont where alpine tundra environments exist, and part of a very small handful of places on the East Coast.  These places are relicts from the end of the last ice age, extremely sensitive to changes in temperature and moisture, home to plants that are usually found much further north.  The growing season is short, nutrients are in short supply, and wind stresses are high, all of which result in slow growing, long lived plants that do not colonize open areas well.  Visitors are encouraged to walk only on the rocky areas, keeping off of the easily damaged vegetation.

I had been eager to visit the peak of Mt. Mansfield for some time because it is one of the only places in Vermont that a certain small clubmoss lives.  I mentioned this to the botany students and during a rest break one of them got my attention and asked if the little plant he was pointing to was the one I had mentioned.

Appalachian Fir-Clubmoss (Huperzia appalachiana)

It was one of the smallest examples of Appalachian Fir-Clubmoss (Huperzia appalachiana) that I had seen anywhere, but it was indeed the plant I was looking for.

Clubmosses are really cool and predate flowering plants by an embarrassingly large span of time.  They are not really moss of any type, though they bear a superficial resemblance to the true mosses.  Mosses themselves are not true plants, having no vascular tissue, the plant equivalent of our circulatory system.  Mosses rely on diffusion to distribute water and nutrients and this imposes strict limits on their size.  Clubmosses are more akin to ferns and conifers: they have simple hair-like roots (true mosses have no roots), they have vascular tissue, and, at one point in the extremely distant past (300+ million years ago), their close cousins were the dominant large vegetation reaching one hundred feet above the ground.  Now most clubmosses are small, only a few inches tall, although in the Amazon I did encounter one waist high clubmoss near an overgrown pond.

Unknown Peruvian clubmoss. It grew to just above my waist.

I was interested in the Appalachian Fir-Clubmoss, Huperzia appalachiana, because several years ago I spent a summer in Shenandoah National Park, Virginia, climbing about on steep cliffs looking for this plant and trying to figure out how to measure any change populations might experience as the climate changes.  It likes acidic, well drained soils over igneous (or highly metamorphic) bedrock that receive frequent moisture, and, unusual for a clubmoss, direct sunlight.  It hybridizes easily with several other clubmosses, Shining Clubmoss (Huperzia lucidula), Northern Fir-Clubmoss (Huperzia selago), and Huperzia appressa, which some people do not distinguish from Huperzia appalachiana, making the identification question particularly vexing where the ranges overlap.

The Huperzia genus was recently split from the Lycopodium genus, which is where many of the more familiar clubmosses reside.  Like many of the Huperzia, the Appalachian Fir-Clubmoss grows from a dense basal cluster and, unlike many of the Lycopodium, it does not creep about over the ground.

Huperzia appalachiana – note the bands of white spore capsules

No-one is certain how long Appalachian Fir-Clubmoss lives.  The best answer I could get from a sharp fellow at Miami University in Ohio was, “At least seventeen years.”  Not a very satisfying answer, and he knew it.

One way to estimate the age is to count the bands of spore capsules on the stalk, those little white bits that looks like tiny eggs or pale ticks in the image above.  Each band correlates to roughly one year of growth.  Unfortunately, no-one knows how old the plant has to be before it starts producing those, and they don’t always produce them each year.  With some Huperzia species you can count the rings of gemmae, odd little cup-shaped brackets the plant produces that contain a tiny asexually produced plant that is dropped onto the ground in place of a spore when conditions are good.  The gemmae look very different from the microphylls, which is what clubmoss leaves are called.

Huperzia appalachiana
The gemmae are the little 3-part flanges near the top of the plant – further down the empty brackets are visible

Another way to judge the age is to count the bands of vegetation where the microphylls are pressed up close to the stem and where they spread out.  Each spreading ring indicates spring growth.

All that is good in theory, unfortunately Appalachian Fir-Clubmoss produces gemmae in a haphazard fashion and, unlike the photo above, often does not have those nice alternating bands of growth.  Hence the, “At least seventeen years,” answer to my question.

Clubmosses grow in a variety of forms and have been used for some rather unlikely purposes in the past.  The spores they produce are tiny and highly flammable, so much so that they were used as flash powder in old time photography.  Condoms were dusted with clubmoss spores to keep the rubber from sticking to itself, and diapers are sometimes dusted with the spores to prevent rashes.  Today, we mainly use living clubmosses in garlands and ancient clubmosses in our coal burning power-plants.

One of the great things about living in New England is the wonderful variety of local clubmosses.  They are delightfully archaic.  Deceptively so, considering that they have been living for well over 300 million years and are still common in many places world-wide.

Tree Ground-Pine (Lycopodium dendroideum)