HOMOLOGOUS STRUCTURES http://itc.gsw.edu/faculty/bcarter/histgeol/paleo2/homol1.htm There are two reasons why a structure might be shaped similarly in two different objects. One is that the structure is analogous, meaning that it performs the same function. The wings of a dragonfly and of a bird are analogous, and, in fact, are also analogous to the wings of a 747. In order to fly an object needs to generate lift. This is easily done (in various ways) by having large flat surfaces project out from both sides of the object. Therefore flying things have such structures. But knowing the function of a structure doesn't necessarily tell us all we want to know about why the structure exists. Part of why a wing exists is to perform a certain function, but part of it lies in the construction of the wing. Perhaps this question is better worded as HOW the wing came to be rather than WHY it came to be. The picture below shows the basic structure of the limbs of several vertebrates. Notice that all the limbs, whether wings, legs, arms, or flippers are built upon the same basic structure. (Diagram from Monroe and Wicander, The Changing Earth, 3rd Edition.) ALL vertebrate limbs are put together this way, regardless of their use. Clearly there is no analogous similarity between a bat's wing and a horse's leg, and the extreme difference in uses of the two makes the underlying similarity seem unusual. These types of similarity are called homologous and are very interesting indeed. Biologists generally find depictions of angels absurdly funny because they have two sets of forelimbs. Bird wings are homologous to human arms, not completely different structures. Angels would necessarily have two tibia, two radii, two ulnae, and more than five sets of digits if they really looked like that, not to mention all the attachment bones that would have to be duplicated in the torso, like shoulder blades and collar bones. Then there would have to be two sets of muscles, a huge sternum and keel, ... Even wimpy angels would be barrel-chested individuals indeed! Pegasus, the flying horse of Greek myth, represents the same basic misunderstanding of anatomy. Organisms have genes that perform two separate functions. One set of genes controls the existence (or lack thereof) of a particular feature. These are called structural genes. The other set controls if, when, and for how long the structural genes are allowed to function. These are called regulatory genes. the diagram below shows the effects of different regulatory gene control on the development of two vertebrate limbs: a human arm and a bat wing. Examine each corresponding bone and compare their relative sizes after growth. There are many examples of homologies among animals and plants. The limbs of vertebrates are not the only obvious homologies in the skeleton -they are the rule rather than the exception. The number of neck vertebrae is constant among mammals, for example. The neck of a giraffe is made of seven bones, just as yours is, and just as a whale's is. In the whale's neck the bones are exceedingly short, in the giraffe they are exceedingly long. One has to wonder why a structure that basically won't bend is made of seven ridiculously short bones rather than one big one, or why a giraffe doesn't have more bones in a ten-foot neck than a person with a tencentimeter neck. Perhaps if they did they wouldn't have to splay their front legs to get a drink of water. There appears to be some constraint upon the way mammal necks are built. 3. Evolution Makes Sense of Homologies http://www.zoology.ubc.ca/~bio336/Bio336/Lectures/Lecture5/Overheads.html Richard Owen (1848) introduced the term homology to refer to structural similarities among organisms. To Owen, these similarities indicated that organisms were created following a common plan or archetype. That is, although each species is unique, the plans for each might share many features, just as the design plans for a Honda Civic and a Honda Prelude might be similar. Nevertheless, if every organism were created independently, it is unclear why there would be so many homologies among certain organisms, while so few among others. It is also hard to make sense of the fact that homologous structures can be inefficient or even useless. Why would certain cave-dwelling fish have degenerate eyes that cannot see? Darwin made sense of homologous structures by supplying an evolutionary explanation for them: • A structure is similar among related organisms because those organisms have all descended from a common ancestor that had an equivalent trait. Ridley uses a specific definition of homology: "A similarity between species that is not functionally necessary. "I interpret this as: "A similarity between species that exists despite several plausible alternative traits that would function equally well." The plasma cell membranes of all organisms, eukaryotic and prokaryotic, are structurally similar, consisting of a phospholipid bi-layer. Many other possible membrane structures exist. The hydrophobic fatty acid tails could be joined. There could be three hydrophobic fatty acid chains. Other hydrophilic groups could be involved besides glycerol phosphoric acid. The similarity of the plasma membrane (as well as other cell structures) suggests that all living cells have descended from an ancestor with a similar membrane structure. One of the classic examples of a homologous structure is the pentadactyl (= five digit) limb. All tetrapods (= four legged) have limbs with five digits, at least at some stage in development. Certain tetrapods lose some of these digits during development, as in the bird wing shown here. But if the bird wing does not need five digits, why do five initially develop in the growing embryo? The most plausible explanation is that while the five digits are not functionally necessary, they represent a genetic artefact inherited from the ancestors of birds. Homologous structures teach us an important lesson about evolution: Evolution works primarily by modifying pre-existing structures. That is, even when two species function in completely different ways, they often use homologous structures to carry out those functions. For example, birds and bats fly rather than run on all fours, yet their wings are modified fore-limbs rather than completely novel structures. Similarly, the stinger of wasps and bees is a modified ovipositor, rather than an entirely new structure. (Explaining why only female bees sting!!) Homologies http://evolution.berkeley.edu/evolibrary/article/0_0_0/lines_05 Evolutionary theory predicts that related organisms will share similarities that are derived from common ancestors. Similar characteristics due to relatedness are known as homologies. Homologies can be revealed by comparing the anatomies of different living things, looking at cellular similarities and differences, studying embryological development, and studying vestigial structures within individual organisms. In the following photos of plants, the leaves are quite different from the "normal" leaves we envision. Each leaf has a very different shape and function, yet all are homologous structures, derived from a common ancestral form. The pitcher plant and Venus' flytrap use leaves to trap and digest insects. The bright red leaves of the poinsettia look like flower petals. The cactus leaves are modified into small spines which reduce water loss and can protect the cactus from herbivory. Another example of homology is the forelimb of tetrapods (vertebrates with legs). Frogs, birds, rabbits and lizards all have different forelimbs, re flecting their different lifestyles. But those different forelimbs all share the same set of bones - the humerus, the radius, and the ulna. These are the same bones seen in fossils of the extinct transitional animal, Eusthenopteron, which demonstrates their common ancestry. Individual organisms contain, within their bodies, abundant evidence of their histories. The existence of these features is best explained by evolution. • Several animals, including pigs, cattle, deer, and dogs have reduced, nonfunctional digits, referred to as dewclaws. The foot of the pig has lost digit 1 completely, digits 2 and 5 have been greatly reduced, and only digits 3 and 4 support the body. Evolution best explains such vestigial features. They are the remnants of ancestors with a larger number of functional digits. • People (and apes) have chests that are broader than they are deep, with the shoulder blades flat i n back. This is because we, like apes, are descended from an ancestor who was able to suspend itself using the upper limbs. On the other hand, monkeys and other quadrupeds have a different form of locomotion. Quadrupeds have narrow, deep chests with shoulder blades on the sides. Organisms that are closely related to one another share many anatomical similarities. Sometimes the similarities are conspicuous, as between crocodiles and alligators, but in other cases considerable study is needed for a full appreciation of relationships. Modification of the tetrapod skeleton Whales and hummingbirds have tetrapod skeletons inherited from a common ancestor. Their bodies have been modified and parts have been lost through natural selection, resulting in adaptation to their respective lifestyles over millions of years. On the surface, these animals look very different, but the relationship between them is easy to demonstrate. Except for those bones that have been lost over time, nearly every bone in each corresponds to an equivalent bone in the other. Studying the embryological development of living things provides clues to the evolution of present-day organisms. During some stages of development, organisms exhibit ancestral features in whole or incomplete form. Snakes have legged ancestors. Some species of living snakes have hind limb-buds as early embryos but rapidly lose the buds and develop into legless adults. The study of developmental stages of snakes, combined with fossil evidence of snakes with hind limbs, supports the hypothesis that snakes evolved from a limbed ancestor. Baleen whales have toothed ancestors. Toothed whales have full sets of teeth throughout their lives. Baleen whales, however, only possess teeth in the early fetal stage and lose them before birth. The possession of teeth in fetal baleen whales provides evidence of common ancestry with toothed whales and other mammals. In addition, fossil evidence indicates that the late Oligocene whale Aetiocetus (below), from Oregon, which is considered to be the earliest example of baleen whales, also bore a full set of teeth. Homologies: cellular/molecular evidence All living things are fundamentally alike. At the cellular and molecular level living things are remarkably similar to each other. These fundamental similarities are most easily explained by evolutionary theory: life shares a common ancestor. The cellular level All organisms are made of cells, which consist of membranes filled with water containing genetic material, proteins, lipids, carbohydrates, salts and other substances. The cells of most living things use sugar for fuel while producing proteins as building blocks and messengers. Notice the simil arity between the typical animal and plant cells pictured below — only three structures are unique to one or the other. The molecular level Different species share genetic homologies as well as anatomical ones. Roundworms, for example, share 25% of their genes with humans. These genes are slightly different in each species, but their striking similarites nevertheless reveal their common ancestry. In fact, the DNA code itself is a homology that links all life on Earth to a common ancestor. DNA and RNA possess a simple four-base code that provides the recipe for all living things. In some cases, if we were to transfer genetic material from the cell of one living thing to the cell of another, the recipient would follow the new instructions as if they were its own. These characteristics of life demonstrate the fundamental sameness of all living things on Earth and serve as the basis of today's efforts at genetic engineering. Examples of Analogies Analogy: Squirrels and Sugar Gliders Beyond being cute and cuddly, flying squirrels and sugar gliders have many striking similarities: big eyes, a white belly, and a thin piece of skin stretched between their arms and legs, a trait which helps them "glide" and remain stable when leaping from high places. But sugar gliders and flying squirrels also have some key differences. Most importantly, they reproduce and bear their babies in fundamentally different ways: • Flying squirrels are placental mammals. Placental mammals spend a long time developing inside the mother's body being nourished by a placenta before they are b orn. • Sugar gliders are marsupial mammals, like kangaroos. Marsupial mammals may only spend a short time developing inside the mother's body and are very tiny when born. After birth, a baby marsupial crawls into its mother's pouch and is nourished by her milk as it continues to grow and develop. Flying squirrels and sugar gliders are only distantly related. So why do they look so similar then? Their gliding "wings" and big eyes are analogous structures. Natural selection independently adapted both lineages for similar lifestyles: leaping from treetops (hence, the gliding "wings") and foraging at night (hence, the big eyes) Analogy: Desert-dwellers Both of these two plants have thick, water-filled branches and sharp spines. You might guess that they are closely relat ed — but they are not. In fact, one "cactus" is more closely related to a common weed, and the other is more closely related to a carnation. Their similarities are analogies — independently evolved adaptations that aid survival in the desert: succulent, waxy stems help store water, and spines provide shade and protect the plant from herbivores. Analogy: Jaws versus Flipper Although their differences are certainly substantial (e.g., sharks are cold-blooded and dolphins are warm-blooded), sharks and dolphins have some undeniable similarities: side fins, a dorsal fin, and a torpedo-shaped body. These similarities are analogies — traits the two lineages evolved independently as adaptations for moving swiftly in the water. Sharks were gliding through the oceans long before dolphins descended from land-dwelling mammals. Dolphins evolved flippers, a dorsal fin, and a torpedo-shaped body as natural selection shaped them for the life of an ocean predator. Analogy: Kings of the Anthill Anteaters live in Latin America and South America, grow to two meters (about six feet) long, and give birth to live young. Echidnas live in Australia, grow to half a meter (about one and a half feet) long, and, as the only close, living relative of the duck-billed platypus, lay eggs. But if you get up close and personal with them, you will find that anteaters and echidnas have some striking similarities: both are t oothless, with a pointy snout, a long, sticky tongue, and sharp, curved claws. All of these traits are analogies — testaments to the selective power of the anthill. These two rather distantly related lineages independently evolved snout, tongue, and claw traits that allow them to more efficiently pillage and plunder an anthill. Analogy: When is a thumb a thumb? Through careful study, biologist can also identify analogies. For example, panda bears have a "thumb" on their hands. They use this "thumb" to hold onto bamboo as they eat. Is the panda's "thumb" homologous or analogous to the thumb on your own hand? Studying the anatomy of panda hands and human hands shows that these "thumbs" must be analogous. When you look at the bones of each, you see that the thumbs are not very similar at all! The human thumb has joints and is made of many bones. The panda thumb is just one bone sticking out of the side of the hand. Furthermore, the panda thumb is the sixth "finger" on its hand! If you watched the hand of a baby panda grow, you would see that the "thumb" develops from a wrist bone. The panda thumb and the human thumb don't grow from the same bones. This is more evidence that they are analogous structures. In fact, the panda thumb is homologous to a wrist bone in humans, and the human thumb is homologous to the first finger in pandas!