Fusion and sex in protocells & the start of evolution

In 1864, five years after reading Darwin’s On the Origin of Species, Pyotr Kropotkin — the anarchist prince of mutual aid — was leading a geographic survey expedition aboard a dog-sleigh — a distinctly Siberian variant of the HMS Beagle. In the harsh Manchurian climate, Kropotkin did not see competition ‘red in tooth and claw’, but a flourishing of cooperation as animals banded together to survive their environment. From this, he built a theory of mutual aid as a driving factor of evolution. Among his countless observations, he noted that no matter how selfish an animal was, it still had to come together with others of its species, at least to reproduce. In this, he saw both sex and cooperation as primary evolutionary forces.

Now, Martin A. Nowak has taken up the challenge of putting cooperation as a central driver of evolution. With his colleagues, he has tracked the problem from myriad angles, and it is not surprising that recently he has turned to sex. In a paper released at the start of this month, Sam Sinai, Jason Olejarz, Iulia A. Neagu, & Nowak (2016) argue that sex is primary. We need sex just to kick start the evolution of a primordial cell.

In this post, I want to sketch Sinai et al.’s (2016) main argument, discuss prior work on the primacy of sex, a similar model by Wilf & Ewens, the puzzle over emergence of higher levels of organization, and the difference between the protocell fusion studied by Sinai et al. (2016) and sex as it is normally understood. My goal is to introduce this fascinating new field that Sinai et al. (2016) are opening to you, dear reader; to provide them with some feedback on their preprint; and, to sketch some preliminary ideas for future extensions of their work.

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Chemical games and the origin of life from prebiotic RNA

From bacteria to vertebrates, life — as we know it today — relies on complex molecular interactions, the intricacies of which science has not fully untangled. But for all its complexity, life always requires two essential abilities. Organisms need to preserve their genetic information and reproduce.

In our own cells, these tasks are assigned to specialized molecules. DNA, of course, is the memory store. The information it encodes is expressed into proteins via messenger RNAs.Transcription (the synthesis of mRNAs from DNA) and translation (the synthesis of proteins from mRNAs) are catalyzed by polymerases necessary to speed up the chemical reactions.

It is unlikely that life started that way, with such a refined division of labor. A popular theory for the origin of life, known as the RNA world, posits that life emerged from just one type of molecule: RNAs. Because RNA is made up of base-complementary nucleotides, it can be used as a template for its own reproduction, just like DNA. Since the 1980s, we also know that RNA can act as a self-catalyst. These two superpowers – information storage and self-catalysis – make it a good candidate for the title of the first spark of life on earth.

The RNA-world theory has yet to meet with empirical evidence, but laboratory experiments have shown that self-preserving and self-reproducing RNA systems can be created in vitro. Little is known, however, about the dynamics that governed pre- and early life. In a recent paper, Yeates et al. (2016) attempt to shed light on this problem by (1) examining how small sets of different RNA sequences can compete for survival and reproduction in the lab and (2) offering a game-theoretical interpretation of the results.

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Social algorithms and the Weapons of Math Destruction

Cathy O'Neil holding her new book: Weapons of Math Destruction at a Barnes & Noble in NYC.

Cathy O’Neil holding her new book: Weapons of Math Destruction at a Barnes & Noble in New York city.

In reference to intelligent robots taking over the world, Andrew Ng once said: “I don’t work on preventing AI from turning evil for the same reason that I don’t work on combating overpopulation on the planet Mars.” Sure, it will be an important issue to think about when the time comes. But for now, there is no productive way to think seriously about it. Today there are more concrete problems to worry about and more basic questions that need to be answered. More importantly, there are already problems to deal with. Problems that don’t involve super intelligent tin-men, killer robots, nor sentient machine overlords. Focusing on distant speculation obscures the fact that algorithms — and not necessarily very intelligent ones — already reign over our lives. And for many this reign is far from benevolent.

I owe much of my knowledge about the (negative) effects of algorithms on society to the writings of Cathy O’Neil. I highly recommend her blog mathbabe.org. A couple of months ago, she shared the proofs of her book Weapons of Math Destruction with me, and given that the book came out last week, I wanted to share some of my impressions. In this post, I want to summarize what makes a social algorithm into a weapon of math destruction, and share the example of predictive policing.

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Argument is the midwife of ideas (and other metaphors)

In their classic book Metaphors We Live By, George Lakoff and Mark Johnson argue — very convincingly, and as I’ve reviewed before — that “[m]etaphor is one of our most important tools for trying to comprehend partially what cannot be comprehended totally” and that these conceptual metaphors are central to shaping our understanding of and interaction with the world we are embedded in. Based on the authors’ grounding in linguistics, part of their case proceeds by offering examples of, by my count, over 58 different metaphors and metonymies in our everyday language; and given their book’s intentions, they chose a particularly pertinent first case: ARGUMENT is WAR.[1]

They show this metaphor in action through some example of common usage (pg. 4):

What do you want me to do? LEAVE? Then they'll keep being wrong!Your claims are indefensible.
He attacked every weak point in my argument.
His criticisms were right on target.
I demolished his argument.
I’ve never won an argument with him.
You disagree? Okay, shoot!
If you use that strategy, he’ll wipe you out.
He shot down all my arguments.

Notice that the even the xkcd I borrowed for visual reinforcement is titled ‘Duty Calls’, an expression usually associated with a departure for war. With our awareness drawn to this militaristic structure, Lakoff and Johnson encourage the reader to ask themselves: how would discussions look if instead of structuring arguments adversarially, we structured them after a cooperative activity like dance?[2]

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Computational kindness and the revelation principle

In EWD1300, Edsger W. Dijkstra wrote:

even if you have only 60 readers, it pays to spend an hour if by doing so you can save your average reader a minute.

He wrote this as the justification for the mathematical notations that he introduced and as an ode to the art of definition. But any writer should heed this aphorism.[1] Recently, I finished reading Algorithms to Live By by Brian Christian and Tom Griffiths.[2] In the conclusion of their book, they gave a unifying name to the sentiment that Dijkstra expresses above: computational kindness.

As computer scientists, we recognise that computation is costly. Processing time is a limited resource. Whenever we interact with others, we are sharing in a joint computational process, and we need to be mindful of when we are not carrying our part of the processing burden. Or worse yet, when we are needlessly increasing that burden and imposing it on our interlocutor. If you are computationally kind then you will be respectful of the cognitive problems that you force others to solve.

I think this is a great observation by Christian and Griffiths. In this post, I want to share with you some examples of how certain systems — at the level of the individual, small group, and society — are computationally kind. And how some are cruel. I will draw on examples from their book, and some of my own. They will include, language, bus stops, and the revelation principle in algorithmic game theory.
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Systemic change, effective altruism and philanthropy

Keep your coins. I want change.The topics of effective altruism and social (in)justice have weighed heavy on my mind for several years. I’ve even touched on the latter occasionally on TheEGG, but usually in specific domains closer to my expertise, such as in my post on the ethics of big data. Recently, I started reading more thoroughly about effective altruism. I had known about the movement[1] for some time, but had conflicting feelings towards it. My mind is still in disarray on the topic, but I thought I would share an analytic linkdex of some texts that have caught my attention. This is motivated by a hope to get some guidance from you, dear reader. Below are three videos, two articles, two book reviews and one paper alongside my summaries and comments. The methods range from philosophy to comedy and from critical theory to social psychology. I reach no conclusions.

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Cancer metabolism and voluntary public goods games

When I first came to Tampa to do my Masters[1], my focus turned to explanations of the Warburg effect — especially a recent paper by Archetti (2014) — and the acid-mediated tumor invasion hypothesis (Gatenby, 1995; Basanta et al., 2008). In the course of our discussions about Archetti (2013,2014), Artem proposed the idea of combining two public goods, such as acid and growth factors. In an earlier post, Artem described the model that came out of these discussions. This model uses two “anti-correlated” public goods in tumors: oxygen (from vasculature) and acid (from glycolytic metabolism).

The dynamics of our model has some interesting properties such as an internal equilibrium and (as we showed later) cycles. When I saw these cycles I started to think about “games” with similar dynamics to see if they held any insights. One such model was Hauert et al.’s (2002) voluntary public goods game.[2] As I looked closer at our model and their model I realized that the properties and logic of these two models are much more similar than we initially thought. In this post, I will briefly explain Hauert et al.’s (2002) model and then discuss its potential application to cancer, and to our model.
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Mutation-bias driving the evolution of mutation rates

In classic game theory, we are often faced with multiple potential equilibria between which to select with no unequivocal way to choose between these alternatives. If you’ve ever heard Artem justify dynamic approaches, such as evolutionary game theory, then you’ve seen this equilibrium selection problem take center stage. Natural selection has an analogous ‘problem’ of many local fitness peaks. Is the selection between them simply an accidental historical process? Or is there a method to the madness that is independent of the the environment that defines the fitness landscape and that can produce long term evolutionary trends?

Two weeks ago, in my first post of this series, I talked about an idea Wallace Arthur (2004) calls “developmental bias”, where the variation of traits in a population can determine which fitness peak the population evolves to. The idea is that if variation is generated more frequently in a particular direction, then fitness peaks in that direction are more easily discovered. Arthur hypothesized that this mechanism can be responsible for long-term evolutionary trends.

A very similar idea was discovered and called “mutation bias” by Yampolsky & Stoltzfus (2001). The difference between mutation bias and developmental bias is that Yampolsky & Stoltzfus (2001) described the idea in the language of discrete genetics rather than trait-based phenotypic evolution. They also did not invoke developmental biology. The basic mechanism, however, was the same: if a population is confronted with multiple fitness peaks nearby, mutation bias will make particular peaks much more likely.

In this post, I will discuss the Yampolsky & Stoltzfus (2001) “mutation bias”, consider applications of it to the evolution of mutation rates by Gerrish et al. (2007), and discuss how mutation is like and unlike other biological traits.

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Lotka-Volterra, replicator dynamics, and stag hunting bacteria

Happy year of the monkey!

Last time in the Petri dish, I considered the replicator dynamics between type-A and type-B cells abstractly. In the comments, Arne Traulsen pointed me to Li et al. (2015):

We have attempted something similar in spirit with bacteria. Looking at frequencies alone, it looked like coordination. But taking into account growth led to different conclusions […] In that case, things were more subtle than anticipated…

So following their spirit, I will get more concrete in this post and replace type-A by Curvibacter sp. AEP13 and type-B by Duganella sp. C1.2 — two bacteria that help fresh water Hydra avoid fungal infection. And I will also show how to extend our replicator dynamics with growth and changing cell density.

Although I try to follow Arne’s work very closely, I had not read Li et al. (2015) before, so I scheduled it for a reading group this past Friday. I really enjoyed the experiments that they conducted, but I don’t agree with their interpretations that taking growth into account leads to a different conclusion. In this post, I will sketch how they measured their experimental system and then provide a replicator equation representation of the Lotka-Volterra model they use to interpret their results. From this, we’ll be able to conclude that C and D are playing the Stag Hunt — or coordination, or assurance, pick your favorite terminology — game.

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Hadza hunter-gatherers, social networks, and models of cooperation

At the heart of the Great Lakes region of East Africa is Tanzania — a republic comprised of 30 mikoa, or provinces. Its border is marked off by the giant lakes Victoria, Tanganyika, and Malawi. But the lake that interests me the most is an internal one: 200 km from the border with Kenya at the junction of mikao Arusha, Manyara, Simiyu and Singed is Lake Eyasi. It is a temperamental lake that can dry up almost entirely — becoming crossable on foot — in some years and in others — like the El Nino years — flood its banks enough to attract hippos from the Serengeti.

For the Hadza, it is home.

The Hadza number around a thousand people, with around 300 living as traditional nomadic hunter-gatherers (Marlow, 2002; 2010). A life style that is believed to be a useful model of societies in our own evolutionary heritage. An empirical model of particular interest for the evolution of cooperation. But a model that requires much more effort to explore than running a few parameter settings on your computer. In the summer of 2010, Coren Apicella explored this model by traveling between Hadza camps throughout the Lake Eyasi region to gain insights into their social network and cooperative behavior.

Here is a video abstract where Coren describes her work:

The data she collected with her colleagues (Apicella et al., 2012) provides our best proxy for the social organization of early humans. In this post, I want to talk about the Hadza, the data set of their social network, and how it can inform other models of cooperation. In other words, I want to freeride on Apicella et al. (2012) and allow myself and other theorists to explore computational models informed by the empirical Hadza model without having to hike around Lake Eyasi for ourselves.

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