The Discovery of a Living Fossil

In the early 1940’s, the genus Metasequoia was only known scientifically in fossil form. It had, in its day, been a widespread genus, found commonly in many areas across the Northern Hemisphere. It thrived among the dinosaurs. However, sometime during the Pliocene, the genus was thought to have died out. Thousands of fossils were left behind, and that would have been the end of the story had a member of its genus not been discovered still alive in a Chinese province later that decade. Its discovery is easily one of the greatest botanical stories of the 20th century, fascinating in its own right. The circumstances surrounding its scientific description, as it turns out, are equally interesting.

In the January 2016 issue of Landscape Architecture Magazine, Kyna Rubin details the event in an article entitled The Metasequoia Mystery. It’s the type of story that you almost need a crazy wall to sort out. A broad cast of characters interacted at various levels in order to make this profound discovery during a tumultuous time when the world was at war and China was being invaded by Japan.

Speaking of Japan, let’s start there. In 1941, Japanese paleobotanist, Shigeru Miki, published research describing fossils that for decades were thought to be either Sequoia or Taxodium as a new genus, Metasequoia. As Rubin points out, due to the war, “not every Chinese botanist would have had access to recent international research, let alone articles by botanists of an enemy country.” This could explain why in 1943 when Zhan Wang – a professer of forestry at Beijing University and the forest administrator for the Ministry of Agriculture and Forestry – was introduced to a living Metasequoia by an old classmate and local villagers in the Hubei Province, he wasn’t sure what he was looking at.

The tree was obviously important to the local people. They called it shuisa (water fir) and had built a shrine around it. Wang collected several branches and some cones that had fallen on a rooftop. At the time he identified it as Glyptostrobus pensilis (water pine), a tree common to the area; but he may have wondered if this was correct.

Eventually Wang’s samples and the details of his collection were brought to the attention of Wanjun Zheng, a dendrologist at the National Central University. Intrigued, Zheng sent his graduate student, Jiru Xue, to collect more samples from the same tree that Wang had encountered. These samples were more complete, and when they were presented to Xiansu Hu – the director of Fan Memorial Institute of Biology in Beijing – the mystery was solved. Hu had access to Miki’s research and concluded that what they had was a living fossil.

In 1948, Hu and Zheng published a paper describing the species and giving it the official name, Metasequoia glyptostroboides. The discovery ignited the botanical community as well as the general public, and soon seeds of what became commonly known as dawn redwood were being disseminated across the globe. Unfortunately, Wang’s contribution was not mentioned in the original paper, and the exact account of the discovery became convoluted.

photo credit: wikimedia commons

Dawn redwood (Metasequoia glytostroboides) is a deciduous, medium to large tree. Its cones are round and about 1 inch long. Its leaves are oppositely arranged and have a feather-like appearance. Its bark is fibrous, stringy, and red-brown to gray in color.  (photo credit: wikimedia commons)

At some point, a discussion between Zheng and a forester named Duo Gan (also known as Toh Kan) revealed that Gan had come across the tree in 1941, but he did not make any collections. Despite Zheng learning of Gan’s encounter after Zheng and Hu’s original paper had been published, Gan’s story became prominent, further obscuring the role that Wang played.

It’s important to note that none of Wang’s original collections were used as the type specimen – the particular specimen of an organism to which the scientific name is formally attached and is referred to in the scientific literature. The type specimen was collected by Xue. This is not uncommon, as initial collections may not always be in the best condition and may not include all the parts and pieces necessary to identify and describe a new species. But, as Rubin notes, “it was Wang’s specimens [that Zheng and others] had first examined and those specimens brought the tree to their attention to begin with.” So Wang’s contribution is an important part of the story.

Thanks to Wang’s former students, his role in the discovery has received greater exposure. Jinshuang Ma in particular has made it his mission to highlight the part that Wang played in the event. Apart from maintaining a website all about Metasequoia, Ma also spent several years searching for a lost herbarium specimen collected by Wang, which he found in an abandoned herbarium in Nanjing. You can read about his find in this article from the August 2003 issue of the journal Taxon. (Ma’s well researched summary of the events surrounding the Metasequoia discovery is also worth reading.)

Failure to acknowledge Wang’s contribution (at least initially) perhaps didn’t make waves outside of China, but in Rubin’s words, “the omission of Wang’s contribution sparked immediate hullabaloo inside China’s botanical circles in the late 1940’s.” Power and class differences likely played a big role. Hu and Zheng were established scholars that had received their educations in the United States and France respectively. Wang was young, from a remote village, and had not studied abroad. While Wang “went on to become one of China’s most distinguished forestry experts and botanists,” he was early in his career at the time of the Metasequoia discovery.

A deep respect for the elders in his field may be the reason that Wang’s students claim that he “never complained” about his treatment. His students go on to say that Wang “was not interested in personal gain,” and instead was simply satisfied to see that Metasequoia “was now growing successfully all over the world and was better protected.” It is listed as endangered on the IUCN Red List and would likely be extinct in its shrunken native range had awareness of its existence not come about when it did.

Fossil of Metasequoia occidentalis - photo credit: wikimedia commons

Fossil of Metasequoia occidentalis – photo credit: wikimedia commons

There are plenty of other interesting details to this story. Read the full article and check out the links on metasequoia.org to learn more. The account of Jiru Xue (also known as Hsueh Chi-Ju), the graduate student who collected the type specimens, is particularly interesting. Suprisingly, the tree Wang and Xue took their collections from is still alive today and is estimated to be over 400 years old.

Other longform article reviews on Awkward Botany:

Confidential Carnivore

This is a guest post. Words and images by Jeremiah Sandler

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If you live in North America or Europe, chances are you have seen Dipsacus fullonum, commonly called teasel.  Its tall (up to 2 meters), spiky flower stalks with large purple flowers are easy to spot in low-lands, ditches, or along highways.  Since this prolific seeder’s introduction to North America from Europe, it has steadily increased its habitat to occupy nearly each region of the United States. Of course, like all plants, teasel has its preferences and is more frequent in some areas than in others.

dipsacus fullonum_jeremiah sandler

Teasel is an unassuming, herbaceous biennial.  It takes two years to complete its life cycle: First-year growth is spent as a basal rosette, and second-year growth is devoted to flowering.  Standard biennial, right?  As of 2011, an experiment was conducted on this plant that changed the way we see teasel, and possibly all other similar plants.

“Here we report on evidence for reproductive benefits from carnivory in a plant showing none of the ecological or life history traits of standard carnivorous species.” -Excerpt from the report titled Carnivory in the Teasel Dipsacus fullonum — The Effect of Experimental Feeding on Growth and Seed Set by Peter J.A. Shaw and Kyle Shackleton.

We all have favorite carnivorous plants, Venus flytraps, pitcher plants, sundews, etc.. Their showy traps and various means of attracting insects are all marvels of evolution in the plant kingdom.  These insectivorous plants evolved these means of nutrient acquisition in an answer to the lack of nutrients in their environment’s soil.  In some of these plants, there is a direct relationship between number of insects consumed and the size of the entire plant. In others, there is no such relationship.

The unassuming, biennial teasel can now join the ranks of carnivore, or protocarnivore.  It didn’t evolve in bogs or swamps where soil nutrients are depleted.  It has no relationship to the standard carnivorous species. It doesn’t have any flashy traps. In fact, it has no obvious traits which suggest it can gain nutrients from insects. Teasel’s carnivorous habits can be likened somewhat to the carnivorous habits of bromeliads; water gathered in their leaves traps insects.

In Shaw and Shackleton’s experiment (done in two field populations), maggots were placed in water gathered in the center of some first-year rosettes of teasel.  Other rosettes in the same population were left alone as controls.  Not surprisingly, the teasels which were ‘fed’ larvae did not change in overall size.  The size of the overwintering rosette did not offer any predictability towards the size of flower shoots for the coming year. However, something strange did happen:

“…addition of dead dipteran larvae to leaf bases caused a 30% increase in seed set and the seed mass:biomass ratio.”…“These results provide the first empirical evidence for Dipsacus displaying one of the principal criteria for carnivory”

Teasel has some physiology to absorb nutrients from other macroorganisms despite teasel evolving in an entirely different setting than typical carnivorous plants.  Teasel’s already proficient reproductive capacity is enhanced by using insects as a form of nutrients in a controlled setting.  

Many exciting questions have been raised by this experiment. How has this absorption mechanism come about, without the obvious use of lures or other structures to attract insects? And how does teasel maximize upon its own morphology in the wild, if at all?  What would the results be if these experiments were recreated on other similar species?

There are studies being conducted all the time that further the boundaries of what we know about these stationary organisms. There are new discoveries waiting just around the corner. Carnivory in plants is amazing because it transcends common notions about plants; especially in the case of the unassuming teasel.

Selected Resources:

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Jeremiah Sandler lives in southeast Michigan where he works in the plant health care industry. He has a degree in horticultural sciences and is an ISA certified arborist. He is interested in all things plant related and plans to own a horticulture business where he can share his passion with others. Follow Jeremiah on Instagram: @j.deepsea

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Poisonous Plants: Lima Beans

I don’t recall being a picky eater as a child, but one food I could barely stomach was lima beans. The smell, the texture, the taste, even the look of them, really didn’t sit well with me. I know I’m not alone in this sentiment. Lima beans are a popular thing to hate, and I have avoided them ever since I was old enough to decide what was allowed on my plate. To be fair, the only lima beans I remember trying were the ones included in the familiar bag of frozen mixed vegetables, which might explain why I didn’t like them. But little did I know there is another reason to avoid them – lima beans are poisonous.

That’s a strong statement. In case you’ve eaten lima beans recently or are about to, I should ease your concerns by telling you that you have little to worry about. Commonly cultivated lima beans are perfectly safe to eat as long as they are cooked properly, and even if they are eaten raw in small doses, they are not likely to hurt you. But again, why are you eating lima beans? They’re gross.

lima beans in cans

Phaseolus lunatus – commonly known as lima bean as well as a number of other common names – is in the legume family (Fabaceae) and is native to tropical America. It is a perennial, twining vine that reaches up to 5 meters. It has trifoliate leaves that are alternately arranged, and its flowers are typically white, pink, or purple and similar in appearance to pea flowers and other flowers in the legume family. The fruits are hairy, flat, 5 – 10 cm long, and often in the shape of a half moon. The seeds are usually smooth and flat, but are highly variable in color, appearing in white, off-white, olive, brown, red, black, and mottled.

P. lunatus experienced at least two major domestication events – one in the Andes around 4ooo years ago and the other in Central America more than 1000 years ago. Studies have found that the first event yielded large seeded varieties, and the second event produced medium to small seeded varieties. Wild types of P. lunatus have been given the variety name sylvester, and cultivated types are known as variety lunatus; however, these don’t appear to be accepted names by plant taxonomists and perhaps are just a way of distinguishing cultivated plants from plants growing in the wild, especially in places where P. lunatus has become naturalized such as Madagascar.

Distinguishing wild types from cultivated types is important though, because wild types are potentially more poisonous. Lima bean, like several other plants we eat, contains compounds in its tissues that produce cyanide. These cyanide producing compounds are called cyanogenic glucosides and are present in many species of plants as a form of defense against herbivores. The predominant cyanogenic glucoside in lima beans is called linamarin, which is also present in cassava and flax.

Fruits of lima bean (Phaseolus lunatus) - photo credit: wikimedia commons

Fruits of lima bean (Phaseolus lunatus) – photo credit: wikimedia commons

In order for lima beans to poison you, they must be chewed. Chewing brings linamarin and the enzymes that react with it together. Both compounds are present in the cells of lima beans, but they reside in different areas. Once they are brought together, a reaction ensues and hydrogen cyanide is produced. Because cyanide isn’t produced until after the plant is consumed, the symptoms of cyanide poisoning can take a little while to occur – often several hours.

Cyanide poisoning is not a pretty thing. First comes sweating, abdominal pain, vomiting, and lethargy. If the poisoning is severe, coma, convulsions, and cardiovascular collapse can occur. There are treatments for cyanide poisoning, but if treatment comes too late or if the dose is large enough, death results.

Cassava (Manihot esculenta) is particularly well known for its history of cyanide poisonings. It is a staple crop of people living in tropical areas of Africa and South America. Humans can readily metabolize small amounts of cyanide, and processes like crushing and rinsing, cooking, boiling, blanching, and fermenting render cassava safe to eat. However, consuming cassava that isn’t prepared properly on a consistent basis can result in chronic illnesses, such as konzo, which is a major concern among cultures in which cassava is an important food source.

I guess I should reiterate at this point that most cultivated lima beans contain low (read “safe”) levels of cyanogenic glucosides and, particularly when cooked, are perfectly safe to eat. I’m still not totally convinced that I should eat them though. While researching this article I came across numerous sites claiming that lima beans are delicious while offering various recipes to prove it. I even came across this story in which a self-proclaimed “lima bean loather” was converted to the side of the lima bean lovers. I don’t fancy myself much of a cook, so I’m hesitant to attempt a lima bean laden recipe for fear that it will only make me hate them more. If anyone out there thinks they can convince me otherwise with their tasty creation, be my guest.

And now a haiku:

You are lima beans
I despised you as a child
Perhaps unfairly?

Follow these links to learn more about cyanide producing crops and lima beans:

Book Review: Bringing Nature Home

Since Bringing Nature Home by Douglas Tallamy was first published in 2007, it has quickly become somewhat of a “classic” to proponents of native plant gardening. As such a proponent, I figured I ought to put in my two cents. Full disclosure: this is less of a review and more of an outright endorsement. I’m fawning, really, and I’m not ashamed to admit it.

9780881929928l

The subtitle pretty much sums it up: “How You Can Sustain Wildlife with Native Plants.” Ninety three pages into the book, Tallamy elaborates further: “By favoring native plants over aliens in the suburban landscape, gardeners can do much to sustain the biodiversity that has been one of this country’s richest assets.” And one of every country’s richest assets, as far as I’m concerned. “But isn’t that why we have nature preserves?” one might ask. “We can no longer rely on natural areas alone to provide food and shelter for biodiversity,” Tallamy asserts in the Q & A portion of his book. Humans have altered every landscape – urban, suburban, rural, and beyond – leaving species of all kinds threatened everywhere. This means that efforts to protect biodiversity must occur everywhere. This is where the You comes in. Each one of us can play a part, no matter how small. In closing, Tallamy claims, “We can each make a difference almost immediately by planting a native nearby.”

A plant is considered native to an area if it shares a historical evolutionary relationship with the other organisms in that area. This evolutionary relationship is important because the interactions among organisms that developed over thousands, even millions, of years are what define a natural community. Thus, as Tallamy argues, “a plant can only function as a true ‘native’ while it is interacting with the community that historically helped shape it.” A garden designed to benefit wildlife and preserve biodiversity is most effective when it includes a high percentage of native plants because other species native to the area are already adapted to using them.

Plants (and algae) are at the base of every food chain – the first trophic level – because they produce their own food using the sun’s energy. Organisms that are primarily herbivores are at the second trophic level, organisms that primarily consume herbivores are at the third trophic level, and so on. As plants have evolved they have developed numerous defenses to keep from being eaten. Herbivores that evolved along with those plants have evolved the ability to overcome those defenses. This is important because if herbivores can’t eat the plants then they can’t survive, and if they don’t survive then there will be little food for organisms at higher trophic levels.

The most important herbivores are insects simply because they are so abundant and diverse and, thus, are a major food source for species at higher trophic levels. The problem is that, as Tallamy learned, “most insect herbivores can only eat plants with which they share an evolutionary history.” Insects are specialized as to which plants they can eat because they have adapted ways to overcome the defenses that said plants have developed to keep things from eating them. Healthy, abundant, and diverse insect populations support biodiversity at higher trophic levels, but such insect populations won’t exist without a diverse community of native plants with which the insects share an evolutionary history.

That is essentially the thesis of Tallamy’s book. In a chapter entitled “Why Can’t Insects Eat Alien Plants?” Tallamy expounds on the specialized relationships between plants and insects that have developed over millennia. Plants introduced from other areas have not formed such relationships and are thus used to a much lesser degree than their native counterparts. Research concerning this topic was scarce at the time this book was published, but among other studies, Tallamy cites preliminary data from a study he carried out on his property. The study compared the insect herbivore biomass and diversity found on four common native plants vs. five common invasive plants. The native plants produced 4 times more herbivore biomass and supported 3.2 times as many herbivore species compared to the invasive plants. He also determined that the insects using the alien plants were generalists, and when he eliminated specialists from the study he still found that natives supported twice as much generalist biomass.

Apart from native plants and insects, Tallamy frames much of his argument around birds. Birds have been greatly impacted by humans. Their populations are shrinking at an alarming rate, and many species are threatened with extinction. Tallamy asserts, “We know most about the effects of habitat loss from studies of birds.” We have destroyed their homes and taken away their food and “filled their world with dangerous obstacles.” Efforts to improve habitat for birds will simultaneously improve habitat for other organisms. Most bird species rely on insects during reproduction in order to feed themselves and their young. Reducing insect populations by filling our landscapes largely with alien plant species threatens the survival of many bird species.

In the chapters “What Should I Plant?” and “What Does Bird Food Look Like?,” Tallamy first profiles 20 groups of native trees and shrubs that excel at supporting populations of native arthropods and then describes a whole host of arthropods and arthropod predators that birds love to eat. Tallamy’s fascinating descriptions of the insects, their life cycles, and their behaviors alone make this book worth reading. Other chapters that are well worth a look are “Who Cares about Biodiversity?” in which Tallamy explains why biodiversity is so essential for life on Earth, and “The Cost of Using Alien Ornamentals” in which Tallamy outlines a number of ways that our obsession with exotic plants has caused problems for us and for natural areas.

Pupa of ladybird beetle on white oak leaf (photo credit: wikimedia commons)

Pupa of a ladybird beetle on a white oak leaf. “The value of oaks for supporting both vertebrate and invertebrate wildlife cannot be overstated.” – Doug Tallamy (photo credit: wikimedia commons)

Convincing people to switch to using native plants isn’t always easy, especially if your argument involves providing habitat for larger and more diverse populations of insects. For those who are not fans of insects, Tallamy explains that “a mere 1%” of the 4 million insect species on Earth “interact with humans in negative ways.” The majority are not pests. It is also important to understand that even humans “need healthy insect populations to ensure our own survival.” Tallamy also offers some suggestions on how to design and manage an appealing garden using native plants. A more recent book Tallamy co-authored with fellow native plant gardening advocate Rick Darke called The Living Landscape expands on this theme, although neither book claims to be a how to guide.

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2015: Year in Review

Raise your glass. 2015 has come to a close, and Awkward Botany is turning three. Two great reasons to celebrate.

I started the year with the goal of posting at least once per week. Consider that goal accomplished, with a couple of bonus posts thrown in for good measure. I had also deemed 2015 the “Year of Pollination.” The underlying purpose was to teach myself more about pollinators and pollination while also sharing my interest in pollination biology with the wider world. That endeavor yielded 17 posts. There is still so much to learn, but we are making some headway. I started two new series of posts (Poisonous Plants and Botany in Popular Culture) and I continued with two others (Ethnobotany and Drought Tolerant Plants). I also went on a couple of field trips and wrote a few book reviews. All of that is reflected below in “Table of Contents” fashion.

Year of Pollination:

Botany in Popular Culture

Poisonous Plants

Ethnobotany

Drought Tolerant Plants

Book Reviews

Field Trips

Three posts that perhaps didn’t get the attention they deserve:

juniper in the snow

Going forward, I will continue to post regularly – as there is no shortage of plant-related things to write about – but I will likely take a week off here and there. I have other projects in mind – some related to Awkward Botany, some not – that will certainly demand much of my attention and time. I have some big ideas for Awkward Botany and beyond, and I will share those with the wide world in due time. For now, I would just like to say thanks all for reading, for commenting, and for sharing Awkward Botany with your friends. Overall, it has been a great year here at Awkward Botany headquarters, and I have you to thank for that. I feel privileged to be part of a community that is infatuated with plants and is fascinated by the natural world.

Good riddance to 2015. It was good, but it gets better. Now we look ahead to 2016. May it be filled with peace, love, and botany.

Year of Pollination: Botanical Terms for Pollination, part two

“The stage is set for reproduction when, by one means or another, compatible pollen comes to rest on a flower’s stigma. Of the two cells within a pollen grain, one is destined to grow into a long tube, a pollen tube, that penetrates the pistil’s tissues in search of a microscopic opening in one of the ovules, located in the ovary. … The second of a pollen grain’s cells divides to become two sperm that move through the pollen tube and enter the ovule.” – Brian Capon, Botany for Gardeners

“Once pollination occurs, the next step is fertilization. Pollen deposited on the sticky stigma generates a fine pollen tube that conveys the sperm through the style to the ovary, where the ovules, or eggs, have developed. After fertilization, the rest of the flower parts wither and are shed as the ovary swells with seed development.” – Rick Imes, The Practical Botanist

Pollination tells the story of a pollen grain leaving an anther by some means – be it wind, water, or animal – and finding itself deposited atop a stigma. As long as the pollen and stigma are compatible, the sex act proceeds. In other words, the pollen grain germinates. One of the pollen grain’s cells – the tube nucleus – grows down the length of the style, forming a tube through which two sperm nuclei can travel. The sperm nuclei enter the ovary and then, by way of a micropyle, enter an ovule. Inside the ovule is the female gametophyte (also referred to as the embryo sac). One sperm nucleus unites with the egg nucleus to form a zygote. The remaining sperm nucleus unites with two polar nuclei to form a triploid cell which becomes the endosperm. The sex act is complete.

The illustration on the left includes the cross-section of a pistil showing the inside the ovary where pollen tubes have made their way to the ovules. The illustration on the right shows pollen grains germinating on a stigma and their pollen tubes begining to work their way down the style. (photo credit: wikimedia commons)

The illustration on the left includes the cross section of a pistil showing the inside of the ovary where pollen tubes have made their way to the ovules. The illustration on the right shows pollen grains germinating on a stigma and pollen tubes as they work their way down the style. (image credit: wikimedia commons)

The zygote divides by mitosis to become an embryo. The endosperm nourishes the development of the embryo. The ovule matures into a seed, and the ovary develops into a fruit. During this process, the remaining parts of the flower wither and fall away. In some cases, certain flower parts remain attached to the fruit or become part of the fruit. The flesh of an apple, for example, is formed from the carpels and the receptacle (the thickened end of a flower stem – peduncle – to which the parts of a flower are attached).

As the seed matures, the endosperm is either used up or persists to help nourish the embryonic plant after germination. Mature seeds that are abundant in endosperm are called albuminous. Examples include wheat, corn, and other grasses and grains. Mature seeds with endosperm that is either highly reduced or absent are called exalbuminous – beans and peas, for example. Certain species – like orchids – do not produce endosperm at all.

The cross section of a corn kernel showing the endosperm and the embryo (image credit: Encyclopedia Britannica Kids)

The cross section of a corn kernel showing the endosperm and the embryo (image credit: Encyclopedia Britannica Kids)

It is fascinating to consider that virtually every seed we encounter is the result of a single pollen grain making its way from an anther to a stigma, growing a narrow tube down a style, and fertilizing a single ovule. [Of course there are always exceptions. Some plants can produce seeds asexually. See apomixis.] Think of this the next time you are eating corn on the cob or popcorn – each kernel is a single seed – or slicing open a pomegranate to reveal the hundreds of juicy seeds inside. Or better yet, when you are eating the flesh or drinking the milk of a coconut. You are enjoying the solid and liquid endosperm of one very large seed.

Much more can be said about pollination and the events surrounding it, but we’ll save that for future posts. The “Year of Pollination” may be coming to an end, but there remains much to discover and report concerning the subject. For now, here is a fun video to help us review what we’ve learned so far:

 

Also, take a look at this TED talk: The Hidden Beauty of Pollination by Louie Schwartzberg

And finally, just as the “Year of Pollination” was coming to an end I was introduced to a superb blog called The Amateur Anthecologist. Not only did it teach me that “anthecology” is a term synonymous with pollination biology, it has a great series of posts called “A Year of Pollinators” that showcases photographs and information that the author has collected for various groups of pollinators over the past year. The series includes posts about Bees, Wasps, Moths and ButterfliesFlies, and Beetles, Bugs, and Spiders.

Year of Pollination: Botanical Terms for Pollination, part one

When I began this series of posts, I didn’t have a clear vision of what it would be. I had a budding interest in pollination biology and was anxious to learn all that I could. I figured that calling 2015 the “Year of Pollination” and writing a bunch of pollination-themed posts would help me do that. And it has. However, now that the year is coming to a close, I realize that I neglected to start at the beginning. Typical me.

What is pollination? Why does it matter? The answers to these questions seemed pretty obvious; so obvious, in fact, that I didn’t even think to ask them. That being said, for these last two “Year of Pollination” posts (and the final posts of the year), I am going back to the basics by defining pollination and exploring some of the terms associated with it. One thing is certain, there is still much to be discovered in the field of pollination biology. Making those discoveries starts with a solid understanding of the basics.

Pollination simply defined is the transfer of pollen from an anther to a stigma or – in gymnosperms – from a male cone to a female cone. Essentially, it is one aspect of plant sex, albeit a very important one. Sexual reproduction is one way that plants multiply. Many plants can also reproduce asexually. Asexual reproduction typically requires less energy and resources – no need for flowers, pollen, nectar, seeds, fruit, etc. – and can be accomplished by a single individual without any outside help; however, there is no gene mixing (asexually reproduced offspring are clones) and dispersal is limited (consider the “runners” on a strawberry plant producing plantlets adjacent to the mother plant).

To simplify things, we will consider only pollination that occurs among angiosperms (flowering plants); pollination/plant sex in gymnosperms will be discussed at another time. Despite angiosperms being the youngest group of plants evolutionarily speaking, it is the largest group and thus the type we encounter most.

A flower is a modified shoot and the reproductive structure of a flowering plant. Flowers are made up of a number of parts, the two most important being the reproductive organs. The androecium is a collective term for the stamens (what we consider the male sex organs). A stamen is composed of a filament (or stalk) topped with an anther – where pollen (plant sperm) is produced. The gynoecium is the collective term for the pistil (what we consider the female sex organ). This organ is also referred to as a carpel or carpels; this quick guide helps sort that out. A pistil consists of the ovary (which contains the ovules), and a style (or stalk) topped with a stigma – where pollen is deposited. In some cases, flowers have both male and female reproductive organs. In other cases, they have one or the other.

photo credit: wikimedia commons

photo credit: wikimedia commons

When pollen is moved from an anther of one plant to a stigma of another plant, cross-pollination has occurred. When pollen is moved from an anther of one plant to a stigma of the same plant, self-pollination has occurred. Cross-pollination allows for gene transfer, and thus novel genotypes. Self-pollination is akin to asexual production in that offspring are practically identical to the parent. However, where pollinators are limited or where plant populations are small and there is little chance for cross-pollination, self-pollination enables reproduction.

Many species of plants are unable to self-pollinate. In fact, plants have evolved strategies to ensure cross-pollination. In some cases, the stamens and pistils mature at different times so that when pollen is released the stigmas are not ready to receive it or, conversely, the stigmas are receptive before the pollen has been released. In other cases, stigmas are able to recognize their own pollen and will reject it or inhibit it from germinating. Other strategies include producing flowers with stamens and pistils that differ dramatically in size so as to discourage pollen transfer, producing separate male and female flowers on the same plant (monoecy), and producing separate male and female flowers on different plants (dioecy).

As stated earlier, the essence of pollination is getting the pollen from the anthers to the stigmas. Reproduction is an expensive process, so ensuring that this sex act takes place is vital. This is the reason why flowers are often showy, colorful, and fragrant. However, many plants rely on the wind to aid them in pollination (anemophily), and so their flowers are small, inconspicuous, and lack certain parts. They produce massive amounts of tiny, light-weight pollen grains, many of which never reach their intended destination. Grasses, rushes, sedges, and reeds are pollinated this way, as well as many trees (elms, oaks, birches, etc.) Some aquatic plants transport their pollen from anther to stigma via water (hydrophily), and their flowers are also simple, diminutive, and produce loads of pollen.

Inforescence of big bluestem (Andropogon gerardii), a wind pollinated plant - pohto credit: wikimedia commons

Inflorescence of big bluestem (Andropogon gerardii), a wind pollinated plant – photo credit: wikimedia commons

Plants that employ animals as pollinators tend to have flowers that we find the most attractive and interesting. They come in all shapes, sizes, and colors and are anywhere from odorless to highly fragrant. Odors vary from sweet to bitter to foul. Many flowers offer nectar as a reward for a pollinator’s service. The nectar is produced in special glands called nectaries deep within the flowers, inviting pollinators to enter the flower where they can be dusted with pollen. The reward is often advertised using nectar guides – patterns of darker colors inside the corolla that direct pollinators towards the nectar. Some of these nectar guides are composed of pigments that reflect the sun’s ultraviolet light – they are invisible to humans but are a sight to behold for many insects.

In part two, we will learn what happens once the pollen has reached the stigma – post-pollination, in other words. But first, a little more about pollen. The term pollen actually refers to a collection of pollen grains. Here is how Michael Allaby defines “pollen grain” in his book The Dictionary of Science for Gardeners: “In seed plants, a structure produced in a microsporangium that contains one tube nucleus and two sperm nuclei, all of them haploid, enclosed by an inner wall rich in cellulose and a very tough outer wall made mainly from sporopollenin. A pollen grain is a gametophyte.”

A pollen grain’s tough outer wall is called exine, and this is what Allaby has to say about that: “It resists decay, and the overall shape of the grain and its surface markings are characteristic for a plant family, sometimes for a genus or even a species. Study of pollen grains preserved in sedimentary deposits, called palynology or pollen analysis, makes it possible to reconstruct past plant communities and, therefore, environments.”

Scanning electron microscope image of pollen grains from narrowleaf evening primrose (Oenothera fruticosa) - photo credit: wikimedia commons

Scanning electron microscope image of pollen grains from narrowleaf evening primrose (Oenothera fruticosa) – photo credit: wikimedia commons

Botany in Popular Culture: Futurama’s Holiday Spectacular

Matt Groening and David X. Cohen’s animated sitcom, Futurama, is replete with social commentary. Set in the 31st Century, it’s not surprising that much of that commentary involves environmental issues. Episode 13 of season 6 – a special, holiday season episode – addresses a number of such issues, including extinction, global warming, fossil fuel depletion, and Colony Collapse Disorder. The episode is broken up into three, distinct segments; each has its own storyline, but all – apart from being environmentally themed – center around traditional (in the fictional world of Futurama) holiday celebrations. Hence, the title of the episode: The Futurama Holiday Spectacular.

Botany plays a particularly prominent role in the first segment of the episode. In the 31st Century, Christmas has morphed into a holiday called Xmas. In the opening scene, the Planet Express Crew has decorated a palm tree to look like a Christmas tree. Looking despondent, Philip J. Fry (a pizza delivery boy from the 21st century who was inadvertently cryopreserved and thawed 1,000 years later) laments, “Something about Xmas just doesn’t feel like Christmas.”  Just then, the arrival of Santa is announced.

In the 31st Century, Santa Claus has been replaced by a robot called Robot Santa, and instead of gifts and holiday cheer, he brings violence and mayhem. The crew begins to lock down the Planet Express headquarters in preparation for Robot Santa’s arrival. Disturbed by this, Fry demands to know how “this crazy holiday” is celebrated – “preferably in song.”  At which point, Robot Santa bursts out of the fireplace singing, “It’s the violentest season of the year…”

robot santa

After a few violent exchanges between the crew and Robot Santa, Robot Santa sings, “The one thing that you need to make your Xmas Day splendiferous / Is a pine tree – a pine tree that’s coniferous.” The crew agrees; they need “an old-fashioned pine tree.” But there is one problem.

“Pine trees have been extinct for over 800 years,” explains Professor Farnsworth. Apparently, they were all chopped down and turned into toilet paper during something called “The Fifty-Year Squirts.” Yet, the Professor exclaims, “There is one hope and, as usual, it’s Norwegian!” And at that, the crew heads off to Norway.

In Norway, the crew arrives at the Svalbard Global Seed Vault where, as the Professor explains, “since 2008, the vault has preserved seeds of every known plant species in case of extinction.” They are confronted by a seed vault employee who asks why the crew is “pokey-poking about the seed vault – guardian of mankind’s precious botanical heritage there?”

The Professor tells the man that they are there to “rummage about a bit.” The crew notes that there is a Germ Warfare Repository that has been constructed right next to the seed vault and asks if there are any cross-contamination concerns. The man says, “No,” and then lets them inside where he brings them a container marked Pinus xmas. Amy notices some “splork” on the seeds and asks, “It’s not germs is it?” Again the man says, “No.” 

futurama2

The Planet Express crew at the Svalbard Global Seed Vault being presented with the seeds of Pinus xmas.

Back in New New York, Fry plants a pine tree seed outside the Planet Express building. A year later, a sapling as tall as Fry has emerged. Fry declares, “Now that’s a tree worth chopping down.” At that point, President Nixon pulls up in his limousine and sees the tree. “That’s what my poll numbers need, ” he says turning to Vice President Cheney – both of them animated heads in jars. Cheney orders Nixon to steal the tree.

The tree is transplanted in front of the White House. During the Xmas tree lighting ceremony, the tree begins to grow rapidly. Apparently it was contaminated with a weaponized virus after all. It begins to produce cones which then fly off the tree and explode. Shortly after the explosions, more pine trees begin to emerge and grow rapidly, at which point Leela exclaims, “Wait! This could be a good thing. Reforestation has begun!” However, this reforestation is occurring at an extremely rapid pace, and before long all land on Earth is completely covered in pine trees.

Soon, all manner of wildlife is found frolicking among the trees. Again Leela exclaims, “Arguably, this could be a good thing. The planet has returned to its primeval state!” The Professor concurs, “All these pine trees are fighting global warming by producing oxygen.”

But the “good news” doesn’t last long. The oxygen level continues to increase and quickly reaches 80%. Ignorantly, Bender decides to celebrate his own laziness with a cigar. As he lights it, the entire planet bursts into flames. Robot Santa returns to announce, “Ho ho ho! Everyone’s dead!”

Futurama

Similar dark comedy ensues in the other two segments as the crew learns about the holiday traditions of Robanukah and Kwanzaa. Again, both segments explore important environmental concerns in the process. Al Gore’s animated head in a jar makes appearances throughout the episode. If you are looking for some added hilarity during this holiday season – as well as some bleak environmental messaging – you can’t go wrong with Futurama’s Holiday Spectacular.

Interesting fact: In 2011, this episode of Futurama won an Environmental Media Award for best comedic television episode with an environmental message. EMA’s have been awarded since 1991 to “honor film and television productions and individuals that increase public awareness of environmental issues and inspire personal action on these issues.”

Harvester Ants – Seed Predators and Seed Dispersers

“The abundance of ants is legendary. A worker is less than one-millionth the size of a human being, yet ants taken collectively rival people as dominant organisms on the land. …  When combined, all ants in the world taken together weigh about as much as all human beings.” – Journey to the Ants by Bert Hölldobler and Edward O. Wilson

Considering how abundant and widely distributed ants are, it is easy to imagine the profound role they might play in the ecosystems of which they are a part. In fact, in the epilogue to Hölldobler and Wilson’s popular book about ants (quoted above), they conclude that in a world without ants, “species extinction would increase even more over the present rate, and the land ecosystems would shrivel more rapidly as the considerable services provided by these insects were pulled away.” It is no doubt then that ants, through their myriad interactions with their surroundings, are key players in terrestrial ecosystems.

photo credit: www.eol.org

photo credit: www.eol.org

Harvester ants offer a prime example of the important roles that ants can play. In the process of collecting seeds for consumption, harvester ants can help shape the abundance and distribution of the plants in their immediate environment. They do this by selecting the types and amounts of seeds they collect, by abandoning seeds along their collection routes, and by leaving viable seeds to germinate in and around their nests. Hölldobler and Wilson have this to say about harvester ants:

[The] numerical success [of ants] has allowed them to alter not just their nest environments, but the entire habitats in which they live. Harvesting ants, species that regularly include seeds in their diet, have an especially high impact. They consume a large percentage of the seeds produced by plants of many kinds in nearly all terrestrial habitats, from dense tropical forests to deserts. Their influence is not wholly negative. The mistakes they make by losing seeds along the way also disperse plants and compensate at least in part for the damage caused by their predation.

There are more than 150 species of harvester ants, spanning at least 18 genera. They are found throughout the world (except extreme cold locales) and are particularly common in arid to semi-arid environments. Pogonomyrmex is one the largest genera of harvester ants with nearly 70 species occurring throughout North, Central, and South America. Messor is another large genus of harvester ant species that mainly occurs in Europe, Asia, and Africa. Both genera build large nests and move massive amounts of soil in the process.

Seed dispersal by harvester ants (also known as diszoochory) is a type of secondary (or Phase II) seed dispersal. It is a case of serendipity, as the dispersal occurs largely by accident. Some plants, on the other hand, have developed a mutualistic relationship with ants, enlisting them to disperse their seeds by way of an elaiosome – a fleshy, nutritious structure attached to seeds that attracts ants. Seeds with such structures are picked up by ants and brought to their nests where the elaiosome is consumed and the seed is left to germinate. This form of ant-mediated dispersal is called myrmecochory and is typically not carried out by harvester ants.

photo credit: wikimedia commons

photo credit: wikimedia commons

Harvester ant colonies have both direct and indirect influences on their surrounding environments; however, there is a dearth of research elucidating the exact details of such influences. A paper published in the Annual Review of Ecology and Systematics in 2000 by MacMahon et. al. reviewed available studies concerning harvester ants and explored our current understanding of the influences that harvester ants (particularly those in the genus Pogonomyrmex) can potentially have on community structure and ecosystem functions. Following are some of the direct influences the authors listed:

  • Removal and consumption of seeds and other materials – The relative abundance of plant species can be affected by the selective removal of seeds. Harvester ants also collect leaves, twigs, pollen, flowers, vertebrate feces, and arthropod body parts.
  • Storage and rejection of seeds – Collected seeds can be dropped during transport, rejected after arriving at the nest, or abandoned in nest granaries. All result in the transport of seeds away from the parent plant and dispersal beyond the plant’s primary dispersal mechanisms.
  • Construction and maintenance of nests – All vegetation and debris is removed from the area immediately surrounding the nest including mature and emerging plants. This area is kept clear for the duration of the life of the colony and, in some cases, can be quite extensive.

Harvester ants can also influence soil properties and soil food webs within and in the vicinity of their nests. They bring large amounts of organic matter down into the soil and redistribute vast amounts of soil particles. Their actions also influence the amount of moisture in the soil surrounding their nests.

This is a mere distillation of the influences that harvester ants might have; see the paper by MacMahon et al. to learn more.

In an effort to better understand how the seed predation and seed dispersal behaviors of harvester ants might influence plant population dynamics, a research team in Spain used data obtained from field research to build a computer model that would predict changes over time. The study site was described as “open and heterogeneous shrubland” and the vegetation was stated to be in “a very early stage in the secondary succession” after being subject to “recurring fires.” The harvester ant colonies involved in the study consisted of three species in the genus Messor. The plant species selected for the study were three native shrubs whose seeds were known to be collected by the harvester ants. Each plant species differed slightly in the amount and size of seeds it produced and in its primary seed dispersal mechanism, which is important because the researchers hypothesized that “the effect of seed predation and seed dispersal may depend on plant attributes.”

Messor bouvieri (photo credit: www.eol.org)

Messor bouvieri (photo credit: www.eol.org)

Data obtained from simulated scenarios and field observations appeared to support this hypothesis; each shrub species interacted differently with the harvester ants. Coronilla minima benefited from “accidental” seed dispersal. Comparatively, it produces a high amount of large seeds, which are primarily dispersed by gravity. Despite predation, ant-mediated dispersal was an advantage. Dorycnium pentaphyllum produced the highest amount of seeds among the three shrub species; however, seed predation was found to have negative effects on its population dynamics. Its primary seed dispersal mechanism involves ballistics (the mechanical ejection of its seeds), so ant-mediated dispersal may not offer an advantage. Finally, Fumana ericoides, despite its limited primary seed dispersal and its comparatively low production of seeds was not affected by the actions of the harvester ants. The authors concluded that “some unknown factor is driving the population dynamics of this species, more than the action of ants.”

Studies such as this, while leaving many unanswered questions, help us understand the important role that harvester ants play in our world. Harvester ants, and ants in general, are truly among Earth’s most enthralling and influential creatures. Learn more about their complex behaviors and countless interactions with flora and fauna by checking out these three documentaries recommended by ANTfinity.