Optimizing dynastic succession genetically

The traditional rule for succession in a monarchy is to pass from father to son. Much of King Henry VIII’s spousal folly was over his anxiety for producing an heir. However, if the basis of being a successful ruler has a genetic component then this would be the least optimal way to run an empire. For diploid sexually reproducing organisms, such as humans, the offspring inherits equal numbers of chromosomes from both parents and classically the genetic relationship or kinship coefficient between parent and child is assigned the value of 1/2.  However, there is a crucially important asymmetry in that males are heterozygous in the sex chromosomes, i.e. they inherit an X chromosome from their mothers and a Y from their fathers, while females are homozygous, inheriting an X from both. Now the X is about 100 million base pairs longer than the Y, which accounts for about 2 percent of the (father’s) genome (counting chromosomes separately). Additionally, given that everyone has at least one X while only males have a Y, the Y cannot contain genes that are crucial for survival and in fact there are much fewer genes on the Y than the X (~800 vs ~50). The Y has been shrinking in mammals over time and there is a debate about its importance and eventual fate (e.g. see here).

We can compute the sex chromosome adjusted genetic correlation coefficients between parents and children.  Let the father’s genetic content be F=F_S + F_D, where F_S is the genetic content passed to sons (half of the autosomes plus the Y chromosome) and F_D is that passed to daughters (half of the autosomes plus the X) and similarly M=M_S+M_D. The son genetic content is then S=F_S+M_S and daughter is D=F_D+M_D. We can treat F and M as a string of random variables with variance 1/(length of mother’s genome) and assuming that the genetic correlation between fathers and mothers is zero (i.e. no inbreeding and no assortative mating) then the correlation coefficient between father and son is

\langle FS\rangle = \frac{ \langle F_S^2\rangle}{\sqrt{\langle F_S^2\rangle+\langle F_D^2\rangle}\sqrt{\langle F_S^2\rangle+\langle M_S^2\rangle}}=\frac{ 1}{\sqrt{1+\langle F_D^2\rangle/\langle F_S^2\rangle}\sqrt{1+\langle M_S^2\rangle/\langle F_S^2\rangle}}

and similarly:

\langle FD\rangle =\frac{ 1}{\sqrt{1+\langle F_S^2\rangle/\langle F_D^2\rangle}\sqrt{1+\langle M_D^2\rangle/\langle F_D^2\rangle}}

\langle MS\rangle =\frac{ 1}{\sqrt{1+\langle M_D^2\rangle/\langle M_S^2\rangle}\sqrt{1+\langle F_S^2\rangle/\langle M_S^2\rangle}}

\langle MD\rangle =\frac{ 1}{\sqrt{1+\langle M_S^2\rangle/\langle M_D^2\rangle}\sqrt{1+\langle F_D^2\rangle/\langle M_D^2\rangle}}

Now, if you assume that genetic content is homogeneous among all chromosomes then that would mean that the genetic material that fathers pass on to sons is 0.48 of the total and thus \langle F_S^2\rangle = 0.48 while \langle F_D^2\rangle = 0.5, \langle M_S^2\rangle = 0.5, and \langle M_D^2\rangle = 0.5 implying that \langle FS\rangle = 0.49\langle FD\rangle = 0.51\langle MS\rangle = 0.51\langle MD\rangle = 0.5 . Hence, parents are more correlated with their children of the opposite sex and fathers are least correlated with their sons. These numbers also probably underestimate the asymmetry. If genetic relationship is the most important factor for royal succession then a dynasty based on opposite sex succession will be more logical than the father to son model.



Selection of the week

Here’s a concert I wish I could have attended. A young Glenn Gould (greatest Bach interpreter since Bach although I heard Felix Mendelssohn was pretty good too) with Leonard Bernstein in his prime conducting the New York Philharmonic (I think) playing the first movement of JS Bach’s Keyboard Concerto in D minor, BWV 1052.

There is a famous incident of a Gould performance of the Brahms Piano Concerto 1 when he and Bernstein had such a disagreement on the tempo (Gould wanted to play it really slow) that Bernstein got up on stage beforehand to make a disclaimer. That performance with speech is recorded and someone has uploaded it to YouTube.

Gould gave up performing in 1964 at age 31. Notice how low he likes to sit at the piano. He used to bring his chair with him when he toured. One of my favourite films is “Thirty two short films about Glenn Gould,” which I definitely recommend seeing.

Are we in a fusion renaissance?

Fusion is a potentially unlimited source of non-carbon emitting energy. It requires the mashing together of small nuclei such as deuterium and tritium to make another nucleus and a lot of leftover energy. The problem is that nuclei do not want to be mashed together and thus to achieve fusion you need something to confine high energy nuclei for a long enough time. Currently, there are only two methods that have successfully demonstrated fusion: 1) gravitational confinement as in the center of a star, and 2) inertial confinement as in a nuclear bomb. In order to get nuclei at high enough energy to overcome the energy barrier for a fusion reaction, electrons can no longer be bound to nuclei to form atoms. A gas of quasi-neutral hot nuclei and electrons is called a plasma and has often been dubbed the fourth state of matter. Hence, the physics of fusion is mostly the physics of plasmas.

My PhD work was in plasma physics and although my thesis ultimately dealt with chaos in nonlinear partial differential equations, my early projects were tangentially related to fusion. At that time there were two approaches to attaining fusion, one was to try to do controlled inertial confinement by using massive lasers to implode a tiny pellet of fuel and the second was to use magnetic confinement in a tokamak reactor. Government sponsored research has been focused almost exclusively on these two approaches for the past forty years. There is a huge laser fusion lab at Livermore and an even bigger global project for magnetic confinement fusion in Cadarache France, called ITER. As of today, neither has proven that they will ever be viable sources of energy although there is evidence of break even where the reactors produce more energy than is put in.

However, these approaches may not ultimately be viable and there really has not been much research funding to pursue alternative strategies. This recent New York Times article reports on a set of privately funded efforts to achieve fusion backed by some big names in technology including Paul Allen, Jeff Bezos and Peter Thiel. Although there is well deserved skepticism for the success of these companies,  (I’m sure my thesis advisor Abe Bers would have had some insightful things to say about them), the time may be ripe for new approaches. In an impressive talk I heard many years ago, roboticist Rodney Brooks remarked that Moore’s Law has allowed robotics to finally be widely available because you could use software to compensate for hardware. Instead of requiring cost prohibitive high precision motors, you could use cheap ones and use software to control them. The hybrid car is only possible because of the software to decide when to use the electric motor and when to use the gas engine. The same idea may also apply to fusion. Fusion is so difficult because plasmas are inherently unstable. Most of the past effort has been geared towards designing physical systems to contain them. However, I can now imagine using software instead.

Finally, government attempts have mostly focused on using a Deuterium-Tritium fusion reaction because it has the highest yield. The problem with this reaction is that it produces a neutron, which then destroys the reactor. However, there are reactions that do not produce neutrons (see here). Abe used to joke that that we could mine the moon for Helium 3 to use in a Deuterium-Helium 3 reactor. So, although we may never have viable fusion on earth, it could be a source of energy on Elon Musk’s moon base, although solar would probably be a lot cheaper.