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Missing Carpels & the Building Blocks of Science

(Photo by: Domingos Cardoso)

Common Name: Amarelão (Brazil), Grapia (Argentina), Khare Khara (Bolivia)

A.K.A.: Apuleia leiocarpa

Vital Stats:

  • Once considered a genus of three different species, now collapsed down to one by taxonomists
  • The only trimerous (three-parted) flower in the whole legume family
  • Male flowers grow an extra stamen in place of the missing carpel

Found: In the rainforests of central South America

It Does What?!

Nothing quite as bizarre as our usual subjects, actually, but stick with me here. This week, I’m attending the annual conference of the American Society of Plant Taxonomists in Columbus, Ohio. I’ll be giving a talk on some of my research dealing with Apuleia and the development of its flowers. I thought I’d take this week to share some of that research here, and to try to make it interesting for people who aren’t into obsessing over obscure plants. If you still find this entry painfully tedious, though, rest assured, we’ll be back to freaks and oddities next week.

Apuleia leiocarpa is part of the legume family, which, if you’re from a temperate part of the world, brings to mind little annuals like beans and peas and clover. In the tropics, though, legumes are just as often towering trees of the rainforest canopy (like Apuleia) or scraggy shrubs of arid grasslands (such as Acacia). Most of the nearly 20,000 species of legumes have flowers with the same basic groundplan: 5 sepals, 5 petals, 10 stamens (the male organs), and a single carpel (where the fruit and seeds form). There are closely related chunks of the family, though, in which some of these floral organs have been lost over the course of evolution. (Now, ‘lost’ can mean two things; either the organ starts to grow and is suppressed before it finishes developing, or it just never forms at all. To the naked eye, these two kinds of loss look exactly the same. I’ll come back to this later.) Apuleia is one such legume- it has entirely lost two of its sepals, two of its petals, and most of its stamens, making for a very simplified flower.

The Hermaphodite (‘Normal’) Flower
S=sepal, P=petal, A=anther/stamen, C=carpel, St=stigma

What’s more, it now forms two different types of flower (called ‘morphs’). If you were to look closely at a flowering branch on one of these trees, you would see that the vast majority of the flowers were male-only, having no carpel with which to form fruit. Only every fourth or fifth flower would be the hermaphrodite type that we think of as a ‘normal’ flower. Botanists refer to this type of plant as being andromonoecious (pronounced “an-dro-mon-ee-shus”). So why would a tree evolve to become andromonoecious? There are a couple of different theories, based on two different ways that the male-only flowers can be produced.

In the first and most common type of andromonoecy, all the flowers on the plant begin as normal hermaphrodites. There are flowers of all different ages, so while some are beginning to open, others haven’t finished forming yet. Pollination starts on the earlier flowers, and the plant detects that it has far more ovaries (future seeds) than it’s going to need. Maybe the soil isn’t providing enough nutrition to produce all those potential fruits, or maybe there’s a drought in progress. So, according to its needs, the tree simply suppresses the development of the carpels in the younger flowers before they have time to mature, leaving parts of each branch with hermaphrodite flowers and parts with male flowers.

The Male-Only Flower
S=sepal, P=petal (both removed)
Arrow= where the carpel would have been

In scenario two, some flowers never develop carpels; they are male-only from the time they are first formed. This type of andromonoecy is thought to occur because the tree requires large amounts of pollen to reproduce successfully (perhaps the species is wind pollinated and individuals tend to be far apart, for example), and it’s “cheaper” to produce male flowers than hermaphrodites. In this situation, we don’t see the pattern of younger versus older flowers with respect to which ones are male.

That white asterisk in the very middle shows the hole through which the carpel would have emerged. It’s just a small, empty cavity in the male flower.

So which type of andromonoecy does Apuleia have? In order to find out, a colleague and I studied pressed herbarium specimens as well as flowers preserved in alcohol. The flowers, we dissected and viewed under an incredibly powerful microscope called a scanning electron microscope, which allowed us to see minute details, such as where a suppressed carpel might have been. In the end, we found that male Apuleia flowers show no sight of having ever developed a carpel. We also noticed that the hermaphrodite flowers always occurred symmetrically, right in the centre of a group of male flowers, a pattern that we wouldn’t see if the andromonoecy was environmentally influenced.

So in the end, we’re able to say that in this species, the different floral morphs probably arose in evolution due to an increased need for pollen, rather than as a control on fruit production. Groundbreaking… right? Well, maybe not, but obscure little discoveries like this are the building blocks for the big important breakthroughs we read about in the news. If you want to make something huge, you need a good foundation to start from.

Now imagine spending three hours of your life staring at this.
Science is so glamourous.

Says Who?

  • Beavon & Chapman (2011) Plant Systematics and Evolution 296: 217-224
  • de Sousa et al. (2010) Kew Bulletin 65: 225-232
  • Gibbs et al. (1999) Plant Biology 1: 665-669
  • Spalik (1991) Biological Journal of the Linnean Society 42: 325-336
  • Zimmerman et al. (In Press) International Journal of Plant Sciences
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EVOLUTION TAG TEAM, Part 2: Sex & the Synconium

The second in an ongoing series of biology’s greatest duos. (Check out Parts One and Three)

(Via: Mastering Horticulture)

Common Name (Plants): Fig Trees

  • A.K.A.: Genus Ficus

Common Name (Wasps): Fig Wasps

  • A.K.A.: Family Agaonidae

Vital Stats:

  • Approximately 800 species of figs
  • Most are trees, but some are shrubs and vines
  • Approximately 640 species (20 genera) of fig wasps
  • All are obligate pollinators of figs

Found: Throughout the Tropics

It Does What?!

Snacked on any Fig Newtons lately? Tasty, right? Like the ad says, “A cookie is just a cookie, but a Newton is fruit and cake.”  …And wasps.

They must have run out of space on the package for that last part.

Before you toss out your favourite teatime treat, I should point out that without those wasps, the figs themselves wouldn’t exist. [Personally, I love Fig Newtons and will eat them regardless of any insects present.] This plant-insect pairing actually represents one of the most stable symbioses out there, with evidence suggesting it has existed for over 65 million years.

Now with 10% more Wings
(Via: Wikipedia)

While it’s not entirely clear how this arrangement evolved in the first place, fig trees produce a unique structure called a synconium, in which the flowers are actually inside the part we think of as the fruit. This synconium, which can contain up to 7000 flowers, depending on the fig species, has a tiny hole at the tip called an ostiole. In order for the flowers to be pollinated and the fruit to grow, a female wasp must squeeze through that hole, often losing her wings and antennae in the process, and distribute pollen that she carries in a sac on her abdomen. As she does so, she also uses her ovipositor to reach down into some of the female flowers and lay her eggs in their ovaries, where a gall is formed and the larvae can develop. Then she dies and ends up in a cookie. The End.

But hold on, let’s remove humans from the equation for a moment. She dies, but her eggs hatch into little moth larvae which use the growing fig for nutrition. Once they’re old enough, the young wasps mate with one another inside the fig (another nice mental image for snacktime), and the females gather pollen from the male flowers and store it inside their abdominal pollen baskets (yes, that’s actually what they’re called). The wingless male wasps have a simple, three step life: 1) mate with females, 2) chew a hole through the fig so they can leave, 3) die. That’s pretty much it for them. They may escape the nursery with the females, but they’ll die shortly thereafter, regardless. In fact, even the females have a pretty rough deal; from the time they’re old enough to mate, they have about forty-eight hours to get their eggs fertilized, gather pollen, find a new synconium, distribute the pollen, and lay their eggs. Two days, and their life is over. No pursuit of happiness for the fig wasp, I’m afraid.

“What does it all mean?”
(Via: BugGuide.net)

As with any long-standing mutualism, there are, of course, parasites ready and waiting to take advantage of it. These parasites are wasps which are able to enter the synconium and lay their eggs, but which do not pollinate the fig. Although their eggs will crowd out those of the fig wasps, decreasing the number of fig wasp larvae born, they are kept in check by the fact that any unpollinated synconium will be aborted by the tree and drop to the ground, taking the parasite eggs with it. The nonpollinating wasps are therefore kept from being a serious threat to the tree’s pollinators.

So there you have it, another of evolution’s great matches. The wasps get an edible nursery, the trees get pollinated, and we get tasty fruits with suspicious crunchy bits that probably aren’t dead wasp bodies, so just try not to think about it too much…

Seeds, or wasp eggs? You be the judge!
(Via: This Site)

[Fun Fact: The symbiosis between fig species and their corresponding wasp partners is so specific (often 1:1), that the shape of the ostiole actually matches the shape of the head of the wasp species which will pollinate it.]

[For those who would like to read about figs and fig wasps in much greater detail (such as how this works when the male and female flowers are in different figs), check out this excellent site for all you could ever want to know.]

Says Who?

  • Compton et al. (2010) Biology Letters 6: 838-842
  • Cook et al. (2004) Journal of Evolutionary Biology 17: 238-246
  • Kjellberg et al. (2001)Proceedings of the Royal Society of London, Biology 268: 1113-1121
  • Proffit et al. (2009) Entomologia Experimentalis et Applicata 131: 46-57
  • Zhang et al. (2009) Naturwissenschaften 96: 543-549
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Advertising in the Wild… Not So Very Different (Ophrys sp.)

(Via: lastdragon.org)

Common Name: Bee Orchids

A.K.A.: Genus Ophrys

Vital Stats:

  • 30-40 recognised species in the genus
  • Grows to a height of 15-50 cm (6-20”)
  • The name Ophrys comes from a word meaning “eyebrow” in Greek, for the fuzzy edges of the petals
  • First mentioned in ancient Roman literature by Pliny the Elder (23-79 A.D.)

Found: Throughout most of Europe and the British Isles

It Does What?!

We tend to think of animals (including humans) as using plants to serve our ends exclusively- we eat them, clothe ourselves with them, build homes with them, and so on. But for all the obvious ways in which the animal kingdom takes advantage of the plants, there are numerous, more subtle, ways that they use us to do their bidding. One of those ways is as pollinators; plants enlist animals to help them reproduce. And while that enlistment often takes a rather mundane form – a bit of pollen brushed onto a bird’s head as it sips nectar, say – sometimes a group of plants will get a bit more creative about it. Such is the case with the bee orchids.

These highly specialised flowers depend on very specific relationships with their pollinators; often only a single species of bee (or wasp, in some cases) will pollinate a given species of orchid. Without those pollinators, the orchids can’t produce seed and would die out. So how do you control a free-roving creature that has other places to be? Why, sex, obviously. (Isn’t that the basis of most advertising?) The bee orchid has evolved a flower that not only looks, but smells like a virgin female of the bee species which pollinates it.

May not be appropriate for younger readers.
(Via: This Site)

At a distance, the bee detects the pheromones of a receptive female. Once he moves in closer, there she is, sitting on a flower, minding her own business. So he flies in and attempts to do his man-bee thing, only to find that he’s just tried to mate with a plant. Mortified (I imagine), he takes off, but with a small packet of pollen stuck to his head. He’s memorised the scent of this flower now and won’t return to it, but amazingly, the orchids vary their scent just slightly from one flower to the next, even on the same plant, so that the duped bee can never learn to distinguish an orchid from a female. What’s more, because the scent is more different between plants than between flowers on the same plant, he is more likely to proceed to a different plant, decreasing the chances that an orchid will self-fertilise.

Hilariously, researchers have shown that, due to their higher levels of scent variation compared to true female bees (variety being the spice of life, right guys?), male bees actually prefer the artificial pheromones of the orchids over real, live females. In experiments where males were given a choice between mating with an orchid and mating with a bee, they usually chose the flower, even if they had already experienced the real thing.

So there you have it. Plants: master manipulators of us poor, stupid animals.

Who could resist?
(Via: Wikia)

Says Who?

  • Ayasse et al. (2000) Evolution 54(6): 1995-2006
  • Ayasse et al. (2003) Proceedings of the Royal Society, London B. 270: 517-522
  • Streinzer et al. (2009) Journal of Experimental Biology 212: 1365-1370
  • Vereecken & Schiestl (2008) Proceedings of the National Academy of Science 105(21): 7484-7488
  • Vereecken et al. (2010) Botanical Review 76: 220-240
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EVOLUTION TAG TEAM, Part 1: Acacia Domatia

The first in an ongoing series of biology’s greatest duos. (Here’s Part 2 and Part 3)

Home, Sweet Home.
(via: Flickr)

Common Name (Plants): Bullhorn Acacias, Whistling Thorns

  • A.K.A.: Acacia cornigera, Acacia drepanolobium, and several other Acacia species

Common Name (Ants): Acacia Ants

  • A.K.A.: Pseudomyrmex and Crematogaster species

Found: Central America (Bullhorn Acacias) and East Africa (Whistling Thorns)

It Does What?!

Life as a tree is tough, particularly when you live in a part of the world that’s home to the biggest herbivores on Earth and happen to have delicate, delicious leaves. Such is the case for the African acacias. Without sufficient defences, they’d be gobbled up in no time by elephants, rhinos, and giraffes. The trees are known for having huge, sharp thorns, but even that’s sometimes not enough; the lips and tongues of giraffes are so tough and dexterous, they can often strip the leaves right out from between the thorns. So what’s a stressed acacia to do? Recruit a freaking army, that’s what.

Pseudomyrmex ferruginea: the giraffe’s worst enemy.
(Photo by April Nobile)

A few species of acacia in both Africa and Central America (where the herbivores are smaller, but no less voracious) have developed a symbiosis wherein they enjoy the services of ant colonies numbering up to 30,000 individuals, tirelessly patrolling their branches 24 hours a day. Should a hungry elephant or goat wander up and take a bite, nearby patrol ants will call in reinforcements and soon the interloper will be utterly overrun with angry, biting ants. What’s more, the protection extends beyond just animal threats. The ants will go so far as to kill other insects, remove fungal pathogens from the surface of the tree and even uproot nearby seedlings because, you know, they might eventually steal some sunlight from the beloved acacia.

“Trespassers Will Be Drawn and Quartered”
(via Wikimedia Commons)

So what do the troops get out of this? Quite a bit, actually. In ant-protected acacias (‘myrmecophytes’, they’re called), the thorns that normally grow at the base of a leaf swell up. In the Central American species, they grow into something that looks like a bull’s horn (hence their common name), while the African ones become more bulbous. These specialized structures, called domatia, are hollow inside and serve as very convenient housing for the ants. What’s more, the trees produce not one, but two different kinds of nourishment for the colony- regular, and baby food. The adult ants will feed from a sweet liquid exuded by nectaries on the branches. Meanwhile, on the tips of the tree’s leaflets, small white structures called Beltian bodies are formed which are high in the protein every growing child ant-larva needs. These are collected by workers and inserted right into the larval pouches, to be eaten before the ants are even fully formed.

The Bullhorn Acacia, now with more Beltian bodies!
(via Flickr)

Sounds like the perfect partnership, right? Usually, yes, but in nature, a symbiosis is only a symbiosis until one side figures out how to take advantage of the other. From the ants’ side, for example, any energy spent by the tree on reproduction is energy not spent on new homes and sweet, sweet nectar for them. Therefore, the ants will sometimes systematically nip all the flowers off the tree as it attempts to bloom. They’ll also prune the acacia’s outward growth if those new shoots may come into contact with a neighbouring tree, allowing invasion by another ant colony. Conversely, if herbivores become scarce and the acacia no longer requires such a strong protection force, it will begin to produce fewer domatia and less nectar in a move to starve some of the ants out. This has been shown to actually be a bad strategy for the acacia, since the soldiers, not to be outsmarted by a tree, turn to farming and begin raising sap-sucking insects on the bark, thereby getting their sugar fix anyway. And so it goes, oscillating between advantageous partnership and opportunistic parasitism… like so many things in life.

The roomier, more spacious African domatium.
(Image by Martin Sharman)

[Side note: While I’ve never personally encountered ant-acacias, I have disturbed an ant-protected tree of another family in the rainforests of Guyana, and can attest to the fact that the retaliation was both swift and intense. I was in a small boat at the edge of a river collecting botanical specimens, and I nearly jumped in the river to escape the onslaught. Don’t mess with ants.]

Says Who?

  • Clement et al. (2008) Behav. Ecol. Sociobiol. 62: 953-962.
  • Frederickson (2009) American Naturalist 173(5): 675-681.
  • Huntzinger et al. (2004) Ecology 85(3): 609-614.
  • Janzen (1966) Evolution 20(3): 249-275.
  • Nicklen & Wagner (2006) Oecologia 148: 81-87.
  • Stapley (1998) Oecologia 115: 401-405.
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The Stinging Tree, or, Australia Hates Mammals

Can’t Touch This
(via: anhs.com.au)

Common Name: Stinging Tree, Gympie-Gympie

A.K.A.: Dendrocnide moroides

Found: Rainforests of Northeastern Australia

It Does What?!

Australia, which was apparently intended only for the very bravest of human beings, is home to many of the world’s most poisonous snakes, spiders, and scorpions. Even the surrounding ocean is exceptional for the number of ridiculously venomous species it contains. Still, a person could be forgiven for thinking that, so long as they stay out of the water and keep away from the creepy-crawlies, they’ll be okay. Ha ha ha… nope. In Australia, everything is out to get you.

Meet Gympie-Gympie, the Stinging Tree (or to be more accurate, stinging shrub). Growing in rainforest clearings and along creek edges- anywhere the canopy is broken- this two metre (6.5ft) high plant has large, heart-shaped leaves and juicy purple fruit. And every square centimetre of it, from the soil on up, is covered in tiny, poison-filled hypodermic needles. These hollow silicon needles are delicate enough to break off at the slightest touch, leaving them embedded in the skin of whatever creature was unfortunate enough to do so. The skin will often then close over them, making the needles nearly impossible to remove. The substance they’re filled with is a very potent neurotoxin with a very long shelf life- herbarium specimens of the plant collected in 1910 are still able to cause pain. And since the body is unable to break down silicon, this all adds up to a very long punishment for a very small mistake.

Go on, I dare you.
(Photo by Melanie Cook)

A brief brush against a stinging tree produces intense pain that peaks after about half an hour, but can literally take years to subside completely. Numerous dogs and horses have died because the pain was so intense. There is even one official record of a human having died- a Dutch botanist of the 1920s. Oddly enough, no actual tissue damage is done by the neurotoxin- death due to the plant is attributed to heart failure due to the shock of the pain, described by one researcher, Dr. Marina Hurley, as “like being burnt with hot acid and electrocuted at the same time.” An ex-serviceman who fell right into one of the trees while crossing a creek in the 1940s describes having had to be tied down to his hospital bed for three weeks because the discomfort was so intense. One intrepid/insane researcher actually purified the neurotoxin and injected himself with it, suffering terribly and thereby proving that the toxin, rather than the needles, causes the majority of the pain. But not all of it… simply standing near a gympie-gympie for an extended period can cause allergic reactions and nosebleeds as the needles are shed in the wind.

“I eat neurotoxins for breakfast.”
(Via: Billabong Sanctuary)

So this must be just about the best herbivore-defence system ever, right? Amusingly, no. The trees still undergo heavy damage due to hungry spiders, ants, snails, and especially beetles, all of which can avoid its defences. The tree is even prey to one species of marsupial, the red-legged pademelon, which is either immune to the neurotoxin or has enormous pain tolerance. So why develop this extensive arsenal if it’s completely ineffective? One expert has suggested that it may have evolved to protect the plants from the now long-extinct giant Diprotodonts which once inhabited the Australian rainforest, making it one more dangerous relic of a long-ended war. You win, stinging tree, you win.

[Fun Fact: The best way to attempt to remove some of those poisonous silicon needles embedded in your arm?  Wax hair removal strips, according to the Queensland ambulance service.]

Says Who?

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Smells like death, looks like… an Amorphophallus?

Amorphophallus titanum

Common Name(s): Corpse Flower, Titan Arum

A.K.A.: Amorphophallus titanum

Found: Sumatra, Western Indonesia

It Does What?!

Looking like something from an Enterprise away mission, this is a plant you won’t soon forget. For those who imagine that biologists don’t have a sense of humour, the scientific name of the Corpse Flower is Amorphophallus titanum, which is Latin for ‘giant misshapen penis.’ And it’s not a bad description; the plant produces a… well, vaguely penis-shaped bloom that grows up to three feet tall and, as if we needed more to snicker about, produces pulses of heat which move from the base to the tip, reaching temperatures of over 36 degrees Celcius (97 Fahrenheit).

It happens to every Amorphophallus at some point…
(Via: plantae.ca)

It’s actually a bit of a misnomer to call this phallic monstrosity a flower- it’s really an inflorescence, a structure on which smaller, individual flowers grow. In the case of Amorphophallus, that cone in the middle is called a spadix (think calla lilies or jack-in-the-pulpit… same plant family), and holds upwards of 900 tiny flowers, of which about half are male and half are female.

Naturally, all those tiny little flowers need to get pollinated in order to create more giant-penis-plants, and the pollinators of choice for Amorphophallus are carrion beetles and blowflies. How to attract the attention of your favoured pollinators in a busy Sumatran rainforest? You give them what they want – the stench of rotting flesh. Those pulses of heat I mentioned before actually serve a purpose; they work like a convection oven, throwing off a foul odour which rises above the canopy as the warmer air rises. This allows the scent signal to be carried over greater distances. And how bad does it smell? Researchers of the plant note that a principal chemical component of that funk is known to also be the main source of the delicate bouquet that is rotting human flesh. Mmm… For another overly-vivid mental picture, be sure to check out a close relative of the Corpse Flower, Helicodiceros muscivorus, a.k.a. Dead Horse Arum.

Says Who?

  • Barthlott et al. (2009) Plant Biology 11: 499-505.
  • Shirasu et al. (2010) Biosci. Biotechnol. Biochem. 74(12): 2550-2554.