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Posts Tagged plant structure

moss s*x and springtails Alison Campbell Jul 22

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Sexual reproduction in flowering plants is often mediated by the birds & the bees (& other animal agents), but up until now the life cycle has appeared much simpler in plants like the mosses. Until fairly recently it was generally accepted that moss sex was a case of ‘just add water’: this released sperm from the male plants which could then swim in the film of water to where the female plants held their eggs. Of necessity this would mean that sperm dispersal could be only over quite short distances, of a few centimetres at most.

However, Todd Rosensteil and his colleagues (2012) decided to confirm the hypothesis that arthropods known as springtails could be involved in transferring sperm between male and female mosses. (Springtails and mosses evolved at the same time, during the Ordovician period.) They posed a number of questions: were springtails really acting as go-betweens in moss sex? If the answer was ‘yes’, how did the moss plants attract their little helpers? And, were the springtails important only if there was not much water around?

Using a common – & cosmopolitan – moss called Ceratodon purpureus, Rosenstiel and his colleagues first determined that female C.purpureus plants emit a significantly greater number of volatile organic chemicals (VOCs), which could act as signals to springtails, than male mosses do. They then carried out a number of experiments.

First of all, they gave springtails a choice between male and female moss plants – the tiny arthropods were much more likely to go for the female plants. (However, it’s not yet clear why the springtails respond positively to this signal: do they get some sort of a food reward?) The same was true when the springtails were given no visual cues & were simply offered a choice between male and female moss VOC samples.

Then, they set up a series of ‘microcosms’ – miniature ecosystems containing moss plants, and where the presence of water and springtails could be manipulated. This time the research team used both C.purpureus and another moss species, Bryum argenteum, in which earlier work had shown that springtails were implicated in spreading sperm around. Some of their microcosms had only the mosses. Others were sprayed with water but had no springtails, or had springtails but no water spray. And some had both springtails and water. The results were fascinating.

When a female moss plant’s egg is fertilised, the resultant zygote grows into a thin brown stalk with a capsule of spores on top: this structure is called a sporophyte. Unsurprisingly, mosses in the absence of both water and springtails produced very few sporophytes indeed. Both the ‘springtail treatment’ and spraying the mosses with water caused a marked increase in fertilisation, as measured by the number of sporophytes produced. But combining springtails and the water treatment saw the number of sporophytes more than double, compared to each treatment on its own. The researchers commented that

[t]hese results highlight the substantial role of microarthropods in facilitating fertilisation in mosses, presumably through enhanced sperm transport.

So maybe we really are looking at something akin to the relationship between flowering plants and their pollinators. And, given the potential antiquity of this arrangement,

it is important to consider the potential role that a plant-pollinator-like relationship may have had in shaping the evolutionary ecology of moss mating systems.

I will definitely be changing the ‘additional reading’ list for my first-years!

 

T.N.Rosenstiel, E.E.Shortlidge, A.N.Melnychenko, J.F.Pankow & S.M. Eppley (2012) Sex-specific volatile compounts influence microarthropod-mediated fertilisation of moss. Nature published on-line 18 July 2012, doi: 10.1038/nature11330

first supertrees – now super domes Alison Campbell Jul 19

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singapore conservatory dome.jpgAfter goggling (a mixture of gobsmacked & ogling) the supertrees, our little party of escapees from the day’s official IBO program made our way into the Flower Dome, the first of the two great conservatories in Singapore’s Gardens in the Bay. Cue more ‘oh, wow!’ moments as the scale of the building became apparent – this is what it looks like once you’re through the doors (& into the wondrous coolness of the huge space):

singapore - entrance to flower dome.jpg

This dome contains gardens, or garden collections, from around the world, including the displays of flowers that give it its name. I was fascinated by the visual juxtaposition of the gardens with the almost futuristic cityscape beyond the conservatory walls.

singapore - flower dome & cityscape.jpg

singapore - flower dome hibiscus.jpg

From some perspectives the dome’s interior gives the impression of being heavily forested, & it’s at this point I had to keep reminding myself that none of this was here even 4 years ago: all the mature trees were brought onto the site from elsewhere…

singapore - flower dome forest look.jpg

… including a 1000-year-old olive tree. We could only guess at the huge amount of work (by goodness knows how many gardeners) to get all these plants established.

singapore - flower dome with olive tree.jpg

There’s also a wonderful collection of xerophytes: plants adapted to life in a dry environment. The plants in the following photo show a range of interesting adaptations related to this lifestyle.

singapore - xerophytes.jpgAnd scattered through the dome is a range of artwork, including this lovely botanically-based eagle, developed from the roots of a tree. (I am always amazed at how some people are able to ‘see’ the form within something, and work to bring it forth.)

singapore - flower dome eagle.jpg

 

in the lecture theatre – but definitely not giving a lecture! Alison Campbell Apr 02

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This is a post I first wrote for Talking Teaching – but hey! it’s about teaching science!

Today’s class was a real experiment for me, & although I try lots of different things in my classes, it was also a step outside my normal comfort zone. (But hey! life would be a bit boring if we always stayed safely inside that zone!) Why? Because I put into practice an idea I stole from my friend & colleague Kevin Gould (who also very kindly let me use the resources he’d developed): today was “design-a-plant” day, & probably to anyone looking into the lecture theatre during the first 30 minutes or so it would have looked as if chaos definitely ruled.

Last Friday I gave everyone an information sheet: descriptions of the features of leaf, stem & root that you might see in plants adapted to different environments. Today I trotted off to the lecture room with a box full of overhead transparency sheets, overhead pens, & printed scenarios (descriptions of a particular environment). The lecture theatre was already full — everyone had come ahead of time! This definitely wasn’t usual (it’s not that they normally trickle in late, but we’re talking seriouslyearly); obviously they were expecting something special. Gulp.

So I put up these slides:

then once they’d sorted out their groups I dished out pens, transparencies, scenario sheets (& copies of the info sheet for those who’d forgotten them), & away we went on a mutual journey of discovery. After all, this wasn’t myidea & I had no idea how it would really work out.

Well! The class erupted into happy, productive noise. I know it was productive because while they talked, argued, explained & persuaded, I circulated, listened in, & answered the occasional question. Those with computers had them open – looking up information related to their scenario. (Next time someone asks a question that I can’t answer on the spot, I’m jolly well going to get someone else to google it for me!) They drew, & altered their drawings, & drew some more. The original 20 minutes stretched towards 30, & still they were focused on what they were doing. I was almost sorry to interrupt :-)

Then, I called for volunteers. A hand went up almost immediately, & its owner came down to the overhead projector, not looking too nervous. She picked up the microphone, described her group’s scenario, & showed – & explained – their response. The next speakers followed just as quickly, and each speaker received a round of applause as they finished.

But the proof’s in the pudding – just what sort of plant had they designed? Well, they didn’t necessarily look like plants that my botanical colleagues could have put a name to, but nonetheless, the explanations each group gave for their particular design were sound, & science-based. They’d obviously taken on board not only the info on that fact sheet, but also the material we’d been looking at in lectures & that they’d found on line. And they’d had fun doing it. (I particularly liked the Nepalese Death Vine – the eerie noise of the wind passing through its herbivore-deterring spines apparently puts the locals off harvesting it, lol – and the Serengeti “cactus” that traps water in basin-like leaves, but when there’s a fire the plant’s transpirative water loss is such that its tissues become flaccid and it wilts, spilling that water onto the ground where the dampness keeps the worst of the fire at bay.) Plus – so far, the feedback for this exercise on our Moodle page is all positive: students felt it definitely helped their learning about plants.

Thanks, Kevin — your design-a-plant lesson got an A+ from all of us today!

skulls & braaiiinz – what’s not to like? (also, plants) Alison Campbell Jan 03

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The intrepid reporters from Number 8 Network e-mailed the other day. “What are you reading?” they asked; “after all, it’s the holidays & you must have heaps of time to put your nose in a book.” Which is sort of right, it is the Christmas/New Year break, but the days just seem to fly by when you’re doing not very much at all.

However, as it happens I’m working my way through several books at the moment, so I was able to oblige.

First up is Skulls, by Simon Winchester. Strictly speaking it’s not actually a book but an interactive iPad app, based on the enormous personal collection of Alan Dudley. I bought it because I find skulls fascinating (though not so obsessively fascinating as I think they must be for a collector of same) & the blurb at the app store offered me the ability to zoom in, out & around a whole bunch of bony brain protectors. This, I figured, would be quite fun & could also be a useful teaching tool (I’m looking forward to showing it to a colleague who teaches 3rd-year zoology).

And the ability to examine skulls in such detail really is great, although – a minor quibble! – I would have liked to be able to look at them from beneath & above as well as from all sides. You lose some definition at high levels of zoom but apart from that, wow! I would otherwise not have known that the Atlantic wolffish has quite so many teeth (shades of an aquatic Red Riding Hood villain) & such a wonderfully architectural skull. Or that a rabbit could somehow have lived long enough for its incisors to grow backwards & inwards in curves reminiscent of reversed (& miniature) elephant tusks.

The chapters comprising the ‘book’ struck me as a bit of an eclectic mix: we’ve got an interview with the collector himself, followed by an essay on the nature of collecting. Then, after learning about what a skull actually is & the bones that make it up, we find out about the dodo, or the pseudoscience relating to skulls – wherein we can learn about Piltdown man & phrenology but not, to my surprise, the various claims about ‘alien skulls’ from South America. Or the iconography of skulls, or skulls in art & in Mexico, & so on. One of the nice things about apps like this is that – even less than with a ‘real’ book – you don’t have to follow a linear progression through the text but can dip in & out, & that there are nice little visual cues to guide you in your choice of where to go next.

Not a lot in the text that was completely new to me, but well worth the price (at $17.99 this is one of the more expensive iPad apps) for the sheer enjoyment to be gained from viewing the images.

As for the brains – I’m also working my way through a Kindle edition of Carl Zimmer’s Brain Cuttings. Zimmer is one of my favourite science writers & this collection of essays (mostly written originally for Discover magazine) hasn’t disappointed me. The first essay’s title is from a question asked by Charles Darwin of one of his many correspondents. Wondering whether people around the world expressed emotions in the same way, Darwin asked, “Does shame excite a blush?” From this starting point, Zimmer takes us through scientists’ current understanding of the evolution of the face, a feature that began to form around half a billion years ago with the appearance of the earliest fishes. He asks why primates, in particular, have such complex, expressive faces – something that has to do with the complex social behaviour of this group of mammals. It turns out that facial mimicry is part of that social behaviour – and apparently the ability to mimic someone else’s facial expression, however briefly, may well be important in allowing us to understand how that person is feeling. An interesting experiment certainly suggests this:

[Researchers] had volunteers bite down on a pen and then look at a series of faces. They had to pick the emotion they thought the faces were expressing. The volunteers could recognise sad faces and angry ones with teh same accuracy as test subjects who did not have pens in their mouths. But they did a worse job of recognising happy faces.

Biting a pen, it just so happens, requires you to use the same muscles you use to smile. Because the smiling muscles were active throughout the experiment, [the research] subjects apparently couldn’t reel themselves start to mimic happy faces. Without that feedback, they had a more difficult time recognising when people were happy.

And that’s just the first chapter! The second essay, “The googled mind”, is an exploration of just where the mind stops. As Zimmer remarks,

[we] tend to think of the mind as separated from the world. We imagine information trickling into our senses and reaching our isolated minds, which then turn that information into a detailed picture of reality… In fact, teh mind appears to be adapted for reaching out from our heads and making the world, including our machines, an extension of itself.

In other words, it looks like the ‘mind’ is more of a complex system that comprises both the brain and various bits of its environment – books, for example, or computers, iPads – and even the tools we use.

This book’s both entertaining & informative & I’m enjoying reading it, one chapter at a time.

Third on my list is an actual, print-on-paper, hardcopy book: Fifty plants that changed the course of history, by Bill Laws. I bought this on a colleague’s recommendation, because of all the topics I teach at first-year level, botany seems to be the one that students are least engaged with, & I was hoping for some nice new examples to add my list of ‘cool stuff about plants & how they changed our world’.

My impression of this book is that it’s a bit like the curate’s egg: good in parts. My copy is a beautifully presented hard-cover edition, with lovely illustrations & some fascinating snippets of information (for example, that pineapples were grown in England over pits full of fresh dung! (This generated heat as it rotted, & augmented the warmth from stoves.) And the idea of a bamboo bicycle is an intriguing one. I’m enjoying dipping into it, a couple of plants an evening.

But unfortunately that enjoyment is tempered by moments of irritation. OK, I know I’m simply being greedy in wanting to know more about some topics than can be fitted into the 2 or 3 pages accorded them here. That’s a minor one. But saying that plants “absorb carbon dioxide and exhale oxygen” glosses over the fact that plants need that oxygen for cellular respiration as much as we do. (All too many of my first-years share this particular misconception.) And what am I to make of the following statement ?

The oldest names for the coconut are in Sanskrit, pointing to India as the source [of this plant]. However, the discovery of the fossilised remains of a tiny proto-coconut on New Zealand’s North Island suggests it might have been first taken into service here 5,000 years ago.

Five thousand years ago, there weren’t any people in New Zealand; hence no-one to take anything ‘into service’… (Interestingly, Sir Charles Fleming notes that a fossil coconut dating to the middle-late Miocene was discovered in Hawkes Bay; this would give an age of around 5-10 million years.)

So, I’m accumulating some new stories to tell in class, and I am enjoying the read, but – unlike Skulls and Brain Cuttings – I probably won’t be recommending Laws’ book to students looking for a bit of extra reading. At least, not until I’ve finished it & identified any other potential pitfalls. Although… I guess the error I’ve picked up on here would be a useful jumping-off point for a discussion of New Zealand’s botanical history.

B.Laws (2010) Fifty plants that changed the course of history. Firefly Books. ISBN: 978-1-742372-18-1

S. Winchester (2011) Skulls. Touch Press.

C.Zimmer (2010) Brain Cuttings: fifteen journeys through the mind. Scott & Nix Inc. NY ISBN (Kindle): 978-1-935622-16-1

biological oddities, including the naughty bits Alison Campbell Aug 05

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Last night I gave a talk up in Auckland, on various biological oddities (mostly from the animal kingdom and, all right, mostly to do with s*x). You can slip a lot of serious science in once the audience’s attention has been captured by the naughty bits! (I would hate folks to think that biologists are totally obsessed with s*x. This is not true. But related stories do tend to focus the attention.)

Anyway, I was chatting about it with some of our grad students this morning and they said, oooh, we wouldn’t might reading more about that. Various people (including me & Grant) have blogged them all before, so I’ll bring all the links together in one place but won’t fill in too many of the gaps.

First up, zombies! More particularly, the use of the z-word to capture public attention & direct it to a serious subject: modelling (& more recently, how to deal with) the spread of infectious disease. The outcome of the modelling work was brought to the world’s attention by a paper with the eye-catching title, When zombies attack. More recent advice on getting through an infectious disease outbreak – things like stocking up on food & water & staying home – was presented by the US Centres for Disease Control under the heading: Preparedness 101: Zombie Apocalypse. (The daughter, who pays attention to such things, felt the advice was sadly lacking in that it doesn’t actually say anything about how to dispose of the zombies who might actually get into your house. Practicalities, people!)

This naturally segued into the tale about zombie ants – zombified by a parasitic fungus. Parasites can have quite marked effects on their hosts’ behaviour – changes that maximise the reproductive success of the parasite. I first got interested in this topic years ago, when I read Carl Zimmer’s excellent book, Parasite Rex. In the case of the ants (Camponotus leonardi), infection with the fungus Ophiocordyceps unilateralis causes the ants to leave their usual habitats, hang upside down off leaves or stems, bite on to the plant – & die. Then when the fungus produces a fruiting body it can rain down spores onto the heads of unsuspecting ants passing underneath.

Then we moved on to the slightly risque stuff, beginning with the interesting observation that female crayfish release urine during courtship. This influences the males’ behaviour & allows the females to assess the quality of their suitors. The original report includes a link to a video – using fluorescent dyes allowed the researchers to visualise the timing of urine release & so relate this to the more obvious behaviour patterns displayed by their subjects.

Couldn’t leave out the tree shrews living in montane forests in Borneo, who use the ‘pitchers’ of some pitcher plant species as toilets. This is quite a cool example of coevolution, where Nepenthes lowii plants gain up to 100% of their nitrogen requirements from shrew faeces, while Tupaia montana (the shrew) gains sugars from licking the plant’s nectaries, enticingly displayed on the inside of the pitcher’s lid. (Well, enticing if you’re a shrew…)

Also in the forest, we have fruit bats. As Ed Yong describes, in one species of fruit bat, the duration of copulation is affected by whether or not, & how long, the female licks the male’s penis during copulation! Presumably this would have an impact on mating success. (In empid flies, for example, duration of mating is affected by the size of the food gifts that males bring for females, & longer copulations tend to produce more offspring.)

On the other hand, duration of copulation would have no impact at all on breeding success in the sole recorded example of homos*xual necrophilia, involving two mallard drakes (one of them very very dead). Not that this stopped the living drake from mating vigorously with the corpse – for 75 minutes!!

Mallard drakes are randy little beggars, with their activity extending to forced copulations with hapless females. This is usually later in the season & often involves multiple drakes, & can be so physical & prolonged that the females may drown. This promiscuous behaviour in waterfowl has a morphological correlate. Males of highly promiscuous species, where there are high levels of sperm competition, have long & tightly coiled penises (matched by long & tightly coiled vaginas in the females). At the other end of the spectrum are the monogamous species like black swans, who are much less-well endowed in the genital department. Females in the promiscuous species are able to control who they mate with by contracting or relaxing muscles that allow them to shorten the vagina, so that in a forced copulation the male may not actually be successful in passing on his genes, as his sperm may not be deposited high enough in the female’s reproductive tract.  Fascinating stuff – & caught on film (again, hat tip to Ed Yong).

As you may imagine, the discussion after last night’s talk was extremely animated :-)

pesky little hoppers Alison Campbell Jan 09

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With the new house came a long drive lined with agapanthus. My mother would have said, “the dreaded agapanthus”, & she wouldn’t have been far wrong. I don’t like the things very much; they spread very vigorously & I tend to view them as a weed. (I see from Te Ara that Biosecurity New Zealand was looking at calling for a nationwide ban on the plants, back in 2007. I wonder what happened with that? Where we live now, every second house has agapanthus in the garden.) Still, we haven’t really given any thought to what we might replace them with, so the agapanthuses (agapanthi?) have had a reprieve for the moment. And this means that I have to cut back all the spent flowerheads – a bit before they’ve finished flowering, actually, so as to minimise the chances of them setting (& spreading) seed.

This is a bit of a back-breaking job, as it happens, bending down to cut the stems off low. (The Significant Other suggested going hell-for-leather with the pruning shears but that would make an awful mess.) But it’s given me a good view of what’s living in the leaf clumps – & as far as I can tell, it’s mostly passion-vine hoppers (Scolypopa australis. The pesky little things keep flying up & landing on the inside of my glasses & it’s a bit unnerving, I can tell you, to have blurry brown things creeping round in front of your eyes! (For those not familiar with these little beasties, there are some lovely images by Phil Bendle here.)

The adults are about 10mm long, with dark grey-brown bodies & transparent wings supported by a lattice of dark brown veins. The nymphs (juvenile form) are quite different, with creamy white bodies marked with brown spots, and the most wonderful collection of bristles sticking out of their rear ends – reminiscent of an arthropodan shaving brush. Both nymphs & adults feed by inserting long pointed mouthparts (rather like hypodermic needles) into the phloem of plant stems, an efficient if rather vampiric way of getting nutrients as they don’t even have to suck: because phloem is moved around the plant under positive pressure, the animals just have to sit there & breakfast, lunch & dinner simply flow into their guts.

This is quite a cool trick, actually, because the layer of phloem tissue in a stem is very thin, just a few cells thick, & before modern technology came along it was very difficult for an enquiring botanist to insert a needle into the phloem for purposes of measuring flow & nutrient content. This problem was overcome by using aphids, which are also little plant-suckers. Once an aphid had got its mouthparts nicely into the phloem, the scientists wouldcut the insect’s body off just behind its head. The disembodied head remained attached to the plant by its embedded mouthparts, & phloem sap would continue to flow through it for several days.

Because aphids, hoppers & the like are tapping directly into a plant’s nutrient transport system, a heavy infestation can be quite debilitating for the plant. (Not that this seems to be the case for our dreaded agapanthi; their growth is revoltingly vigorous.) The female hoppers cut little slits in the stem in which to lay their eggs, thus creating an opening for infection which can result in die-off of the affected parts. And, as with human drug addicts who share needles, as the hoppers move from plant to plant they can spread disease from one feeding station to the next. In other things, they’re not something you want in large numbers in your garden.

I suspect it’s a vain hope that they’ll stay on the agapanthuses & leave our nice new passionfruit vines alone…

_________________________________________________________________________________

In other news: the goldfish must like their new pond :) There was a lot of piscine hanky-panky going on when I went out to feed them this morning.

stunning biological image #2 Alison Campbell Nov 26

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I teach a bit of botany, to our first-year students. I really enjoy the subject & hopefully some of that rubs off :) Anyway, I’m always on the lookout for new images to use in my lectures, & tonight I came across this stunning photograph by Eckhard Völcker, who has very kindly given his permission for me to share the image with you.  

While it looks like a lovely piece of lace in contrasting colours, it’s actually a section through a willow shoot, stained with dyes that (for example) highlight the lignified tissues in xylem and the fibre caps on vascular bundles.

You can see more of these beautiful photos here, or visit Herr Völcker’s webpage for more of his work.

(Your turn, Grant!)

yes, we have some bananas Alison Campbell Oct 29

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Prominent creationist Ray Comfort once (in)famously commented that the ‘design elements’ that make up a banana, including its so-convenient shape, are evidence for the existence of a Designer. A comment that has been pretty resoundingly debunked – unsurprisingly, since the banana-as-we-know-it is due in large part to the hand of man, selecting for those features of bananas that make them desirable as a food – lack of seeds (wild-type, uncultivated bananas have almost more seeds than flesh) & that wonderfully unzippable peel. Something that last year’s Schol Bio examination asked students to think about.  

Inside a wild-type banana

Image: inside a wild-type banana (from Wikipedia)

Bananas belong to the genus Musa. If you think back to the last banana you ate, you’ll rememember that it’s seedless, unlike the ‘wild’ banana shown above. All commercially-grown banana plants are produced asexually, from suckers or sprouts. The varieties we import also tend to be rather large, but the fruit can be much smaller (apparently the name ‘banana’ derives from an Arabic word for finger, banan). They’re imported as unripe fruit, & ripened before sale by being exposed to ethylene. Ripe fruit (not just bananas) release this gas, but for commercial ripening the fruit are placed in a room that’s then flooded with it. 

Incidentally that lack of any sort of sex life places banana crops at some risk: because they’re clonal, if a pathogen comes along to which they have no resistance, much of the crop may fail. For example, the ‘Black Sigatoka fungus’ can lower production by up to 50% in infected crops. This problem is magnified by the loss of genetic diversity in wild bananas as rainforests are felled. Another pathogen, bacterial banana Xanthomonas wilt (BXW) disease, poses a significant threat in East Africa. While we may view the loss of bananas as something of a misfortune, for others it would be much worse: after rice, maize & wheat, bananas are the world’s 4th most important food crop.

Now, back to the question. The examiner tells us that “Many different species of banana exist today, all of which are descended from one or other of the ‘wild’ Asian species: Musa acuminata (AA:2n = 22) and Musa balbisiana (BB: 2n = 22). Students needed to pay careful attention to the AA & BB. They represent the genome – the full chromosome set – rather than alleles at a particular locus. This is important, because the question goes on to give you information about the genomes of several of the major cultivars in modern bananas:

Species Genome

 

Cultivars

 

AA

 

Sucrier

 

Jari Buaya

 

AAA

 

Gros Michel

 

Grande Naine

 

Cavendish

 

BB

 

Abuhon

 

Chuoi Hot Qua Lep

 

AB

 

Njalipoovan

 

ABB

 

Awak

 

Pelipita

 

AABB

 

Kluai Ngoen

 

So, the examiner asked scholarship candidates to ”[d]iscuss the sequence of events and processes that have resulted in the three different species of banana with the followign genomes, arising from the original ‘wild’ species of banana: AAA, AB, and ABB,” and advised using annotated flow diagrams in doing this. The discussion also needed to include ”the genetic processes that could have occurred to produce the different cultivars of Gros Michel, Grande Naine and Cavendish within the one species of the AAA genome.” (The question went on to ask candidates to design an experiment to investigate the action of ethylene on ripening rate in bananas, but I think we have enough to look at without going into that right now!)

O-kay. I am not one of those clever people who can do all sorts of flowcharts on their webpages, but if I was, I’d be drawing a flow diagram or two showing things like polyploidy & hybridisation. This is because that first AAA genome must be the result of a cross between 2 M.acuminata individuals where one produced normal (i.e. n = 11) haploid gametes, A, while the other produced gametes (at least some!) which were diploid (AA) due to complete non-disjunction in meiosis. The 3n (AAA) individuals thus produced are autopolyploids (both parents from the same species. Because as triploid organisms you’d predict they’d have difficulty producing gametes of their own, these AAA plants are sterile & can reproduce only asexually.

The second (AB) species is the result of a hybridisation event involving one M.acuminata parent and one from M.balbisiana, both producing haploid gametes. And tthe third, ABB, species is another polyploid organism. This time it’s an allopolyploid (2 different parents) with the M.acuminata parent contributing a haploid gamete (A: n = 11) while the M.balbisiana parent must have produced some BB (2n = 22) gametes, again through complete non-disjunction during meiosis. (You could probably get away with not drawing flow diagrams, but in that case you’d need to give a comprehensive & logical description of what was happening.)

What about those different cultivars with the one genotype (AAA)? Two possibilities here. One is mutations after the AAA triploid was formed. In some cases mutations – in different plants – could result in different phenotypes. And if those phenotypes were viewed as ‘desirable’ by farmers, they’d be propagated by cuttings, ending up as the different cultivars.

The other possibility is that the original parent plants (those AA individuals) had differences in their genotypes. This is really only to be expected, given the fact that independent assortment, crossing-over & recombination routinely shuffle alleles between homologous chromosomes, and homologues assort independently when gametes are formed. Again, when the AAA polyploids were produced (I think it’s safe to assume this could happen more than once) then they’d receive these variations from their parents. And again, any ‘desirable’ phenotypes would be selected for.

There! Isn’t that a rather more satisfying explanation than Mr Comfort’s?

northern rata – first it’s an epiphyte, then it’s not Alison Campbell Aug 03

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Last year’s Level 3 paper on ‘plant responses & animal behaviour’ (AS 90716) had a question on northern rata - rather a lovely tree; I remember that we had one on our section back in Wairoa, when I was a kid. For some reason that tree & the big totara next to it had been left when the rest of the section was cleared.. Anyway, this question began (as all the questions do) with a bit of contextual information:

Northern rata (Metrosideros robusta) is found in lowland forest throughout the North Island and near the northwest coast of the South Island. It is much more common as an epiphyte than a ground plant, and is mostly found growing on established tress such as the rimu (Dacrydium cupressinum). Both the rata and its host require plenty of light.

The rata epiphyte develops tuber-like swellings on its roots, which help with water storage.

Eventually roots from the epiphyte grow down the trunk of the supporting tree to the ground, forming a massive trunk and root system. This system replaces and eventually kills the supporting tree.

This is a really interesting relationship. When it’s young, the rata gains considerable benefits from its epiphytic lifestyle. It’s up at canopy level (not down on the ground like most seedlings) & as a result is exposed to much higher light levels than would be experienced on the forest floor. It doesn’t expend a lot of energy in growing taller, but can put that energy into producing leaves instead. (Those water-swollen roots mentioned in the original context statement are an adaptation to this lifestyle.) At this point what we’re looking at is a commensal relationship: the rata gains from its perch high in the canopy, while the rimu may not gain from it but it’s not significantly harmed, either.

But eventually things turn nasty – for the rimu, anyway. Over time there’s a structural change in the epiphyte ( is this just due to size? or age? – hopefully a better botanist than me can enlighten all of us!). Once its roots hit the ground, & it can tap into a constant groundwater supply, the rata is on the way to becoming an independent canopy tree with a hollow trunk formed by the fusion of its ground-seeking roots (just like the strangler figs we saw in Queensland last year). Ultimately the rimu dies – & rather than its death opening up a space for other seedlings to grow, it’s replaced by the rata. This fate isn’t something where natural selection might see the rimu population adapt, because the rimu is already a reproductively mature plant before the rata takes it out – it’s already reproduced & passed on its genes. Normally, as I’ve said, the open, sunlit space left by a fallen rimu would quickly see the growth of seedlings, including those of the rimu. But with the rata already a tall canopy tree (they can grow up to 25m tall), there is no sunlit opening in the forest, & so germination & growth of other seeds must wait until the rata itself comes crashing down.

However, all is not rosy for the rata – in many areas they are under severe browsing pressure from possums. Apparently it can take as little as 3 years to kill a mature tree. When a lot of trees are being hit quickly, this opens up the canopy on a fairly large scale, wih the potential for future damage due to windthrow in storms, and attack by insects & disease organisms. At the same time, it’s getting more expensive to control possums, something that’s compounded by growing public opposition to the use of 1080 for this purpose. Yet without some efficient, cost-effective means of controlling these furry pests, we stand to lose this beautiful tree, & many more besides.

azolla & endosymbiosis Alison Campbell Aug 01

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ResearchBlogging.org

There are other photosynthesisers besides Volvox, living in our fishpond. Bigger plants include waterlilies, various sedges, & Elodea. And at this time of year the surface is covered by a carpet of duckweed, but when summer comes the Azolla will tend to take over. Sometimes called ‘water fern’, Azolla contains an endosymbiont, a cyanobacterium (blue-green alga) that lives within the plant’s ‘body’ but not within its cells (Ran et al. 2010).

This cyanobacterium is actually a fairly recent fellow-traveller for the fern. Almost all eukaryotes (with the exception of Archaezoans like Giardia) contain intracellular endosymbionts – the mitochondria. These & the chloroplasts of plants formed as the result of endosymbiotic events that occurred perhaps two billion years ago, when the free-living ancestors of these organelles were engulfed by other prokaryote cells but for some reason weren’t digested. Instead, they continued to do their thing, churning out sugars (in the case of the proto-chloroplasts) & ATP well in excess of what the ‘host’ could generate alone (mitochondria). Lyn Margulis developed this endosymbiotic theory for the origins of mitochondria & chloroplasts on the basis of a range of observiations: both organelles contain their own, circular, DNA & (just like bacteria) are able to manufacture their own proteins; their ribosomes & tRNA molecules are like what you’d find in bacteria; and they’re enclosed in a double membrane. Interestingly, many of the genes that would once have been on that circular chromosome of a mitochondrion or chloroplast have ended up in the ‘host’ cell’s nucleus – the host can to some degree control the organelles’ functioning. (Ran et al. note that the chloroplast genome is one of the smallest known, at only 150-200,000 base pairs long.)

Ran & his colleagues were keen to delve further into the process of endosymbiosis as it relates to chloroplasts in plants. To do this, they chose to study a cyanobacterium living inside a species of Azolla – inside, but outside the actual Azolla cells, tucked away into litle ‘cavities’ in cells on the water fern’s upper surface. While there are other symbioses between plants & cyanobacteria, this one’s unusual on two counts: the cyanobacterium involved can’t grow outside the host, & it’s passed on from one generation of Azolla to the other (‘vertical transmission’). The oldest fossils of Azolla date back 140 million years, & it’s possible that this endosymbiotic relationship goes back that far in time, The team hypothesised that “… genome reduction may… act on cyanobacteria in symbiosis with plants”, mirroring what appears to have happened in the evolution of chloroplasts.

Figure 1 from Ran et al. (2010): A) fronds of Azolla filiculoides;. B) Close up of the upper surface of an Azolla branch. C) Light micrograph of the cyanobiont. The larger cells represent nitrogen-fixing heterocysts. Scale bar = 5 µm. D) Transmission electron micrograph of the cyanobiont. E) A snap-shot in the vertical transmission process of the cyanobiont between Azolla plant generations, using fluorescence microscopy. Pairs of megasporocarps (blue) develop at the underside of the cyanobacterial colonized Azolla leaves. Filaments of the motile cyanobacterial cell stage (red), the hormogonia (h), are attracted to the sporocarps, gather at the base and subsequently move towards the tip, before entering the sporocarps via channels (white arrows). Once inside the sporocarp the hormogonia differentiate into individual thick walled resting spores (or akinetes; ak), seen as the intensively red fluorescing small inoculum on top of the megaspores (sp).

The team sequenced the cyanobacterium’s genome – and found it to be ‘eroding’. Thirty-one percent of its genes are pseudogenes (they’re either not transcribed, or they don’t produce functional proteins), & there are a lot of transposons – ‘jumping genes’ - in the genome. Significantly, some of the genes that are essential for a free-living bacterium have been ‘pseudogenised’, which means that the cyanobacterium must be dependent on the Azolla for things like DNA repair proteins. The ‘DNA replication initiator’ gene is also pseudogenised,  which is important as it means that Nostoc azollae can divide & grow only very slowly. And the same is true for genes involved in glycolysis and taking nutrients into the cell.

On the other hand, the cyanobacterial genes involved in nitrogen fixation are still functioning well ie the cyanobacterium is able to differentiate to produce heterocysts, where nitrogen-fixation occures, meaning that the symbiont is a key source of nitrogen for its plant host. There’d be quite strong selection pressure for continuing this endosymbiotic relationship: nitrogen is a limiting factor for plant growth, and the host (the Azolla) would gain a big selective advantage over competitors that lacked N-fixing endosymbionts.

The team conclude that over time, the Nostoc azollae genome will erode to the point that ‘ulimately may cause NoAz to resemble a plant organelle (devoted to nigtrogen fixation) more than a free-living organism.’  They also point out that for such organelles to evolve, there would initially have to have been some form of vertical transmission process (seen in this example) & eventually  the symbiont would become intracellular.

So perhaps, in the relationship between Azolla filiculoides and its cyanobacterial partner, we are looking at the evolution of a fully endosymbiotic relationship.

L.Ran, J.Larsson, T.Vigil-Stenman, J.A.A.Nylander, K.Ininbergs, W-W.Zheng, A.Lapidus, S.Lowry, R.Haselkorn & B.Bergman (2010). Genome erosion in a nitrogen-fixing vertically transmitted endosymbiotic multicellular cyanobacterium PLoS ONE, 5 (7) : 10.1371/journal.pone.0011486

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