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How do trees transport water such long distances? Part 2: the mechanism remains a mystery (to me)

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and I still haven’t found what I’m looking for…

So scientists have learned a lot, though not everything, about water’s travels from soil to leaf in a plant or tree. It’s a fascinating story, and I’m keen to learn more. But the real mystery for me is about energy. As the excellent Nature article, upon which I’m mostly relying, points out, animals have a pump-based circulatory system to distribute nutrients, oxygen and so forth, but plants are another matter, or another form of organised matter.

I actually posed two questions in my last post. How do plants – and I think I should specify trees here, because the massive distance between the soil and their top leaves makes the problem more dramatic – move water such large distances, and how do they know they have to transport that water and how much water to transport?

So let’s look at the Nature Education explanation:

The bulk of water absorbed and transported through plants is moved by negative pressure generated by the evaporation of water from the leaves (i.e., transpiration) — this process is commonly referred to as the Cohesion-Tension (C-T) mechanism. This system is able to function because water is “cohesive” — it sticks to itself through forces generated by hydrogen bonding. These hydrogen bonds allow water columns in the plant to sustain substantial tension (up to 30 MPa when water is contained in the minute capillaries found in plants), and helps explain how water can be transported to tree canopies 100 m above the soil surface.

Notice how we’re again returning to the explanations questioned by Wohlleben – transpiration and capillary action. But we’re introduced to something new – the C-T mechanism. The thesis is that water’s cohesiveness through hydrogen bonding creates a tension (the tension that makes for capillary action) that enables water to be shifted up to 100 metres – all because of the minuteness of capillaries found in plants. And trees? Somehow, I just can’t see it. Perhaps the key is in the phrase ‘helps explain’.  There must surely be more to this. The thesis also mentions ‘negative pressure’ generated by transpiration. This is the signalling I wrote about before. Somehow the plant’s chemistry recognises that there’s an imbalance, and of course this happens in all living things, regardless whether they have a complex nervous system. So maybe there’s no need to worry about ‘knowing’. All living organisms respond to their ever-changing environment by altering their internal chemistry, by opening or closing barriers, by selectively adding or subtracting nutrients, and there are unknowns everywhere about precisely how they do that. It’s a kind of organised chemistry that seems like everyday magic from the outside, whether we’re focusing on a beech tree or our own intestines.

The C-T mechanism is only new to me I should add. It can actually be traced back to 1727 and a book by Stephen Hales, in which he pointed out that without what he called perspiration the water in a plant would stagnate, and that it was also required to allow for the capillary movement of water, because ‘the sap-vessels are so curiously adapted by their exceeding fineness, to raise [water] to great heights, in a reciprocal proportion to their very minute diameters’. But this ‘reciprocal proportion’, according to Wohlleben, as quoted in the last post, can only account for a maximum of 3 feet of upward force in ‘even the narrowest of vessels’.

The water transport system, referred to in the last post as the water potential difference or gradient, also has another name, the Soil Plant Atmosphere Continuum (SPAC). I also mentioned something about an ‘apoplastic pathway’. Water enters the tree by the roots, which are divided and subdivided much like branches and twigs above-ground, with the thinnest examples being the fine root hairs. Water enters through the semi-permeable cell walls by osmosis. Cell-to-cell osmosis carries the water deeper into the root system, and thence into an apoplastic pathway. According to this video, this pathway provides an uninterrupted flow of water (no cell wall barriers) which allows a mass flow ‘due to the adhesive and cohesive properties of water’. This is the cohesion-tension theory again. Apparently, due to evaporation, a tension is created in the apoplast’s continuous stream, leading to this ‘mass flow’.

This makes absolutely no sense to me. What I’m so far discovering is that it’s pretty hard to start from scratch as an amateur/dilettante and get my head around all this stuff, and in my reading and video-watching I’ve yet to find a straightforward answer to the how of long distance, fast transport of water in plants/trees – there probably isn’t one.

I’ll try again after a diet of videos – so far I’ve found a large number of videos in Indian English, and their accents defeat me, I’m sad to say. No transcripts available. Meanwhile, I’ve compiled a little glossary (from various sources) to help myself…

apoplast – within plants, the space outside the plasma membrane within which material can diffuse freely. It is interrupted by the Casparian strip in roots, by air spaces between plant cells and by the plant cuticle.

Casparian stripa band of cell wall material deposited in the radial and transverse walls of the endodermis, which is chemically different from the rest of the cell wall – the cell wall being made of lignin and without suberin – whereas the Casparian strip is made of suberin and sometimes lignin.

cortical cells – in plants, cells of the cortex, the outer layer of the stem or root of a plant, bounded on either side by the epidermis (outer) and the endodermis (inner).

exudation – An exudate is a fluid emitted by an organism through pores or a wound, a process known as exuding.

guttation – water loss, when water or sap collects (at times of low evaporation, dawn & dusk), at tips of grass, herbs (not to be confused with dew, caused by condensation).

hydrostatic pressure – the pressure exerted by a fluid at equilibrium at a given point within the fluid, due to the force of gravity. This increases in proportion to depth measured from the surface because of the increasing weight of fluid exerting downward force from above.

lignin – a class of complex organic polymers that form important structural materials in the support tissues of vascular plants and some algae. Lignins are particularly important in the formation of cell walls, especially in wood and bark, because they lend rigidity and do not rot easily.

osmosis – the movement of water from an area of high to low concentration through a semi-permeable membrane. ‘Pumps’ in the cell membrane transport the specific ions into the cell which means water moves in by osmosis thus maintaining hydrostatic pressure.

phloem – the living tissue that transports the soluble organic compounds made during photosynthesis and known as photosynthates, in particular the sugar sucrose, to parts of the plant where needed. This transport process is called translocation.

plasmodesmata – narrow threads of cytoplasm that pass through the cell walls of adjacent plant cells and allow communication between them.

root pressure – the transverse osmotic pressure within the cells of a root system that causes sap to rise through a plant stem to the leaves. Root pressure occurs in the xylem of some vascular plants when the soil moisture level is high either at night or when transpiration is low during the day

sap – a fluid transported in xylem cells (vessel elements or tracheids) or phloem sieve tube elements of a plant. These cells transport water and nutrients throughout the plant.

suberin – an inert impermeable waxy substance present in the cell walls of corky tissues. Its main function is as a barrier to movement of water and solutes.

symplast – the network of cytoplasm of all cells interconnected by plasmodesmata. The movement of water occurs from one cell to another through plasmodesmata

tracheid – a type of water-conducting cell in the xylem which lacks perforations in the cell wall.

vascular (plants) – also known as tracheophytes and also higher plants, form a large group of plants (over 300,000 accepted known species) that are defined as those land plants that have lignified tissues (the xylem) for conducting water and minerals throughout the plant.

xylem – one of the two types of transport tissue in vascular plants, phloem being the other. The basic function of xylem is to transport water from roots to shoots and leaves, but it also transports some nutrients.

 

On the Trump’s downfall. What a memo. One wonders if the DoJ is running out of patience with the wannabe dictator and his imbecilities, which may bring things to a head sooner rather than later. But those in the know say that Mueller is always thorough and unlikely to be distracted, so I shouldn’t project my own impatience onto him. Dog give me strength to suffer the horrorshow for a while longer.

 

Written by stewart henderson

February 5, 2018 at 3:48 pm

Posted in biology, botany

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How do plants transport water? Part 1: xylem, transpiration and a mysterious water potential difference

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roots, xylem, upward flow, transpiration – but how does it work? Find out in the next thrilling episode, maybe.
Stolen from Nature Education, with apologies

This post could fit well in the ‘How Stuff Works’ series, always a useful resource, but I doubt if they’ve done a piece on today’s subject. Maybe I’ll check later.

I’ve been reading a book called The hidden life of trees, by Peter Wohlleben, a Chrissy present from a good friend. One of its shortest chapters is titled ‘The mysteries of moving water’. The reason for its brevity is essentially that there’s as yet no solution to the mystery of how water gets from the soil to the leaves of a tree, or any plant for that matter. At least, according to Wohlleben.

This strikes me as amazing, if true. After all, it’s a simple, everyday scenario for any home gardener. You notice on a hot summer day that the leaves of your capsicum plant are wilting. You apply a two-litre dose of H2O to the base, et voilà, within an hour or two (I don’t know, I’ve never timed it), those leaves have become as turgid as much of my writing. And it just may cross your mind that it’s pretty miraculous how plants can do that. But if it’s true that we don’t know how plants manage such an everyday miracle, surely working it out is Nobel Prizeworthy for any ambitious team of botanico-chemists out there, or whatever.

Of course it’s much more likely that botanists have been trying to solve this mystery for decades – isn’t it? But before I look into it, here’s what Wohlleben says in his book:

…water transport is a relatively simple phenomenon to research – simpler at any rate than investigating whether trees feel pain or how they communicate with one another – and because it appears so uninteresting and obvious, university professors have been offering simplistic explanations for decades… Here are the accepted answers: capillary action and transpiration.

Upon reading this I tried to recall what I knew of these terms. With capillary action I drew a blank, though I feel sure I knew about it once. Transpiration, though, was clear enough: it was like perspiration, the evaporation of water from the leaves, rather than the skin (or is perspiration the secretion of water through the pores rather than the evaporation? Later). So transpiration is only about the movement of water from the surface of a leaf to the atmosphere by means of solar energy; it surely has nothing to do with movement through the stem or trunk, though the loss of water from the leaves is presumably a signal to the plant to draw up more water from the earth, but how can we talk of signals when a plant has no brain or command centre to receive them? And how can water be ‘drawn up’ when it has no muscle power or other obvious energy source?

As to capillary action, Wohlleben explains:

Capillary action is what makes the surface of your coffee stand a few fractions of an inch higher than the edge of your cup. Without this force, the surface of the liquid would be completely flat. The narrower the vessel, the higher the liquid can rise against gravity. And the vessels that transport water in deciduous trees are very narrow indeed: they measure barely 0.02 inches across. Conifers restrict the diameter of their vessels even more, to 0.0008 inches. Narrow vessels, however, are not enough to explain how water reaches the crown of trees that are more than 300 feet tall. In even the narrowest of vessels, there is only enough force to account for a rise of 3 feet at most.

Needless to say, plenty of research has been done on the subject of water transport in plants, but I have to agree with Wohlleben that there’s a lot that’s missing. The key to the process is a material called xylem, a structure made from hollow, dead, reinforced cells. Here’s how a BBC science site tries to explain it:

Transpiration explains how water moves up the plant against gravity in tubes made of dead xylem cells without the use of a pump.

Water on the surface of spongy and palisade cells (inside the leaf) evaporates and then diffuses out of the leaf. This is called transpiration. More water is drawn out of the xylem cells inside the leaf to replace what’s lost.

As the xylem cells make a continuous tube from the leaf, down the stem to the roots, this acts like a drinking straw, producing a flow of water and dissolved minerals from roots to leaves.

Water doesn’t flow upwards, however. It has to be pumped up, or sucked, as we do when we apply our lips and energy to a straw. The BBC also describes the whole process as transpiration, which just seems wrong to me. Obviously much transpires here, but it isn’t just transpiration. What?

What obviously needs explaining is where the energy comes from to draw the water up against gravity, and how the plant ‘knows’ that water needs replenishing.

A more comprehensive, and richly referenced, attempt at an explanation is provided by Nature, the well-known science magazine, on one of its educational websites. There we’re told that ‘plants retain less than 5% of the water absorbed by roots for cell expansion and plant growth’. This is fascinating, as is the reason for the lack of retention – photosynthesis. Water is lost to the atmosphere from the leaves’ stomata, which are like our pores. These stomata are used to absorb CO2 for the photosynthesis of sugars, but their openness to CO2 increases the transpiration rate, so there’s a tricky balance between the two – water loss versus CO2 and sugar gain.

The xylem mentioned above doesn’t reach down all the way to the base of the root system. First the water must pass through several cell layers that act as a filtration system. But how does it do this? What is the force being applied and where does it come from? The Nature article gives this complex explanation:

The relative ease with which water moves through a part of the plant is expressed quantitatively using the following equation:

Flow = Δψ / R,

which is analogous to electron flow in an electrical circuit described by Ohm’s law equation:

i = V / R,

where R is the resistance, i is the current or flow of electrons, and V is the voltage. In the plant system, V is equivalent to the water potential difference driving flow (Δψ) and i is equivalent to the flow of water through/across a plant segment. Using these plant equivalents, the Ohm’s law analogy can be used to quantify the hydraulic conductance (i.e., the inverse of hydraulic R) of individual segments (i.e., roots, stems, leaves) or the whole plant (from soil to atmosphere).

Got that? I may be wrong, but isn’t this just an analogy? Don’t analogies tend to break down with a little bit of analytic pressure? The idea of hydraulic conductance is clearly drawn from electrical conductance, but electrical conductance relies on a power source, doesn’t it? What is the plant’s power source? Yes, I can see that certain parts of the plant have a greater resistance to the water’s mostly upward movement than others, and that this resistance is measurable by examining the time it takes for water to pass through the different parts with their particular structure and chemistry, but it says nothing about the energy source. In Ohm’s law, V, voltage is the amount of power, which comes from a source of that power, such as a battery. In the above analogy, Δψ is described as the water potential difference that drives flow. I’m possibly being dumb, but how does that happen? What’s meant by ‘water potential difference’?

The Nature article, I must say, is very good at telling us about the materials and obstacles negotiated by water molecules on their journey. First they pass through the root’s epidermis, then the cortex and the endodermis and then on to the xylem. They travel by an apoplastic pathway (more of that next time), or else a cell-to-cell pathway (C-C), and the role of ‘water-specific protein channels embedded in cell membranes (i.e., aquaporins)’ is mentioned, but this role is apparently still much of a mystery. Anyway, the xylem continues into the petiole, to which the leaves are attached, and then into the mid-rib, the main central vein of the leaf. From there the water passes into the smaller branching veins of a dicot leaf, which also contain tracheids – elongated xylem cells for the transport of water and mineral salts. It’s from this network of veins that transpiration takes place.

So I’m learning a lot, but the ‘water potential gradient’ and how it pulls or pushes water upwards, that’s still very much a mystery to me. But there’s more to come.

References

https://www.nature.com/scitable/knowledge/library/water-uptake-and-transport-in-vascular-plants-103016037

http://www.bbc.co.uk/schools/gcsebitesize/science/add_gateway_pre_2011/greenworld/planttransportrev1.shtml

Peter Wohlleben, The hidden life of trees, Collins 2017

 

Ok, the usual update on Trump’s downfall. Some are saying that the Mueller enquiry is winding up (and I’m not talking about GOP hardheads), but I’m hoping not, because I reckon the financial stuff alone will take years to wade through properly. In the meantime though, I’m hoping that more really dramatic developments occur to light a fire under Trump’s capacious backside, sooner rather than later. The latest news is that the Mueller team are looking at the cover-up re Trump Jr’s meeting with Russian agents. So maybe the cover-ups and the endless obstructing will lead to some justice action soon, while the ‘follow the money’ aspect will continue for some time, and hopefully do the really lasting and permanent damage to the Trump horrorshow.

 

Written by stewart henderson

February 1, 2018 at 11:13 pm

how evolution was proved to be true

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The origin of species is a natural phenomenon

Jean-Baptiste Lamarck

The origin of species is an object of inquiry

Charles Darwin

The origin of species is an object of experimental investigation

Hugo de Vries

(quoted in The Gene: an intimate history, by Siddhartha Mukherjee)

Gregor Mendel

I’ve recently read Siddhartha Mukherjee’s monumental book The Gene: an intimate history, a work of literature as well as science, and I don’t know quite where to start with its explorations and insights, but since, as a teacher to international students some of whom come from Arabic countries, I’m occasionally faced with disbelief regarding the Darwin-Wallace theory of natural selection from random variation (usually in some such form as ‘you don’t really believe we come from monkeys do you?’), I think it might be interesting, and useful for me, to trace the connections, in time and ideas, between that theory and the discovery of genes that the theory essentially led to.

One of the problems for Darwin’s theory, as first set down, was how variations could be fixed in subsequent generations. And of course another problem was – how could a variation occur in the first place? How were traits inherited, whether they varied from the parent or not? As Mukherjee points out, heredity needed to be both regular and irregular for the theory to work.

There were few clues in Darwin’s day about inheritance and mutation. Apart from realising that it must have something to do with reproduction, Darwin himself could only half-heartedly suggest an unoriginal notion of blending inheritance, while also leaning at times towards Lamarckian inheritance of acquired characteristics – which he at other times scoffed at.

Mukherjee argues here that Darwin’s weakness was impracticality: he was no experimenter, though a keen observer. The trouble was that no amount of observation, in Darwin’s day, would uncover genes. Even Mendel was unable to do that, at least not in the modern DNA sense. But in any case Darwin lacked Mendel’s experimental genius. Still, he did his best to develop a hypothesis of inheritance, knowing it was crucial to his overall theory. He called it pangenesis. It involved the idea of ‘gemmules’ inhabiting every cell of an organism’s body and somehow shaping the varieties of organs, tissues, bones and the like, and then specimens of these varied gemmules were collected into the germ cells to produce ‘mixed’ offspring, with gemmules from each partner. Darwin describes it rather vaguely in his book The Variation of Animals and Plants under Domestication, published in 1868:

They [the gemmules] are collected from all parts of the system to constitute the sexual elements, and their development in the next generation forms the new being; but they are likewise capable of transmission in a dormant state to future generations and may then be developed.

Darwin himself admitted his hypothesis to be ‘rash and crude’, and it was effectively demolished by a very smart Scotsman, Fleeming Jenkin, who pointed out that a trait would be diluted away by successive unions with those who didn’t have it (Jenkin gave as an example the trait of whiteness, i.e. having ‘white gemmules’, but a better example would be that of blue eyes). With an intermingling of sexual unions, specific traits would be blended over time into a kind of uniform grey, like paint pigments (think of Blue Mink’s hit song ‘Melting Pot’).

Darwin was aware of and much troubled by Jenkin’s critique, but he (and the scientific world) wasn’t aware that a paper published in 1866 had provided the solution – though he came tantalisingly close to that awareness. The paper, ‘Experiments in Plant Hybridisation’, by Gregor Mendel, reported carefully controlled experiments in the breeding of pea plants. First Mendel isolated ‘true-bred’ plants, noting seven true-bred traits, each of which had two variants (smooth or wrinkled seeds; yellow or green seeds; white or violet coloured flowers; flowers at the tip or at the branches; green or yellow pods; smooth or crumpled pods; tall or short plants). These variants of a particular trait are now known as alleles. 

Next, he began a whole series of painstaking experiments in cross-breeding. He wanted to know what would happen if, say, a green-podded plant was crossed with a yellow-podded one, or if a short plant was crossed with a tall one. Would they blend into an intermediate colour or height, or would one dominate? He was well aware that this was a key question for ‘the history of the evolution of organic forms’, as he put it.

He experimented in this way for some eight years, with thousands of crosses and crosses of crosses, and the more the crosses multiplied, the more clearly he found patterns emerging. The first pattern was clear – there was no blending. With each crossing of true-bred variants, only one variant appeared in the offspring – only tall plants, only round peas and so on. Mendel named them as dominant traits, and the non-appearing ones as recessive. This was already a monumental result, blowing away the blending hypothesis, but as always, the discovery raised as many questions as answers. What had happened to the recessive traits, and why were some traits recessive and others dominant?

Further experimentation revealed that disappeared traits could reappear in toto in further cross-breedings. Mendel had to carefully analyse the relations between different recessive and dominant traits as they were cross-bred in order to construct a mathematical model of the different ‘indivisible, independent particles of information’ and their interactions.

Although Mendel was alert to the importance of his work, he was spectacularly unsuccessful in alerting the biological community to this fact, due partly to his obscurity as a researcher, and partly to the underwhelming style of his landmark paper. Meanwhile others were aware of the centrality of inheritance to Darwin’s evolutionary theory. The German embryologist August Weismann added another nail to the coffin of the ‘gemmule’ hypothesis in 1883, a year after Darwin’s death, by showing that mice with surgically removed tails – thus having their ‘tail gemmules’ removed – never produced tail-less offspring. Weismann presented his own hypothesis, that hereditary information was always and only passed down vertically through the germ-line, that’s to say, through sperm and egg cells. But how could this be so? What was the nature of the information passed down, information that could contain stability and change at the same time?

The Dutch botanist Hugo de Vries, inspired by a meeting with Darwin himself not long before the latter’s death, was possessed by these questions and, though Mendel was completely unknown to him, he too looked for the answer through plant hybridisation, though less systematically and without the good fortune of hitting on true-breeding pea plants as his subjects. However, he gradually became aware of the particulate nature of hereditary information, with these particles (he called them ‘pangenes’, in deference to Darwin’s ‘pangenesis’), passing down information intact through the germ-line. Sperm and egg contributed equally, with no blending. He reported his findings in a paper entitled Hereditary monstrosities in 1897, and continued his work, hoping to develop a more detailed picture of the hereditary process. So imagine his surprise when in 1900 a colleague sent de Vries a paper he’d unearthed, written by ‘a certain Mendel’ from the 1860s, which displayed a clearer understanding of the hereditary process than anyone had so far managed. His response was to rush his own most recent work into press without mentioning Mendel. However, two other botanists, both as it happened working with pea hybrids, also stumbled on Mendel’s work at the same time. Thus, in a three-month period in 1900, three leading botanists wrote papers highly indebted to Mendel after more than three decades of profound silence.

Hugo de Vries

The next step of course, was to move beyond Mendel. De Vries, who soon corrected his unfair treatment of his predecessor, sought to answer the question ‘How do variants arise in the first place?’ He soon found the answer, and another solid proof of Darwin’s natural selection. The ‘random variation’ from which nature selected, according to the theory, could be replaced by a term of de Vries’ coinage, ‘mutation’. The Dutchman had collected many thousands of seeds from a wild primrose patch during his country rambles, which he planted in his garden. He identified some some 800 new variants, many of them strikingly original. These random ‘spontaneous mutants’, he realised, could be combined with natural selection to create the engine of evolution, the variety of all living things. And key to this variety wasn’t the living organisms themselves but their units of inheritance, units which either benefitted or handicapped their offspring under particular conditions of nature.

The era of genetics had begun. The tough-minded English biologist William Bateson became transfixed on reading a later paper of de Vries, citing Mendel, and henceforth became ‘Mendel’s bulldog’. In 1905 he coined the word ‘genetics’ for the study of heredity and variation, and successfully promoted that study at his home base, Cambridge. And just as Darwin’s idea of random variation sparked a search for the source of that variation, the idea of genetics and those particles of information known as ‘genes’ led to a worldwide explosion of research and inquiry into the nature of genes and how they worked – chromosomes, haploid and diploid cells, DNA, RNA, gene expression, genomics, the whole damn thing. We now see natural selection operating everywhere we’re prepared to look, as well as the principles of ‘artificial’ or human selection, in almost all the food we eat, the pets we fondle, and the superbugs we try so desperately to contain or eradicate. But of course there’s so much more to learn….

William Bateson

Written by stewart henderson

June 14, 2017 at 5:42 pm

bonobo society, sex and females

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sexual dimorphism - a difference on average, but massive individual variation

sexual dimorphism – a difference on average, but massive individual variation

Men are bigger than women, slightly. That’s how things evolved. It’s called sexual dimorphism. It happens with many species, the genders are different in size, shape, coloration, whatever. With humans there’s a size difference, and something of a shape difference, in breasts and hips, but really these aren’t significant. Compare, say, the deep-water triplewort seadevil, a type of anglerfish, in which the female is around 30 cms long, and the male a little over a centimetre. The difference in mass would be too embarrassing to relate.

Among our primate cousins the greatest sexual dimorphism, in size as well as other features, is found in the mandrills, with the male being two to three times the size of the females. In some gorillas there’s a substantial size difference too in favour of the males, and in fact in all of the primate species the male has a size advantage. But size isn’t everything, and the bigger doesn’t have to always dominate.

Female bonobos are smaller than the males, even more so than in humans, yet they enjoy a higher social status than in any other primate society, probably including humans, though it’s hard to compare, since humanity’s many societies vary considerably on the roles and status of women. So how have females attained this exalted status within one of the most highly socialised primate species?

Bonobos and chimpanzees are equally our closest living relatives. It isn’t clear when exactly they separated from each other, but some experts claim it may have been less than a million years ago. Enough time for them to become quite distinct physically, according to the ethologist Franz De Waal. Bonobos are more gracile with longer limbs and a smaller head, and they have a distinctive hairstyle, with a neat parting down the middle. They’re also more easily individuated by their facial features, being in this sense more like humans. And there are also major differences in their social behaviour. Male chimps are dominant in the troupe, often brutally so, whereas bonobo society is less clearly hierarchical, and considerably less violent overall. De Waal, one of the world’s foremost experts on both primates, became interested in bonobos primarily through studies on aggression. He noted that sometimes, after a violent clash, two chimps would come together to hug and kiss. Being interested in such apparent reconciliations and their implications, he decided to look at reconciling behaviours in other primates. What he discovered in bonobos (at San Diego Zoo, which in 1983 housed the world’s largest captive colony) was rather ‘shocking’; their social life was profoundly mediated by sex. Not that he was the first to discover this; other primatologists had written about it, noting also that bonobo sex was far more human-like than chimp sex, but their observations were obscurely worded and not well disseminated. There are other aspects of the physical nature of sexual relations in bonobos that favour females, such as female sexual receptivity, indicated by swelling and a reddening of the genital area, which pertains for a much longer period than in chimps. Female bonobos, like humans and unlike other primates, are sexually receptive more or less all the time.

This isn’t to say that bonobos are oversexed, whatever that may mean. Sexual relations are far from constant, they are casual, sporadic and quickly done with. Often they’re associated with finding food, and it seems likely that sexual relations are used to reconcile tensions related to food availability and other potential causes of conflict.

So how does this use of sex relate to the status of females in bonobo society. I’ll explore this further in the next post.

bonobo relations - more than just sex

bonobo relations – more than just sex

Written by stewart henderson

September 4, 2016 at 1:32 pm

Abiogenesis – LUCA, gradients, amino acids, chemical evolution, ATP and the RNA world

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chemical-evolution-1

Jacinta: So now we’re thinking of the Earth 4 billion years BP, with an atmosphere we’re not quite sure of, and we want to explore the what and when of the first life forms. Haven’t we talked about this before?

Canto: Yeah we talked about the RNA world and viroids and abiogenesis, the gap between chemistry and biology, inter alia. This time we’re going to look more closely at the hunt for the earliest living things, and the environments they might’ve lived in.

Jacinta: And it started with one, it must have. LUA, or LUCA, the last universal common ancestor. Or the first, after a number of not-quite LUCAs, failed or only partially successful attempts. And finding LUCA would be much tougher than finding a viroid in a haystack, because you’re searching through an immensity of space and time.

Canto: But we’re much closer to finding it than in the past because we know so much more about what is common to all life forms.

Jacinta: Yes so are we looking definitely at the first DNA-based life form or are we probing the RNA world again?

Canto: I think we’ll set aside the world of viroids and viruses for now, because we want to look at the ancestor of all independently-existing life forms, and they’re all DNA-based. And we also know that LUCA used ATP. So now I’m going to quote from an essay by Michael Le Page in the volume of the New Scientist Collection called ‘Origin, Evolution, Extinction’:

How did LUCA make its ATP? Anyone designing life from scratch would probably make ATP using chemical reactions inside the cell. But that’s not how it is done. Instead energy from food or sunlight is used to power a protein ‘pump’ that shunts hydrogen ions – protons – out of the cell. This creates a difference in proton concentration, or a gradient, across the cell membrane. Protons then flow back into the cell through another protein embedded in the membrane, which uses the energy to produce ATP.

Jacinta: You understand that?

Canto: Sort of.

Jacinta: ‘Energy from food or sunlight is used..’ that’s a bit of a leap. What food? The food we eat is organic, made from living or formerly living stuff, but LUCA is the first living thing, its food must be purely chemical, not biological.

Canto: Of course, not a problem. I believe the microbes at hydrothermal vents live largely on hydrogen sulphide, and of course sunlight is energy for photosynthesising oganisms such as cyanobacteria.

Jacinta: Okay, so your simplest living organisms, or the simplest ones we know, get their energy by chemosynthesis, or photosynthesis. Its energy, or fuel, not food.

Canto: Semantics.

Jacinta: But there are other problems with this quote re abiogenesis. For example, it’s talking about pre-existent cells and cell membranes. So assuming that cells had to precede ATP.

Canto: No, he’s telling us how cells make ATP today. So we have to find, or synthesise, all the essential ingredients that make up the most basic life forms that we know cell membranes, proteins, ATP and the like. And people are working towards this.

Jacinta: Yes and first of all they created these ‘building blocks of life’, as they always like to call them, amino acids, in the Miller-Urey experiments, since replicated many times over, but what exactly are nucleic acids? Are they the same things as nucleic acids?

Canto: Amino acids are about the simplest forms of organic compounds. It’s probably better to call them the building blocks of proteins. There are many different kinds, but generally each contain amine and carboxyl groups, that’s -NH2 and -COOH, together with a side chain, called an R group, which determines the type of amino acid. There’s a whole complicated lot of them and you could easily spend a whole lifetime fruitfully studying them. They’re important in cell structure and transport, all sorts of things. We’ve not only been able to create amino acids, but to combine them together into longer peptide chains. And we’ve also found large quantities of amino acids in meteorites such as the Murchison – as well as simple sugars and nitrogenous bases. In fact I think we’re gradually firming up the life-came from-space hypothesis.

Jacinta: But amino acids and proteins aren’t living entities, no matter how significant they are to living entities. We’ve never found living entities in space or beyond Earth. Your quote above suggests some of what we need. A boundary between outside and inside, a lipid or phospho-lipid boundary as I’ve heard it called, which must be semi-permeable to allow chemicals in on a very selective basis, as food or fuel.

Canto: I believe fatty acids formed the first membranes, not phospho-lipids. That’s important because we’ve found that fatty acids, which are made up of carbon, hydrogen and oxygen atoms joined together in a regular way, aren’t just built inside cells. There’s a very interesting video called What is Chemical Evolution?, produced by the Center for Chemical Evolution in the USA, that tells about this. Experimenters have heated up carbon monoxide and hydrogen along with many minerals common in the Earth’s crust and produced various carbon compounds including fatty acids. Obviously this could have and can still happen naturally on Earth, for example in the hot regions maybe below or certainly within the crust. It’s been found that large concentrations of fatty acids aggregate in warm water, creating a stable, ball-like configuration. This has to do with the attraction between the oxygen-carrying heads of fatty acids and the water molecules, and the repulsion of the carbon-carrying tails. The tails are forced together into a ball due to this repulsion, as the video shows.

fatty acids, with hydrophobic and hydrophilic ends, aggregating in solution

Jacinta: Yes it’s an intriguing video, and I’m almost feeling converted, especially as it goes further than aggregation due to these essentially electrical forces, but tries to find ways in which chemical structures evolve, so it tries to create a bridge between one type of evolution and another – the natural-selection type of evolution that operates upon reproducing organisms via mutation and selection, and the type of evolution that builds more complex and varied chemical structures from simpler compounds.

Canto: Yes but it’s not just the video that’s doing it, it’s the whole discipline or sub-branch of science called chemical evolution.

Jacinta: That’s right, it’s opening a window into that grey area between life and non-life and showing there’s a kind of space in our knowledge there that it would be exciting to try and fill, through observation and experimentation and testable hypotheses and the like. So the video, or the discipline, suggests that in chemical evolution, the highly complex process of reproduction through mitosis in eukaryotic cells or binary fission in prokaryotes is replaced by repetitive production, a simpler process that only takes place under certain limited conditions.

Canto: So under the right conditions the balls of fatty acids grow in number and themselves accumulate to form skins, and further forces – I think they’re hydrostatic forces – can cause the edges of these skins to fuse together to create ‘containers’, like vesicles inside cells.

Jacinta: So we’re talking about the creation of membranes, impermeable or semi-permeable, that can provide a safe haven for, whatever…

Canto: Yes, and at the end of the video, other self-assembling systems, such as proto-RNA, are intriguingly mentioned, so we might want to find out what’s known about that.

Jacinta: I think we’ll be doing a lot of reading and posting on this subject. I find it really fascinating. These limited conditions I mentioned – limited on today’s Earth surface, but not so much four billion years ago, include a reducing atmosphere lacking in free oxygen, and high temperatures, as well as a gradient – both a temperature gradient and a sort of molecular or chemical gradient, from more reducing to more oxidising you might say. These conditions exist today at hydrothermal vents, where archaebacteria are found, so researchers are naturally very interested in such environments, and in trying to replicate or simulate them.

Canto: And they’re interested in the boundary between chemical and biological evolution, and reproduction. There are so many interesting lines of inquiry, with RNA, with cell membranes….

Jacinta: Researchers are particularly interested in alkaline thermal vents, where alkaline fluids well up from beneath  the sea floor at high temperatures. When this fluid hits the ocean water, minerals precipitate out and gradually create porous chimneys up to 60 metres high. They would’ve been rich in iron and sulphide, good for catalysing complex organic reactions, according to Le Page. The temperature gradients created would’ve favoured organic compounds and would’ve likely encouraged the building of complexity, so they may have been the sites in which the RNA world began, if it ever did.

a hydrothermal vent off the coast of New Zealand. Image from NOAA

a hydrothermal vent off the coast of New Zealand. Image from NOAA

Canto: So I think we should pursue this further. There are a lot of researchers homing in on this area, so I suspect further progress will be made soon.

Jacinta: Yes, we need to explore the exploitation of proton gradients, the development  of proton pumps and the production of ATP, leaky membranes and a whole lot of other fun stuff.

Canto: I think we need to get our heads around ATP and its production too, because that looks pretty damn complex.

Jacinta: Next time maybe.

 

Written by stewart henderson

July 29, 2016 at 8:51 am

clever Charlie Darwin

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A photo taken by me! King Charles seated in state in the Musuem of Natural History, London. It was a thrill to be granted an audience

A photo taken by me! King Charles seated in state in the Musuem of Natural History, London. It was a thrill to be granted an audience

I recently decided to reread Darwin’s Origin of Species, which was really reading it for the first time as my first reading was pretty cursory, and I could barely follow the wealth of particular knowledge he used for cumulative effect to adduce his theory. This time I’ve been doing a closer reading, and becoming increasingly impressed, and I’ve only read the first chapter, ‘Variation under Domestication’.

Darwin’s argument here of course is that domesticated horses, dogs, birds and plants have been artificially selected over long periods of time, and often unconsciously, to suit human needs and tastes. This might seem screamingly obvious today, and to a degree it was recognised in Darwin’s time, but because of an inability to take the long view, and also because of the then-prevalent paradigm of the fixity of species, breeders and nurserymen tended to under-estimate their own cumulative powers, and to claim, for example, that dogs and pigeons had always come in many varieties. Even Darwin was uncertain, and was willing to concede – writing of course before the advent of Mendelian genetics, never mind the revolution wrought by the identification and analysis of DNA as the molecule of inheritance – that in some cases the breeders might be right:

In the case of most of our anciently domesticated animals and plants, I do not think it is possible to come to any definite conclusion, whether they have descended from one or several species.

He was even prepared to concede that it was ‘highly probable that our domestic dogs have descended from several wild species’, while at the same time arguing that the breeding of dogs, in Egypt, other parts of Africa and Australia (where, in his Beagle travels, he observed dingoes, which he may have seen as semi-domesticated by the Aborigines) extended back far further in time than most people suspected. We now know that Darwin’s concession here was ‘premature’. The latest research strongly suggests that our domesticated dogs trace their ancestry to a group of European wolves dating from 19,000 to 32,000 years ago, and probably now extinct. That’s a time-frame Darwin would’ve baulked at, and it’s both funny and kind of tragic that this is something I’ve ‘discovered’ after 30 seconds of selective internet searching. There’s no doubt, though that Darwin’s bold but always informed speculations were heading in the right direction.

Particularly informed –  and bold – were his speculations about pigeons. This is hardly surprising as he spent several years studying and breeding them himself. Interestingly, he started doing so because he’d become convinced that all the fancy pigeons then on show were most likely derived from one common species, the rock pigeon or rock dove (Columba livia), a view already held by some naturalists but few breeders.  He devotes several pages in Chapter 1 to arguing his case, for example pointing out that the ‘several distinct species’ argued for by breeders can be crossed with complete success, that’s to say with no signs of sterility or more than usually defective offspring.

So, as with dogs, I decided to look up what the latest research was on the ancestry of English carriers, short-faced tumblers, runts, fantails, common tumblers, barbs, pouters, trumpeters and laughers, to name some of the pigeons Darwin mentions in the chapter, and was excited to find that a piece of research published as recently as 2013 has confirmed Darwin’s hypothesis. Cheaper and faster genome sequencing technologies have enabled researchers to sequence the genomes of many wild and domesticated birds, and they’ve found that all of the latter are clearly closer to C livia than to any other wild species. It only took just over 150 years for Darwin to be proven correct.

Close reading like this really does reap some fun rewards, and I’ll finish with two more examples. Darwin wrote of how in the world of breeding, quite a drastic change can be brought about in one breeding step, as in the case of the fuller’s teasel with its hooks. He goes on:

So it has probably been with the turnspit dog; and this is known to have been the case with the ancon sheep.

Not knowing wtf he was talking about, I irritatedly decided to look up these unknown creatures. The turnspit dog is a now-extinct breed, bred specifically from around the 16th century to provide the dogpower to turn meat on a spit, the only conceivable way of cooking large joints of meat in your average fancy household for a couple of centuries. The dog, or dogs, because the system worked better if you had two of them engaged in shift work, turned a wheel by running inside it, rat-like, until the meat was cooked. They were known to be long-bodied and short-legged, but details of how they were bred aren’t known, as they were apparently beneath scholarly consideration. They certainly weren’t seen as cuddly pets – if you treat creatures as slaves it heightens your contempt for then (cf Aristotle) – and they were even taken to church as foot-warmers. They’d disappeared entirely by the end of the 19th century.

It's a dog's life?

It’s a dog’s life?

The ancon sheep was a short-legged type, apparently bred from a single individual in the USA in the late nineteenth century, its short legs having the singular advantage, to some, of curtailing its hopes of freedom by jumping the fence. The term ‘ancon’ has since been used by breeding researchers to describe strains of creatures arising from an individual with the same phenotype.

Achondroplastic_sheep

Written by stewart henderson

June 4, 2016 at 11:00 am

What is a trisomy?

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Canto: So I happened to watch an excellent video from the Royal Institute recently, a talk by the beautifully named and beautifully voiced Irish geneticist, Aoife Mclysaght…

Jacinta: How do you pronounce that?

Canto: It’s pronounced Aoife Mclysaght…

Jacinta: Oh right.

Canto: So the theme was that everything in biology makes sense only in the light of evolution, and she was illustrating this through her area of interest and research, gene duplication. And along the way she talked about trisomies, particularly trisomy 21, usually referred to as Down Syndome.

Jacinta: A trisomy involves having an extra copy of a chromosome, in this case chromosome 21.

Canto: Very good, and the extra copy is a perfectly good copy, but having that extra copy causes major problems, obviously.

Jacinta: The term ‘trisomy’ refers of course to three – having three rather than two sets of a particular chromosome. Humans normally have two sets of 23 chromosomes. I have a relative who has a rare and unnamed form of trisomy, or at least a rare form of chromosomal disorder, which, when looking into it, I decided must be a trisomy. But since then I’ve discovered that Williams syndrome – which I learned about from another person I know with that condition – isn’t a trisomy, but the result of genes missing from chromosome 7. So now I’ve gone from thinking that trisomies accounted for all or most sorts of genetic intellectual disabilities to… I don’t know.

Canto: To a position of deeper ignorance. So people with trisomies have 47 chromosomes, with Down syndrome being the most common. Others include Edward syndrome (trisomy 18) and Patau syndrome (trisomy 13). Interestingly, though, there’s another rarer form of Down syndrome that’s due to translocation – that’s when a part of a chromosome – in this case chromosome 21 – migrates to another chromosome, usually chromosome 14, during cell division

Jacinta: That complicates matters… So do we know what causes these trisomies, and these translocations? They seem to be very specific, occurring for only particular chromosomes, or bits of them.

Canto: Well you’re right in that trisomies 21, 18 and 13 are relatively common – I mean rare but more common than a trisomy 9 or 15 or 19, just to pick out any numbers less than 23. We do know that trisomies become more common with older egg cells. As you know, your egg cells are as old as you are, and they become a little decrepit with age like yourself.

Jacinta: We’re both slouching to oblivion.

Canto: It’s also the case that most trisomies don’t survive to term, in fact they mostly miscarry so early that the mother doesn’t even know she’s been pregnant. So presumably those trisomies I just picked at random, if they occur at all, have more fatal consequences. It seems in any case that a trisomy occurs when cells divide but one chromosome somehow sticks to its homologue and is carried with it into the new cell. So maybe some chromosomes are more ‘sticky’ than others?

Jacinta: I think we need to do a deeper dive, as one pundit likes to say, into meiosis and aneuploidy.

Canto: Aneuploidy?

Jacinta: That’s just when you have an abnormal number of chromosomes per cell: it could be less or more. Actually trisomy 16 is the most common form, but fatal in its full-blown version. It can exist in mosaic form – when not all the cells have it.

Canto: So can you explain meiosis for us?

Jacinta: A long story but fascinating of course. It’s the basis of sexual reproduction for all eukaryotes. So before eukaryotic germ cells or gametes divide they need to replicate their chromosomes so that the resulting pair of cells has an equal share. This period of replication is known as the S phase.

Canto: Wait a minute, does this mean that in the S phase humans have 92 chromosomes per cell instead of 46?

Jacinta: Don’t bog me down with clever questions. Taking another step back, we have this whole process called the cell cycle, which we divide into phases. We can start anywhere, since it’s a cycle, if you know what I mean, but if you need a beginning it’s the prophase. Anyway, the S phase comes after the G1 phase and before the G2 phase. S, by the way, stands for synthesis, and G here stands for gap. Together these three phases make up the interphase, at the end of which we have the prophase of a new cell cycle, though actually meiosis isn’t a cycle the way mitosis (non-sexual reproduction or cell division) is. To be accurate, the next phase is called prophase 1, which is followed by metaphase 1, anaphase 1 and telophase 1 before we have prophase 2….

Canto: Stop cycling I’m getting dizzy.

Jacinta: Well yes believe me it’s complicated, and I haven’t begun yet. But you did ask for it.

Canto: Can you give the simplified version?

Jacinta: Not really.

Canto: Okay, we’ll leave that for another day. Focusing in on the part of meiosis when these trisomies and other anomalies occur, it seems that the problem isn’t so much stickiness as non-stickiness. Think of gametes. In mammals such as humans there are two types, egg and sperm cells. They’re differentiated by their sex chromosomes, chromosome 23…

Jacinta: And also by the fact that the egg cell is like the sun and the sperm cell is like the earth.

Canto: Well, sort of, in terms of volume. Now, after meiosis – which occurs in phases, meiosis 1 and meiosis 2, creating two daughter cells then four grand-daughter cells, so to speak – each of these grand-daughter gametes should be haploid. That’s to say, they should contain only one of each of the 23 chromosomes. But nothing’s perfect and sometimes there are errors, and we’re not clear about why, though the chances of error rise with the age of the female as mentioned before. Mostly the problem is that the chromosomes didn’t properly separate, a state called chromosome nondisjunction. Something to do with the spindle apparatus not functioning properly due to a lack of cohesion of the chromosome. This occurs rather more frequently in female meiosis, or oogenesis, than in male meiosis, or spermatogenesis, they’re not sure exactly why.

chromosome_nondisjunction_meiosis

Jacinta: Well I must say that’s all very enlightening, and salutary, as it’s made me aware of how little I know about genetics in general. Now I know a teensy bit more. As to trisomies and other such chromosomal problems, what they know just makes me keen to know more about how we might detect them and possibly in the deep future rectify them at source. But the science is clearly a long way from that…

Canto: Well you never know. Genetics is a fast-moving field.

Jacinta: we must explore it more. It’s serious fun.

Written by stewart henderson

January 31, 2016 at 10:06 pm