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Biology 30 Module 3 Reproduction and Genetics Lesson 11 Types of Reproduction and Development: Single Organisms to Vertebrates Copyright: Ministry of Education, Saskatchewan May be reproduced for educational purposes Biology 30 53 Lesson 11 Biology 30 54 Lesson 11 Lesson 11 Reproduction: Cells and Organisms Directions for completing the lesson: Text References for suggested reading: Read BSCS Biology 8th edition Sections 6.6-6.8 7.3-7.6 Pages 133-139, 146-152 OR Nelson Biology Pages 509-531, 563-564 Study the instructional portion of the lesson. Review the vocabulary list. Do Assignment 11. Biology 30 55 Lesson 11 Vocabulary amniocentisis amniotic egg blastocyst chorion chorionic villus biopsy epididymis estrogen external fertilization feedback fetus FSH germ layers gestation internal fertilization LH menopause menstruation morula ovaries Biology 30 oviducts oviparous ovoviviparous ovulation progesterone semen teratogens testes testosterone tuballigation ultrasound umbilical cord uterus vagina vasdeferens vasectomy viviparous yolk sac zygote 56 Lesson 11 Lesson 11 – Reproduction and Development of Vertebrates Introduction In plants and other organisms below the vertebrate level, reproduction of new individuals is commonly accomplished by asexual methods or a combination of asexual and sexual phases. This last form is known as alternation of generations. In the vertebrate groups, the haploid (n) phase is represented only by gametes (eggs and sperm). There is neither multicellular development of haploid cells nor a gametophyte stage which eventually produces gametes. For this reason, vertebrates are considered to lack alternation of generations and to reproduce entirely by the sexual method. This is also evident in some other taxonomic groups, such as arthropods. Relying almost exclusively on sexual reproduction to produce new organisms, species also developed the characteristic of individuals being female or male, but not both. While hermaphrodites are common in less complex multicellular organisms, they are not numerous in arthropods or vertebrates. Appearances of hermaphrodites in these groups tend to be accompanied by physical (structural) and behavioral abnormalities. The development of separate sexes in sexually reproducing species introduced a number of difficulties. Some of these relate to actions of trying to successfully bring eggs and sperm close enough together so that fertilization could occur. After fertilization, other possible difficulties arise. These have to do with the successful development of a zygote into an independently functioning individual. This lesson will look at reproduction as it applies to vertebrates, from pre-fertilization to the hatching or birth of young and what conditions face the young of various groups. Biology 30 57 Lesson 11 After completing this lesson you should be able to: • identify some of the difficulties of organisms in carrying out successful sexual reproduction and conditions which can increase the success rates of external and internal fertilization. • explain some of the different adaptations among species which increase the chances of survival for the young after they are hatched or born. • discuss some pre-fertilization conditions and their significance. • name and explain the four major ways in which embryos could develop among different groups. • compare amniotic egg of birds and reptiles with the structures that form in the uterus of a pregnant mammal. • describe some of the features related to viviparous reproduction, with emphasis on humans. In particular, · · · · · female and male structural differences. female reproductive cycles. production of semen the fertilization process. the establishment of an embryo inside the uterus. • summarize some of the stages in embryological and fetal development and the general birth process most placental females go through. • discuss the biofeedback mechanisms involved in the reproductive cycle. • discuss the relationship between diet and health of mother and the development of the fetus. • describe how the use of hormones in birth control pills alter the reproductive cycle. • discuss some other aspects related to human reproduction, such as · birth control · amniocentesis · ultrasound Biology 30 · sexual diseases · fertility drugs 58 · genetic screening · in vitro fertilization Lesson 11 Requirements for Successful Sexual Reproduction In biological terms, the sexual process relates specifically to an action where two reproductive cells or gametes fuse together. These gametes, haploid in number, combine to form a diploid zygote which can then undergo mitotic divisions and differentiations in forming another organism. One of the initial requirements of successful reproduction is to bring the different gametes close enough together so that the fusion process can take place. Critical to this is the fact that individual eggs and sperm often do not have long survival times. In the process of oogenesis, the first meiotic division produces a large cell and a smaller polar cell or body. The smaller polar body may or may not disintegrate immediately. The next meiotic division involving the larger cell again results in a large cell-small cell combination. The larger cell remains as the egg. Having more cytoplasm from the unequal divisions, this cell or egg can sustain itself for a longer period of time. However, in most instances, this "longer" period of time is still only a few days. Spermatogenesis goes through similar division processes, except that four, equally-sized cells or sperm result. This leaves each sperm with very little cytoplasm so it is incapable of sustaining the cell for very long. Sperm develop flagella and are capable of swimming movements. This energy-draining action contributes to the very short life spans for the sperm of vertebrates. These characteristics of eggs and sperm therefore make it important that the formation and release of the two types of gametes be timed so that they occur almost together. Also important is that the sperm be placed close enough to the eggs so that they may meet quickly. A zygote, resulting from a successful fertilization, is the beginning or the initial cell of a new organism. As with any other organism, basic processes and requirements must be satisfied if life is to continue for the zygote and then the embryo. The other major requirements for successful reproduction include: the provisions of a suitable environment a sufficient source of nourishment for the developing embryo until it is ready for hatching or birth. The degree to which different species satisfy or meet these conditions is reflected by the mortality rates of the embryos and the very young of those species. Biology 30 59 Lesson 11 External-Internal Fertilizations For less complex, multicellular organisms, the sexual structures or organs involved in reproduction generally experience only limited developments. The structures could include just the sexual organs or gonads (ovaries and testes) and oviducts and sperm ducts leading from these to the body surfaces. These limited developments are carried on into a number of taxonomic groups and extend into the vertebrate level. Initial reproductive actions could be either one of two processes. 1. Both eggs and sperm can be released to the outside where fertilization may occur; or 2. Eggs can be retained in the body cavities or chambers of one individual and can be fertilized by sperm released from another (as in clams). Water plays a very important role in external fertilization. Unless a sperm cell is directly deposited onto the surface of an egg, there must be a way for it to reach the egg. The method usually involves a fluid medium, which enables the flagellated sperm to swim about until a chance meeting with an egg may occur. It can be rightly assumed that external fertilization is not a very efficient process in bringing an egg and a sperm together. Success rates of just some fertilizations happening increase with the production of very high numbers of eggs or sperm, or both. Such a characteristic is quite common among externally fertilizing species. For instance, each spawning season, fresh and salt water fish can produce eggs or sperm which can number in the millions. Despite this, the reproductive rate can still be quite low. Many eggs are not fertilized, while those which are could be eaten by other organisms or fail to develop in unfavorable conditions. Even after hatching, predation rates upon the young are high. The chances of eggs being fertilized increase as sperm are deposited closer to the eggs. The spawning actions of many fish have females and males side by side or the male following the female so that sperm is released at almost the same time and as close to the eggs as possible. In some amphibians, such as frogs and toads, a male will sometimes grasp the female from the back in an action called amplexus. This stimulates an egg laying action in the female. As the eggs are released from the female's cloaca, the male discharges sperm over them. Water is still needed as a transporting agent and this is one of the reasons why amphibians must still return to water to reproduce. Biology 30 60 Lesson 11 Closer contacts eventually see adaptations in species where males actually transfer sperm into females' bodies. Any action of this type is part of internal fertilization. The "simplest" actions have two individuals coming together and lining their bodies up so that certain body openings are opposite each other. Hermaphroditic earthworms carry out this action so that one individual releases sperm which enter a special cavity or seminal receptacle of its partner, while receiving a transfer of sperm into its own seminal receptacle in return. In other species, a male and female line up to carry out a similar action of sperm transfer from opening to opening. Slight body grooves could serve as pathways in some instances. Some species (spiders) have males using their limbs or walking legs to transfer sperm from themselves to the openings of seminal receptacles in females. In many insects, females and males bring the ends of their abdomens (and abdominal openings) together, allowing for direct sperm transfer to a seminal receptacle. In some situations, sperm can be kept alive in females for months and are used at whatever times that eggs are in the process of being laid. In birds and reptiles, close contact between female and male cloaca permit sperm to pass from one to the other. In these individual groups, sperm are used almost immediately for fertilization as they work their way to the oviducts, rather than being stored. In mammals, further developments exist in addition to the gonads and ducts for egg and sperm production and transfer. Males have special copulatory organs for inserting into females' bodies and depositing sperm. This method of carrying out internal fertilization makes it more likely that a high number of sperm will have been internally deposited, closer to the eggs. Although internal fertilization does take place in some aquatic species, its greatest benefit is to land or terrestrial organisms. Without the medium of water to be used for sperm transfer from one individual to another, external fertilization would not be able to take place. On land, this difficulty is overcome by placing sperm on the surface or inside the body of a female. Even though water is not as important for internal fertilization, fluids still play a part in the process. Body fluids of males and females still provide a means for sperm and eggs to move about inside reproductive canals. Biology 30 61 Lesson 11 Pre-Fertilization What has been mentioned previously should have given an indication that the success of fertilization depends very much on two things. These are: 1. timing in the formation and release of eggs and sperm and 2. having the gametes placed as closely together as possible. Both of these conditions are necessary because of the gamete cells' short life spans, which may average only several days. It is therefore important that females and males find each other and carry out mating behaviors at the proper times. A number of different factors could have influences in bringing individuals together and initiating reproductive behaviors. Hormones play a large part in reproductive cycles. For many species, hormone levels themselves could fluctuate according to photoperiods. Certain periods of light and darkness over a length of time can affect the hypothalmus of the brain and the pituitary gland. These then either increase or suppress the levels of certain hormones which affect the reproductive organs. Females and males exposed to the same environmental conditions should experience body changes which tend to match each others' in terms of levels of development or where they are in the reproductive cycle. Photoperiods and hormone levels set up distinctive annual cycles for many species. Examples: The male members of the deer and goat families go into an autumn rut. This is a reproductive behavior in which they establish territories, actively seek out females and try to establish dominance over other males by rituals or by fighting. The annual reproductive behavior for waterfowl and other birds takes place in the spring. In other species, a number of reproductive cycles could take place over certain time periods. Mice could produce two to four litters from spring to fall. Just as changes in hormone levels initiate reproductive behavior at certain times, they also cause it to decline and stop. Many species will even stop producing eggs and sperm and are, in effect, sterile for certain periods of time. Male deer or bucks, at times other than the fall rut, are usually in this condition. Fish and many birds also experience this after spring spawning or egg laying. many domesticated livestock and pets have lost much of the behaviors to breed only at certain times of the year. Horses and cattle can breed and reproduce at any time of the year, although there is still a tendency to follow a certain yearly pattern. Biology 30 62 Lesson 11 Courtship behaviors often arise as a result of certain hormone levels. Largely inherited or innate, these behaviors can be used to find possible mates at a reproductive stage where breedings can occur. Songs of male birds are not only intended to warn other rivals, but are used to attract females. Displays of brightly colored plumage, "dancing", head bobbing or other actions can all do the same thing. The importance of courtship behaviors to successful fertilization is especially noticeable in some groups which follow very ritualistic behaviors. Individual pairs of some fish, birds and other organisms have series of responses and counter responses made by females and males. If one of the partners is not reproductively ready and fails to carry out a particular action in a sequence, the entire courtship ritual usually stops at that point. The other partner may start all over again at the beginning or lose interest and move away. Should a female and male be at a stage where egg(s) and sperm are fully developed and ready for fertilization, it is more likely that a proper sequence of actions would be followed through. This would be culminated by a mating or spawning process where fertilization could occur. Many courtship behaviors are based on vision. Displays are intended to be seen by other individuals and reacted to. The kinds of reactions made would determine the sequence of events to follow. But sight is just one of a number of possible senses which could be important to reproductive processes. Pheromones, or chemical "messengers" released from bodies, are used to attract potential mates through their sense of smell. Females of large numbers of species, in a state of reproductive readiness, release chemicals or body odors which attract males. Male odors could also stimulate some females and initiate or accelerate their reproductive behaviors. Sounds and hearing serve as other forms of communication between individuals. Even touch or contact can be used as a means to either attract or repulse other members of a species. It should be emphasized again that all of the conditions mentioned to this point, as well as others, could play a part in bringing females and males and then eggs and sperm together. A successful fertilization is one of the first steps to a possibly successful reproductive process. Variations in Fertilization and Embryo Development Vertebrate reproduction can follow one of four major patterns: External Fertilization Internal Fertilization – Oviparous Internal Fertilization – Ovoviviparous Internal Fertilization – Viviparous Biology 30 63 Lesson 11 External Fertilization – Aquatic Development External fertilization is carried out primarily by fish and amphibians. Their eggs are fertilized externally almost simultaneously as they are being laid. A female and male are usually close together when these actions are taking place. The eggs often have a surrounding protein material which soon absorbs water and swells to form a jellylike coating. As well as holding the eggs together, this coating helps to protect the eggs and also helps to maintain a more stable temperature around them. A supply of yolk in each egg sustains the developing embryo for the one to three week time period it normally takes from egg laying to hatching. Aquatic development is necessary for such eggs as an exposure to air would dry and kill them. In addition, respiration by means of gills is common to most species – at least for part of their life cycles. Internal Fertilization – Oviparous Development (Amniotic Eggs) Organisms which lay eggs for external incubation are known as oviparous. Depositing eggs for incubation on land could happen only when the eggs are covered with shells relatively impermeable to water, but still allowing gases to pass through. This would permit an embryo to develop within an enclosed, watery environment, despite being exposed to air. Such an adaptation appeared in birds, reptiles, the duckbill platypus and spiny anteater. However, enclosing an egg in a leathery or hard shell not only stops the movement of water, but also prevents the passage of sperm. Therefore, these groups must experience internal fertilization before the egg shell is formed. For the majority of organisms, sperm transfer is accomplished when females and males come together and establish contact with their cloaca. Cartilaginous fish, such as sharks, also carry out internal fertilization. Sperm transfer occurs along the grooves of fin-like claspers of a male shark to the female. Once an egg has been fertilized near the upper end of an oviduct, it begins moving down. Modified regions along the oviducts then form the shells around the eggs as they move through. In birds, the process from fertilization to when the egg is actually laid could take three or four days. Biology 30 64 Lesson 11 Shelled eggs show the first appearance of membranes important to embryo development. Just inside the shell is the chorion. The chorion is a membrane richly supplied with blood vessels. This is where exchanges of oxygen and carbon-dioxide between the shell and the embryo occur. Also present is a membrane around the embryo called the amnion. The amnion contains the amniotic fluid which serves as a protective cushioning agent for the developing embryo. Also contained within the chorion is the allantois and yolk sac. The allantois, like the chorion, is also involved in gas exchanges with the embryo. Another function is to serve as an enclosure for waste material from the embryo. These toxic wastes are largely converted to insoluble, less harmful uric acid which is then held within the membrane. The yolk sac membrane contains the supply of food necessary to sustain the developing individual until hatching and, in some instances, for a short time after. In summary, an amniotic egg with its membranes has a number of ways of increasing the success of land reproduction. An external shell reduces or stops the loss of water, preventing an embryo from drying out. The nature of the shell and membranes enables gas exchanges to occur between an embryo and the outside. A supply of food enclosed within membranes serves to nourish the developing embryo. Membranes also remove and store its wastes. Biology 30 65 Lesson 11 Internal Fertilization – Ovoviviparous Development Organisms which fertilize eggs internally and retain the eggs within their body until after hatching are Ovoviviparous. Ovoviviparous development is prominent in some fish and reptiles (guppies, garter snakes...). The eggs are fertilized internally and have membranes or shells formed around them, and are retained in the females’ body until after hatching. The hatched young are then expelled from the mothers' bodies. From the time that eggs are fertilized to the time that the young are born, a mother's only role is that of providing a more stable and secure environment for them. Nourishment for a developing embryo comes from stored yolk within the egg. Most young are independent and go their separate ways right after birth. In some species, young have to be particularly careful as their own mothers could turn cannibalistic and devour them. Internal Fertilization – Viviparous Development Internal fertilization and incubation, with dependence on the mother after birth is Viviparous development. Most mammals give birth to their young after they have experienced internal development with some degree of nourishment provided by a mother's body. This form of reproduction is characteristic of marsupials and placentals. Among the marsupials, the period of internal development can range from 8 to 40 days for various groups. Most of the nourishment comes from a small quantity of yolk. A yolk sac partially embeds into the lining of a mother's uterus to provide for some movements of nutrients and wastes between mother and embryo. Development is still very incomplete when a young organism is born. After birth, it must make its own way through the mother's fur to her marsupium or pouch and, once inside, fasten onto a nipple. Development is completed inside the marsupium. The particularly difficult journey from birth canal to marsupium and then a somewhat precarious existence inside the pouch places marsupials at a reproductive disadvantage when compared to placentals. Biology 30 66 Lesson 11 Initial embryonic beginnings of marsupials and placentals are similar. The manner in which placental embryos finish their developments gives this group a decided survival advantage over marsupials. In placentals, embryos initially have only a very limited amount of yolk material which is quickly used up. However, the membranes of the yolk sac and allantois (amnion) are modified so that they form a connection between an embryo and the uterine lining of the mother. This connection is called the umbilical cord and the area where the end of it embeds into a portion of the uterus is the placenta. Blood vessels from the mother and the embryo are close to each other in the placenta, but there is no direct connection between the two circulatory systems. Nutrients and wastes pass between them by diffusion through the lymph between the capillary systems. The placental connection enables a young individual to be fairly well formed at birth, after a longer period of internal development. The membranes in mammals are similar to those found in the amniotic egg of birds and reptiles. Insert image comparing membranes of birds and humans if desired. Biology 30 67 Lesson 11 Viviparous Reproduction – A Closer Look at Placentals The dominance of embryo development with a placental connection among mammals warrants a closer examination of this reproductive group. Although many of the references and descriptions will be made to humans, it should be kept in mind that structures and processes are very similar among placentals in general and that references could just as easily be made to other specific groups. Female-Male Structural Differences Female Reproductive System The major structures in a placental female's reproductive system are a pair of ovaries, oviducts (also called fallopian tubes) leading from these to an enlarged uterus, and a canal or vagina leading from this to the outside. Image by Conruyt A separate tube, the urethra, leads from the bladder to a separate opening just in front of the vaginal opening. In primates and some other placentals, the single uterus could be considered as an enlargement and then joining of the two fallopian tubes or oviducts. In other placentals, especially where multiple young are common (as in rodents), the enlargements of the two oviducts or uterine tubes remain separate until close to the terminal end or vagina. Embryos could develop in both of the uterine tubes or "horns" in such organisms. Biology 30 68 Lesson 11 Male Reproductive System In males, a pair of testes has many seminiferous tubules that are lined with cells that undergo meiosis to produce sperm. (The production of sperm is referred to as spermatogenesis.) These tubules lead to a coiled mass, the epididymis (contained in the testes), where sperm cells mature and are stored until ejaculation or release occurs. image by Xiong Chiamiov Just as in a female, a duct network is important in the transfer of the sperm, or male gametes. Upon release, the sperm move out of the epidiymis, and into the vas deferens tube. This tube circles near the bladder and then unites with the urethra before continuing as a single tube which transports both urine or sperm to the outside. Along the way, two seminal vesicles, a pair of Cowper's glands (also called Bulbourethral gland) and a prostate gland, all add fluids to the duct network. These fluids, along with the sperm, form the semen which a male ejaculates or releases during mating. Ejaculation occurs from a penis, a structure containing three cylindrical bodies of spongy tissue located along the sides and top. There are many blood vessels in this spongy tissue and when the vessels dilate, blood enters the penis. As this happens, pressure increases on veins leading out and this slows or stops the movement of blood out. The two combined actions cause the penis to lengthen and become firm enough to penetrate the vagina so that ejaculation can occur within the female body. Biology 30 69 Lesson 11 Structural Differences The male duct network has two major differences from those of females 1. In females, the network is open in that oviducts are not directly connected to the ovaries, but are separated by an open space. The male ducts are closed or complete from the testes to the body surface opening. 2. In females, the urinary and reproductive canals remain separate from one another and have their own separate external openings. However in males, the urinary and reproductive canals are united, having one external opening. Hormonal control of Reproductive Cycles Hormone productions and concentrations are responsible for establishing female reproductive cycles. For many placental females which are reproductive during only part of a year, the same day-night characteristics or photoperiods which affect the males also affect them. This increases the likelihood of both sexes being fertile in the same time period. The hypothalmus of the brain and the pituitary gland are the major agents setting up a reproductive cycle. The hypothalmus directs the pituitary in its production of FSH (follicle-stimulating hormone) and LH (luteinizing hormone). These two hormones are responsible for the production of eggs and sperm. In a male, after puberty, these hormones (FSH and LH) are responsible for the formation and release of the steroid hormone, testosterone. Testosterone promotes sperm production in the testes and the development of secondary sex characteristics. As well, testosterone accelerates or promotes the development of extra muscle tissue. This last effect is a reason some athletes have used this steroid. In a female, FSH initiates the maturation of an ovum (egg cell) inside a fluid-filled sac or a follicle in the ovary. FSH & LH stimulate the follicle to secrete estrogen which in turn causes an increased blood supply and a build-up in the lining of the uterus. This continues for approximately 14 days. Just before the 14th day there is a sudden increase (caused by positive feedback of estrogen on the pituitary and hypothalamus) in LH causing the follicle to burst and the ovum to be released. This is called ovulation. LH stimulates the ruptured follicle to develop into the corpus luteum. The corpus luteum releases estrogen (stimulated by FSH) and progesterone (stimulated by LH). (It becomes a temporary gland releasing hormones.) Progesterone acts to cause more build-up in the uterine lining. The increased levels of estrogen and progesterone create negative feedback on the hypothalmus and the pituitary so that FSH and LH levels fall. This stops development of other follicles. If fertilization does not occur, the corpus luteum Biology 30 70 Lesson 11 begins to disintegrate and the levels of estrogen and progesterone drop. This decreases the blood supply to the uterine lining and menstruation begins. If the egg is fertilized and does establish itself in the uterus, developing membranes secrete hormones which maintain the uterine lining. Female Hormone Cycle INSERT DIAGRM OF HORMONE CYCLE HERE The female reproductive cycle is an excellent example of feedback regulating certain body conditions and trying to maintain a balance according to particular situations. Both positive feedback (where a particular action results in an effect which further intensifies the action of the first.) and negative feedback (where the product of one action will begin to suppress the action of the first) are demonstrated in the female reproductive cycle. For many hoofed placentals, such as deer, elk, the female reproductive cycle repeats about once a month during the breeding season. In some rodents such as rats and mice, it can repeat every few days, unless an impregnation takes place. In other species, the cycle may repeat in 2 to 4 weeks, but it may only happen two or three times in a limited breeding season. Many female placentals will show a marked change in behavior around the time of ovulation, when an egg can be fertilized. In this period of sexual excitement or estrous, a female may actively seek out a mate and initiate breeding behavior. Biology 30 71 Lesson 11 Primate females (including human) experience regular reproductive cycles throughout the year. These menstrual cycles keep repeating unless interrupted by pregnancy. The human menstrual cycle is shown in the following illustration. It is based on 28 days, which is common to many individuals. However, it should be kept in mind that cycles can vary from 21 days to 48 days for others. Even an individual can experience fluctuations in her cycles. The Menstrual Cycle - Centre Interactions among the nervous system and several organs, glands, and hormones regulate the cycle. Note the graph of the rise and fall of hormone levels in the regular monthly cycle. (Should be familiar with this.) * follicular phase – The development of the follicle occurs. Lasts from Day 6 to 14. ** luteal phase – occurs from day 15-28 of the menstrual cycle. The corpus luteum develops. It produces progesterone which causes the buildup of the uterine lining. By Lyrl Ovulation or egg release in many human females takes place about midway through a menstrual cycle – about day 13 or 14, with a possible variation of a couple of days either way. A sharp increase in LH (positive feedback of estrogen) appears to initiate the release of an egg from a follicle. If fertilization does not occur within 12 to 48 hours in the fallopian tube, the egg will degenerate and be resorbed by the body. Menstruation will follow. With age there are fewer functioning follicles, leading to a decrease in progesterone and estrogen. This decrease leads to menopause, and ovulation and menstruation cease to occur. Biology 30 72 Lesson 11 The Fertilization Process Some of the different factors which could lead up to a mating action were described in an earlier section. Hormones play a major part. However, different senses such as sight and smell, and also various behavioral actions by a female and male could influence the timing of a mating action. When a female is receptive, a male will deposit anywhere from millions to billions of sperm into the vagina at each ejaculation of semen. The seminal fluid from the seminal vesicles, Cowper's glands and prostate gland in which the sperm are mixed serves a number of purposes: It actively transports the sperm upon initial ejaculation and later it provides a medium in which the flagellated sperm can swim. In many females, the vaginal and uterine environments are quite acidic, which is harmful to sperm. Seminal fluids lower this acidity and give the sperm a better survival chance. Seminal fluid has a sugar base and this also prolongs the lives of sperm by supplying them with some energy. Despite the high numbers, the mortality rate among sperm cells is extremely high. All sperm may be dead in less than a day. On the other hand, some could survive up to five or six days in the female reproductive tract. The cervix is a major obstacle to many swimming sperm. This is a narrowing between the uterus and the vaginal canal. Many sperm fail to get past this point. Some female bodies could produce an extra amount of mucus which accumulates in this area and stops sperm movement. Those sperm which do get past the cervix continue swimming in the uterus and into the oviducts or fallopian tubes. Ovulation of one or more eggs from an ovary, or both ovaries, does not release them directly into the oviducts or fallopian tubes. A space between the fallopian tube and ovary requires that eggs somehow be drawn into the funnel-like opening of the tube. This is accomplished by cilial action and fluid movements. It is in the fallopian tube that an egg is usually fertilized. The process must take place within several days of the egg entering the tube, otherwise it will degenerate. Fertilization in humans is most frequent when sperm enter the fallopian tubes within a time span of anywhere from 2 days before, to a day after, ovulation. An egg in a fallopian tube could be fertilized in as short a time as 15 minutes from the moment that ejaculation of semen into a female's body takes place. The limited survival times of both sperm and egg cells make the timing of actual mating actions very important in determining reproductive success. Biology 30 73 Lesson 11 Most eggs are fertilized when they are located at the top, or near the beginning, of the oviduct or fallopian tube. Each egg has a protective membrane around it. An enzyme carried by the cap at the top of the sperm cells is needed to help break down this membrane to permit entrance. Once a single sperm has entered the egg, a chemical barrier to all other sperm is established. The sperm which was successful in entering the egg loses its tail and combines its chromosomes with those of the egg. This new cell, with the normal diploid chromosome number, is the zygote. Establishment of an Embryo in the Uterus Successful fertilization does not necessarily ensure successful reproduction. A fertilized egg or zygote takes about two to three days to move the rest of the way through an oviduct before entering the uterus. During the movement, the zygote has been dividing mitotically and also forming a chorion membrane around itself. This membrane produces and releases enzymes which help clear the way to the uterus. In the uterus, the enzymes then enable the embryo to work into the thickened uterine wall. A successful implantation of the embryo into the uterine wall means that a successful pregnancy will have been established. Embryology and Fetal Development Almost as soon as an egg is fertilized to become a 2n zygote, mitotic divisions and cleavage begin dividing it into a mass of smaller cells. These mitotic divisions go on as the fertilized body makes its way along the fallopian tube to the uterus. When the embryo reaches the uterus and is a mass of approximately 60 to 80 cells, it is called a morula. Some variations exist among species after this, with respect to the cell numbers, cell arrangements and yolk content. In general, the next stage produces a rounded layer of cells with a hollow interior. This is the blastula or blastocyst (as it is more commonly called in mammals). Implantation into the uterine wall begins to occur sometime around this stage. Biology 30 74 Lesson 11 INSERT DIAGRAM HERE Development from Ovulation through Implantation As the embryo embeds itself into the uterine wall, its cells continue to divide, with part of the cell layer folding inward. This marks the beginning of a gastrula stage, in which an embryo develops distinct germ layers: ecotoderm, endoderm and mesoderm. A third layer of cells forms when cells split off near the junctures of the ectoderm and endoderm, to form the mesoderm. Biology 30 75 Lesson 11 The three germ layers which result are significant in that each will produce, or differentiate into, the different kinds of tissues and organs which will make up a body. Ectoderm cells differentiate into skin cells, nerves, brain and spinal cord. Endoderm gives rise to digestive structures. The mesoderm produces muscles, bones and the major body organs. From the blastula stage onward, other developments take place. One of the more important of these is the development of extraembryonic membranes and, in particular, the placenta and umbilical cord linking the embryo to the uterine lining. The placenta, as part of its functions, acts as a temporary endocrine gland. Soon after it begins forming, it starts to release (human) chorionic gonadotropin hormone or HCG. This hormone stimulates the corpus luteum and its production of progesterone which, in turn, helps to maintain the uterus and pregnancy. The early presence of HCG is the basis of a home pregnancy test. A kit can be used to determine whether or not urine contains the hormone and if a pregnancy exists. After a placenta has become established, it will begin producing and releasing progesterone itself. The placenta and the umbilical cord permit nutrients and waste materials to pass back and forth between the mother and embryo(s). Exchanges between mother and embryo take place at the placenta. There is no direct joining of the two circulatory systems. Rather, substances diffuse back and forth through the lymph between the two systems. The graph on the right shows the hormone levels as they are during a pregnancy. Compare this graph back to the graph on the top of page 72 that shows the hormone levels in the regular monthly cycle. (You should be familiar with this graph as well.) Biology 30 76 Lesson 11 Occasionally, in the blastula or early gastrula stages, embryonic cells could divide roughly in half and then go on developing as separate individuals. This action gives rise to identical twins or, if divisions are not complete, Siamese twins (joined together). Human embryological development is generally studied on the basis of trimesters, or three month developmental periods. First Trimester: During the first trimester, most human embryos will be forming all their limbs and other organs. For this reason, the mother's health, diet and whatever enters her body in the early stages of pregnancy can have significant final results. At the end of the first trimester, all the major body organs, systems and external body features will have been formed. Usually weighing about one gram, a 2 to 3 cm embryo begins to be called a fetus before the first trimester is over. Second Trimester: Remaining growth in the uterus largely consists of adding more cells to the tissues and organs which were developed in the embryonic stages. Leg, arm and general movements of the fetus begin about the fourth month of pregnancy and increase in frequency and force to about the sixth month. At the end of this second trimester, the baby is "almost" ready to be born, with all systems complete. Third Trimester: Most vital systems are still frequently too weak to function independently if a premature birth took place at the end of the sixth month. Movements tend to decrease after the second trimester as the fetus' increased size gives it less room to maneuver in a more confined space. Large weight gains take place in the last three months, slowing only in the latter stages of the ninth month. The last number of weeks of a pregnancy have a fetus assuming a lower position in the uterus with the head normally positioned at the lower end, near the cervix. During this time the placenta will usually have started to degenerate as well. Biology 30 77 Lesson 11 Fetal Health The development of a fetus is dependant on the health of the mother. As well as providing beneficial substances, such as vitamins and minerals, to the baby, a mother can transfer harmful substances as well. These harmful substances can greatly affect the embryo’s development. Whatever the mother ingests or inhales can end up in her blood, which in turn can be passed to the fetus. The developing fetus is at greatest risk during the first 9 weeks of development when the organs are developing. INSERT DIAGRAM HERE ILLUSTRATING THE EXCHANGE Placenta and Substance Exchange Substances harmful to the fetus, causing structural abnormalities, are termed teratogens. Teratogens can include: cigarette smoke – may lead to underweight babies alcohol – can affect the fetus’s brain and nervous system. Alcohol consumption can lead to babies which have Fetal Alcohol Syndrome, FAS. Children with FAS can have smaller weight, height, and head size, malformations of the head and face, and varying degrees of mental retardation some prescription and over the counter drugs - thalidominde was a drug in the 1950’s that was used to treat morning sickness. This resulted in many babies being born with missing or deformed limbs. Radiation – such as X-rays. X-rays are generally not performed on pregnant women, unless the benefit outweighs the risks Proper nutrition along with avoidance of teratogens is essential for a pregnant women in order to signficantly decrease the chance of having a child with a birth defect. However, some birth defects are due to heredity or other unknown, unavoidable factors. Biology 30 78 Lesson 11 The Birth Process The actual development period or gestation in placentals varies considerably. In rodents, the time from fertilization to birth could be only three to four weeks. Members of the dog family have gestation times of approximately two months. Many other mammals, including humans, have times averaging about 9 months. One of the longer gestation periods is that of an elephant, which could be up to 22 months. The factor(s) which actually start the birth process are still not clear. Whatever the triggering agent(s), hormone levels seem to be part of the action. During pregnancy, estrogen and progesterone hormones are in balance with one another. The first has a tendency to cause contractions of the smooth, uterine muscle. The second, progesterone, prevents contractions. Just before birth, levels of progesterone drop. About this time, the pituitary begins increasing its output of another hormone called oxytocin. The concentrations of oxytocin and estrogen (whose levels now become higher that those of progesterone) finally cause uterine contractions to begin. In certain situations, doctors will sometimes inject oxytocin to induce contractions and labor. The first part of the birth process generally takes the longest. While uterine contractions increase in frequency, muscles of the cervix and vagina relax. This dilation or expansion permits easier passage of the young animal or human baby. Dilation could last from approximately an hour to over fifteen hours. Females who gave birth previously generally have shorter dilation times. The end of the dilation period comes with the passage of a mucus plug which had been in the cervical opening during pregnancy. The amnion membrane will have been ruptured by muscle contractions just before this so that amniotic fluids will be expelled as well. This "water breaking" is followed by the second part of birth, or true labor, in which the fetus is pushed out. This could last from several minutes to over an hour. True labor sees another example of a feedback effect on the body. Usually, feedbacks have negative or inhibitory effects on the initial actions. During this time, there is a positive feedback, where a particular action results in an effect which further intensifies the action and effect. As the baby's head presses against the smooth (involuntary) muscles, the pressure actually causes those muscles to contract even more frequently and with more strength. This effect goes on until the baby is finally pushed out. With a decrease in the pressure on the smooth muscles, the contractions will begin to slow down, although still continuing for a time afterward. Emergence of a young animal or baby could cause the umbilical cord to tear from the placenta still in the uterus. (In humans it is usually cut just after emergence of the baby.) The torn or cut umbilical cord quickly seals so that there is no loss of blood. Biology 30 79 Lesson 11 After developing in a warm, watery world, the newborn will be stimulated to begin breathing by the licking or touching actions of the mother and increasing CO2 levels in its own blood. The final stage of birth is the expulsion of the placenta and any remaining fetal matter. This is accomplished by the contractions of the smooth muscles which still continue, but at diminishing rates. The contractions also help to seal off broken blood vessels in the uterine wall. In a week to several weeks after birth, the mother's uterus returns to normal size. Any tissue damage to the vaginal walls which occurred during birth heal. Hormone levels and concentrations return to conditions as they were prior to pregnancy – unless a mother is actively feeding young with her own milk. In such a situation, there could be higher levels of progesterone in the mother's body. This could delay the return of estrus or menstrual cycles until weaning of the young occurs. Offspring Survival In many populations, mortality rates are frequently highest among the very young – during embryonic development and shortly after birth. Therefore, to consider the success of reproduction, one should also include a time period shortly after hatching or birth. Different species have different adaptations to increase survival chances of their young. For many, parental involvement is very limited. Many fish, amphibians and reptiles deposit their eggs and then leave them. For such species, the poor survival rates among the young are compensated for by very high egg numbers. Eggs are laid by individuals in numbers ranging from thousands into the millions. Predation upon the young could still leave some unharmed. Large numbers could also be an advantage in providing early warning signals to a group. As well, sometimes the scattering action of many individuals can be confusing to predators. Internal fertilization and internal development of embryos offers greater protection for the young. Limited internal space reduces the numbers of offspring produced, but the better protection increases their survival chances. Mothers, and often both parents, offer varying amounts of care and protection to the young. Parental involvement frequently increases as number of offspring decreases. One or both parents could build burrows, nests or other shelters for the young. Protection is provided by the parents acting as decoys to draw predators away or actually attacking them. Biology 30 80 Lesson 11 Offspring themselves can show different survival techniques. Young of many species become quiet and almost completely motionless when there is danger nearby. Some have the added protection of having very little body scent initially, so as not to attract possible predators. Body markings could provide effective camouflage with the environment. Some young can "bluff" very effectively with threatening postures, hisses, loud squeals or snapping and biting actions. Fluffing feathers or holding wings outward, or having body fur rise, have the effect of making some young organisms seem much bigger than what they are. In species with longer gestation periods, offspring usually show a dependency on the parent(s) for longer periods of time. The parent(s) not only provide food and protection, but may teach their youngsters survival skills. Some offspring remain closely associated with their parents until they become sexually mature. Other Aspects Related to Human Reproduction Monitoring Human Reproduction and Embryo Development The births of children with various structural defects or defective body functions, along with their ensuing consequences, have long been a concern to many people. Such concerns have led to efforts to find ways of determining the characteristics of young before they are even born. Currently, there are a number of ways of carrying out some examinations. Biology 30 81 Lesson 11 Ultrasound Imaging Utilizes sound waves which pass through a baby's body to produce an image on a screen. Images or outlines of various body features may indicate possible structural abnormalities. Routinely performed at 16 weeks gestation. Amniocentesis Drawbacks Some individuals express concern with this procedure, due to the lack of information as to what effect(s) sound waves may have on an embryo. Involves the insertion of a syringe through the abdominal wall of a mother and into the fetal sac. A quantity of amniotic fluid, containing some of the embryo's cells, is removed for analysis. Some of these embryo cells are cultured in a special medium until a karyotype can be prepared. A karyotype shows the number of chromosomes present, their sizes and shapes. An examination of the chromosomes may determine any possible abnormality about them and may perhaps indicate some consequence of this. An examination of the amniotic fluid itself is also carried out to see if it contains any unusual proteins. Tests on the cells and fluid are able to identify certain specific disorders which may be present in a developing embryo or fetus. Generally performed only on pregnant females over the age of 35 (risk of genetic abnormailites increases after age 35) Chorionic villus biopsy Involves the insertion of a thin tube through the mother's vagina and into the uterus. A quantity of cells can be removed from the chorionic membrane surrounding the embryo. As the chorion originates from the same cells as the embryo, it has the same genetic component. A karyotype can then be examined Can be performed at about two months into the pregnancy. Drawbacks The risk of a possible spontaneous abortion is slightly higher with this procedure than with amniocentisis Drawbacks: requires care, to avoid any possible injury to the fetus and to avoid a possible spontaneous abortion. not usually performed before the fourth or fifth month of a pregnancy due to the small size of the amniotic sac and small quantity of amniotic fluid prior to that. Since testing is not done until later during gestation, discovery of an abnormality limits the kinds of actions which may be taken to deal with it. Biology 30 82 Lesson 11 The results of tests, such as these and others, are generally positive and confirm normal pregnancies and developments. There are some, however, that require special medical or genetic counselling and possibly specific actions. Embryonic disorders could require such actions as blood transfusions to embryos, having mothers put on special diets or drugs, or various other procedures. Genetic counselling is frequently sought out by individuals who have some knowledge of certain disorders, their genetic backgrounds or their family histories. People may want extra information with respect to such things as: how certain disorders progress; probabilities of their children inheriting or experiencing certain conditions; genetic or diagnostic tests which could be carried out; interpretation of certain results; and, possibly, advice or recommendations on particular courses of action. Birth Control For a variety of reasons, individuals may choose not to have children or to delay having children until particular times. As numerous as the reasons may be, so may the ways of preventing reproduction. Permanent surgical sterilizations are used to prevent the movements of gametes to areas where fertilization could occur by cutting off access within the duct networks. In females, abdominal incisions can be made so that access to the oviducts permits them to be tied off somewhere along their lengths. This tubal ligation prevents the movement of eggs from ovaries to the uterus. INSERT DIAGRAM OF TUBAL LIGATION HERE Biology 30 83 Lesson 11 In a vasectomy, a male can have small incisions made in the sides of the scrotum or sac holding the testes. The vas deferens along the sides are exposed, tied off in two places and then surgically cut between the two tie-offs. This prevents sperm from moving out of the testes. INSERT DIAGRAM HERE Vasectomies and tubal ligations are reversible, but the reversal may or may not be successful. In other male animals, the practice of castration involves the removal of the entire testes. Doing this means that there will be no sperm and also no more hormone production from certain cells located in the testes. The following table summarizes some common methods of birth control. Methods of Birth Control Insert table here if desired that compares methods such as abstinence, vasectomy, birth control pill, tubal ligation, needle injections or implants. Sterility On the opposite end of the spectrum from birth control, is the situation where some individuals or couples are unable to have children. Causes of Female Sterility Causes of Male Sterility blocked fallopian tubes failure to ovulate, due to hormonal imbalance Obstruction in the vas deferens or epidydymus Low sperm count endometriosis Abnormal sperm damaged eggs Biology 30 84 Lesson 11 There are a number of options available to overcome sterility. Often surgical techniques can be used to remove blockages Hormonal problems can also be corrected. Low progesterone could prevent an egg from embedding in the uterine wall. Taking extra progesterone around the time of ovulation would correct this problem. Sometimes, follicle cells have low sensitivities to the hormones FSH and LH. Fertility drugs, which mimic the hormone put out by the hypothalmus to stimulate FSH and LH production, can be taken. Much greater levels of these two hormones could then initiate follicle development. Often, these fertility drugs are too effective and there are multiple follicle developments and multiple births. Low sperm counts could be enhanced by combining a number of ejaculations in a laboratory to prepare one sample with an about normal sperm Other technologies are summarized below. Insert table here that compares AI, IVF, IVM, superovulation, surrogate or cryopreservation if desired. Some of these procedures have moral or "ethical" issues associated with them. A fertilized egg could be implanted into a different female than the one from which it was removed. It could be that a father's identity may not even be known, with donor sperm coming from sperm banks. Individuals could be born without ever knowing the identity of either biological parent. It is also possible to determine the sex of the embryo and to manipulate or to modify it genetically, before implantation. Sexual Diseases (STD’S) Internal fertilization and contact between sexual partners bring with them the possibilities of transferring viruses, bacteria or other parasites between partners. Syphilis (bacterium), gonorrhea (bacterium), chlamydia (bacterium), herpes (viral) and AIDS (viral), are just some diseases. Sexually transmitted, or venereal, diseases increase in frequency where there are multiple partners. Prevention is the most effective way of dealing with such diseases and reducing the number of partners is one preventative measure. Aside from abstinence, the condom is the most effective method for prevention of sexually transmitted diseases. Most diseases can be cured if diagnosed early; however, immunity usually doesn't last. Initial infections may not be noticed or may seem too minor to be of any consequence. Embarrassment can also be a factor in not seeking medical attention. Lack of medical attention and help can have undesirable effects in a number of ways. Without treatment, other partners and even unborn babies are at risk of being infected. Development of a specific disease can possibly lead to sterility, more serious internal damage and even death. Biology 30 85 Lesson 11 Summary Vertebrate reproduction is accomplished through the sexual process, where different gametes fuse together. The sexual process restores the normal diploid chromosome number of the species in a new zygote. Fertilization can be either external or internal. Water for sperm transfer is especially important in external fertilization, but body fluids are also necessary for transporting sperm and eggs when internal fertilization is carried out. Embryonic development can take place within eggs. Some eggs are fertilized externally and develop in an aquatic or moist environment so that they will not dry out. Reptiles and birds have amniotic eggs, with outer protective shells and inner membranes enclosing watery environments. The membranes are important in the movements of nutrients and wastes in and out of embryos. Some ovoviviparous species retain the eggs internally until they hatch. Young are then born to begin independent lives. Yolk material is fairly prominent and is important in embryo developments within eggs. Marsupial embryos have only a limited yolk supply. Young are born only after brief internal developments. A precarious journey must be made by the underdeveloped young to a marsupium or pouch where growth can be completed. In terms of security, vertebrate placentals provide the best conditions for embryo and fetal development. Extraembryonic membranes, modified into placenta and umbilical cord, serve as the lifeline between mother and young until birth. Once viviparous development is complete, youngsters are born into conditions where their parents often provide some degree of support for a time afterwards. Species with longer gestation times and where young are slower to mature, generally show longer parent-offspring relationships. Although differences do exist in reproductive methods, there are basic similarities among all species. Small parts of parent organisms develop into new individuals. These small parts or initial beginnings contain all the necessary material and genetic information that is needed to maintain species survival. In this way, a continuity of life is maintained generation after generation, as long as reproduction remains successful. Biology 30 86 Lesson 11