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paleoanthropology, genetics and evolution

stature

  • Statures of fossil Homo

    Tue, 2011-09-13 00:25 -- John Hawks
    Synopsis: 
    A laboratory exercise that applies regression equations to estimate the statures of some fossil hominin femora.

    Homo erectus and Neandertals were more or less human-sized. That may not be saying much, since we are so variable in stature ourselves.

    In this case, the fossils don't entirely speak for themselves. To estimate the sizes of ancient people, working with long bones, we must apply some kind of regression or other estimation method.

    1. KNM-ER 1481 is a complete femur from Koobi Fora, Kenya, approximately 1.9 million years old. Without any associated skull or teeth, we can't be sure what species it represents. Many scientists attribute it to early Homo because of its differences from known australopithecine femora.
    2. The Trinil femur was found by Eugene Dubois in 1892 as he excavated fossil beds at Trinil, Java. He had found a human skullcap the year before, and after finding the femur's humanlike anatomy, Dubois named a new species, Pithecanthropus erectus. This is the original Homo erectus femur. Today, we are less certain about its age and association with the partial skull. It may be a million years old, but it may be substantially younger.
    3. The femur from Spy, Belgium, represents a Neandertal who lived around 45,000 years ago. This femur is part of a more complete skeleton, and exhibits many of the characteristic features of Neandertal long bones, including the great thickness and curvature of the shaft and very large joint surfaces.
    4. What to do: Here you will examine the fossil cast femora, using regression equations to predict stature of the individual.

      1. Determine the sex of the individual. The femur head diameter is a relatively good indicator of sex. If it is less than 44 mm, the individual is likely to be a female. More than 46 mm, and the individual is likely to be a male. In between these values, you may need more information — either from the rest of the skeleton or from the size and robusticity of the femur itself.
      2. Measure the maximum length of the femur. This measurement is taken using the osteometric board, and represents the maximum distance from any points on the proximal and distal ends of the bone. Take your measurement in centimeters.
      3. Apply the correct regression equation. These are specific to sex and race. The femora at this station come from donated anatomy collections from the early 20th century, and represent people of European ancestry. The male and female regression equations for this population are listed at right.
    Study questions: 
    1. What are some weaknesses of estimating body size for fossil humans by applying a regression drawn from a contemporary human population?
  • Heritability and stature

    Mon, 2011-09-12 01:42 -- John Hawks
    Synopsis: 
    Heritability is the proportion of variance in a phenotype explained by additive genetic variance.

    Tall parents tend to have tall children.

    That's a simple generalization, not an absolute statement. You may be a short person with two tall parents. If you know many families, you'll probably know someone who is an exception to the rule. Two short parents may have a very tall daughter, and two siblings may be very different heights.

    Still, if we look at many families we will find that the stature of the parents lets us make some fair predictions about the statures of their children. We can look at data to quantify just how parent and offspring heights are related.

    For example, here is a plot of the statures of students in my Anthropology 105 class in 2010, compared to the mean of their mothers' and fathers' statures. The average of mother and father's heights are the midparent stature.

    Young men in this class have a taller average stature than young women. The picture separates men and women, and both taller sons and daughters tend to come from taller parents.

    We can do a bit better than this to quantify the relationship of the parents' and sons' and daughters' statures: We can put lines on the graph to show how the sons' and daughters' heights tend to increase with the midparent stature:

    Each of these lines is called a linear regression between midparent stature and offspring stature. The linear regression is the line that has the smallest squared distance from all of the points, added together.

    The squared distance is special. In statistics, the average squared distance from all points to the mean is called the variance. When we consider a trait like stature, there are many possible biological causes that can contribute to individuals being taller or shorter than the average. In statistical terms, these causes are all factors that contribute to the variance of stature.

    In the chart above, the linear regression represents the amount of the variance of the stature of the offspring that was contributed by the variance of their midparent statures. Parent statures can predict offspring statures and the linear regression is the prediction.

    It's not a perfect prediction. Take a look at the midparent value of 160 cm. When the midparent average is 160 cm, the regressions predict that daughters will be 156 cm and sons will be 168 cm. Out of Anthropology 105 students last year, two men had parents with an average of 160 cm, and both of them were very close to 168 cm in height — a good prediction! But the two women with parents this stature have very different heights. One is only 148 cm, the other 162, both more than 2 inches from the predicted value, in different directions. The regression gives the best prediction we can make from the parents, but there is obviously a lot of variation that can't be predicted in that way.

    Let's look more closely at the female students. The following chart adds together females from several years of Anthropology 105, with their midparent statures.

    The slope of the regression across these 300 women is 0.72. That means that roughly 72 percent of the variance of the students' stature can be attributed to variance in their midparent stature.

    This regression is special in genetics. Daughters resemble their parents because they inherit genes from them. The regression between the midparent and daughters' statures gives us a way to estimate the effects of those genes. In fact, the proportion of variance in a trait that can be explained by genes is the same as the slope of the regression. Geneticists call this proportion the heritability of stature. In my Anthropology 105 classes over the last few years, we would estimate the heritability of stature as 0.72, or 72%.

    Again, there are exceptions. Sometimes parents give other things to their children besides genes. The right foods, the right resources can make a difference to growth and development. And some genes can have unexpected effects. Most obviously, some genes make sons a lot taller than daughters, which is why we've considered the two sexes separately.

    The heritability of a trait is a powerful concept. It's important to understand some of its strengths and limits:

    1. Heritability refers to a population. A female in my Anthropology 105 class may be taller or shorter than her parents. We can't predict 72% of that student's stature; instead, 72% of the variance in the students can be explained by the variance among their parents.
    2. Traits with higher heritability have a lower influence from the environment. Traits that are highly influenced by the environment have a lower heritability.
    3. The response of a population to selection on a trait is determined by the heritability.
    4. Geneticists can look at other relatives besides parents to estimate heritability. Some of the strongest estimates come from comparing identical twins (who have the same genes) to fraternal twins (who share on average half their genes).
    5. Estimating heritability doesn't require us to know anything about the actual genes themselves.

    The last point is very important. We can quantify the influence of genes without knowing anything about which genes even affect a trait. Francis Galton invented the parent-offspring method of estimating heritability, more than twenty years before the word "gene" was coined. Remembering this helps to remind us that the concept of heritability is limited. It is not a guide to how genes work, it is a simple scale of which traits have a stronger or weaker genetic influence.

    Because it is a proportion, heritability varies only between zero and one. A trait with zero heritability is one for which none of the variance can be explained by the parents' variance.

    Heritability may be very low because the individuals in the population have little genetic variability. For example, an orchard of apple trees may consist of genetically identical individuals that are clones of a single parent tree. The apples (hopefully) all taste the same. But the trees may be very different in height, branch form, and the number of apples. These traits may be strongly influenced both by the environments of individual trees and by chance. By contrast, wild populations of oak trees are not made up of clones. The heights of individual trees are still strongly influenced by environments, but they may also be influenced by differences in genes.

    Study questions: 
    1. Make a list of three traits that have low heritability in humans. Why are these mostly influenced by the environment and not genes?
    2. Are there environments inhabited by human populations that would make the influence of genes seem to be less on some traits?
    3. Suppose that the linear regression between midparent and offspring for a trait has a slope of 0.56. What would you estimate as the heritability of this trait?
  • The normal distribution and anthropometrics

    Wed, 2011-09-07 12:31 -- John Hawks
    Synopsis: 
    A combination of random genetic and environmental factors cause individuals to cluster near the average for many traits we measure.

    Organisms within a population are variable; they are not all the same. When you measure a lot of organisms, you begin to notice that you can predict some things about their variation.

    For example, stature is a very simple measurement so it may be surprising how much it can tell anthropologists a lot about individuals and populations. Stature changes as an individual grows and matures, and as she ages. The rate at which stature changes during growth reflects health and nutrition. The average stature in different populations reflects their environment and their evolutionary history. Stature can even tell us about ancient climates and migrations of peoples in the past.

    To get at such interesting information, anthropologists have to understand how stature varies within a single population of people. This involves understanding some statistics.

    For many traits that we measure including stature, a simple rule is at play: Most individuals will be near the average, and few individuals will be very far from the average. Here's a plot called a histogram, showing the stature of female students from my course last year:

    Female student stature histogram

    A histogram of female students' stature in my Anthropology 105 course. The curving line represents the normal distribution with the same mean and standard deviation as the women in the class.

    In this plot, the x axis is stature (body height) in 5 cm increments. The height of each bar represents the number of women who have statures within that 5 cm bin. For example, 28 women had statures between 160 and 165 cm, but only 4 women had statures between 145 and 150 cm. No one was shorter than 145 cm, and no one was taller than 185 cm. The histogram shows how many individuals fall in each bin, a quick way to see the distribution of the observations.

    The mean stature of women in the class was 163.4 cm. The standard deviation of their statures was 6.7 cm. The standard deviation is how different women were on average from the class mean. How often are women far from the average stature? We can make another histogram to show this for the class:

    Most of the women had statures between 157 and 170 cm — in other words, within one standard deviation of the mean. As we look farther and farther from the mean, we see fewer and fewer women with statures that extreme.

    Patterns like this one are very common in natural populations. It is in fact so common that we call it the normal distribution. Most individuals have traits near the average, and we see fewer and fewer individuals as we look far from the average. In that sense, there's something about the individuals in a human population that seems like it's not random: If you draw any individual by random chance and measure him or her, you're more likely to find a person near the average height than far from the average. You can't predict the stature, but if you guess the stature will near the average, you'll be right a good fraction of the time. It's not like flipping a coin or rolling a die.

    Why does this pattern emerge?

    Suppose we have a fair die, and roll it once. We should have an equal chance of rolling a one, two, three, four, five or six. If we roll the die many times, we should start seeing around the same proportion of ones as we see sixes, fives, and threes. This is one way to look at random chance: Each outcome has an equal probability of happening, and we can't predict which will happen in any given trial.

    Now suppose we take two rolls of the die and add them both together. Our trials are still random. We might roll a one and then a three, or a five and then a two, or two sixes. But when we look at the sum of the two numbers, we see a pattern begin to emerge. You will see a lot of sevens, fewer threes and nines, and very few twos or twelves. In other words, you can begin to predict that the outcome will be near the average.

    Why is this? There's only one way to get a sum equal to two: You have to roll two ones, "snake-eyes". But there are lots of ways to get a seven. You can get a one and a six, or a six and a one. You can get a three and a four, or a five and a two. In fact, there are twelve different ways to get a seven, and if you roll many pairs of dice, you'll get on average twelve times as many sevens as twos.

    In dice, if we combine two different random events, we will be more likely to see an outcome near the average than at an extreme. If we roll ten dice, or fifty, we become more and more likely to see an outcome near the average.

    A similar principle applies in biology. A person's body grows by a series of genetic and environmental events. Within a population, some of the factors that affect growth are random. People vary in the combination of genes they carry because of the random chance of inheriting them from their parents. And people vary in the environments they experience because of the random chance of who they are and where they live. A person's stature, in other words, is affected by many, many influences. Some genes may tend to make stature a little taller, others make it a little shorter. Some foods tend to make stature a little taller, others a little shorter. The combination of all these things determines how tall a person will be, and when we consider a population of people all together, they will tend to be clumped near the average.

    This is why many traits follow the normal distribution. But there are exceptions. A person's ABO blood type, for example, is a trait that has essentially no influence from the environment and is entirely determined by a combination of alleles for one particular gene. For blood type, there is no normal distribution in humans. We are A, B, AB, or O, period.

    Also, some biological factors may be very large in their effects. When we look at human stature, the largest single influence on stature is sex. Males average taller than females. Like most things, this is not an absolute — some tall women stand above the average man. But the difference between the sexes is enough to really change the distribution of stature in the population. When we look at females alone, we see a normal distribution, but when we mix individuals of both sexes, the pattern is much flatter. People are less clustered near the average, because half of them average taller and half average shorter.

    Study questions: 
    1. Can you think of some other biological traits that do not vary according to a normal distribution?
    2. If you look at individuals within a single family, should they also have statures that vary according to a normal distribution? Why or why not?
    3. What is the importance of the standard deviation when we interpret variation in the population?
  • Predicting stature from bone measurements

    Tue, 2011-09-06 01:00 -- John Hawks
    Synopsis: 
    A laboratory exercise involving measurement of femora and estimation of stature using regression formulae.

    Anthropologists have collected data from many populations in the world, showing the relationship between the parts of the skeleton and body size and stature. The long bones are the most important elements for estimating overall stature, because each of them contributes to a fairly large segment of the body's length.

    We can use regression equations to give an estimate of the stature from a single long bone. These estimates are not perfect — sometimes a person is taller than you might guess from his femur, sometimes shorter. Moreover, the relationship of bone length and stature varies among human populations, because of differences in body proportions. But allowing for error, the estimates of stature from long bone lengths are among the most important pieces of information we can gather in the process of identification.

    What to do: Here you will examine isolated femora, using regression equations to predict stature of the individual.

    1. Determine the sex of the individual. The femur head diameter is a relatively good indicator of sex. If it is less than 44 mm, the individual is likely to be a female. More than 46 mm, and the individual is likely to be a male. In between these values, you may need more information — either from the rest of the skeleton or from the size and robusticity of the femur itself.
    2. Measure the maximum length of the femur. This measurement is taken using the osteometric board, and represents the maximum distance from any points on the proximal and distal ends of the bone. Take your measurement in centimeters.
    3. Apply the correct regression equation. These are specific to sex and race. The femora at this station come from donated anatomy collections from the early 20th century, and represent people of European ancestry. The male and female regression equations for this population are listed at right.
    Study questions: 
    1. Applying regression equations to estimate stature is a primary component of forensic investigation of the skeleton. If you found a femur, what would you do?
    2. Why do the equations vary between males and females?
  • Measuring stature and proportions

    Fri, 2011-09-02 01:07 -- John Hawks
    Synopsis: 
    An exercise in some basic anthropometric measurements

    At this station, you'll be taking some measurements of your body. This is a graded exercise, but we expect that you'll have no problem getting all the points for this one!

    • The computer station has a spreadsheet to enter your measurements. You'll be using a stature board to measure your full height.
    • We're also interested in your parents' heights because the lecture will use the class statistics to illustrate concepts of inheritance. If you know the heights of your biological parents, we'd like you to fill them in the form. Participation in this part of the exercise is voluntary, and if you don't know your parents' heights or don't want us to know them, feel free to leave those blank.
    • You'll take some measurements of your feet, which you'll compare to your footprints at the next station. Fill in the length and breadth of your foot, and your shoe size. We'll be looking at the correlations between these measurements and stature.
    Study questions: 
    1. Are you the same height as your parents? Are you between them, more like one or the other, or possibly taller or shorter than both?
    2. How different do you think the average height of men and women is in our population?
    Study terms: 
Subscribe to stature

Neandertals

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Malapa

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