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14.3 Factors Limiting Natural Population Growth E X P E C TAT I O N S Differentiate between density-independent and density-dependent population regulating factors, and describe their effect on the growth of populations. Describe various types of interactions among different species of animals and plants, and explain how they affect population growth. Compare and explain the fluctuations of various populations, emphasizing factors such as carrying capacity, fecundity, and predation. Density-independent Factors Temperature or population density Many populations that grow exponentially eventually stop growing quickly, or crash as a result of a very high death rate (recall Figure 14.13 on page 478). Such crashes are frequently the result of abiotic factors such as bad weather. For example, certain species of bark beetles (which cause serious damage to forests in various parts of Canada) grow exponentially until very cold weather in early or late winter kills many of the individuals that would produce the next generation. When winters are mild, or short, enough insects survive to reproduce during the next summer, creating an exponential growth curve. These abiotic limiting factors are referred to as density-independent factors because they affect populations regardless of their density. These factors are effective against small populations as well as large ones. Figure 14.17 illustrates the effect of a density-independent factor. Other density- independent factors include events like floods or droughts. For populations located within a small geographic area, forest fires, hurricanes, or tornadoes can also limit population growth. Figure 14.18 shows a memorable example of a density-independent regulating event. Density-dependent Factors As you have seen, the growth of many populations follows a logistic rather than an exponential growth curve. A population like this is limited by densitydependent factors. The strength with which these factors slow a population’s growth depends on the density of the population. When a population is carrying capacity temperature population crash Time Figure 14.17 Notice that for this hypothetical population, a decline in population size is linked to temperature, a density-independent limiting factor. The decline begins before the population is halfway to its carrying capacity and well before its own density would inhibit its growth. Figure 14.18 A severe ice storm hit eastern Ontario and parts of Québec in January 1998. The devasting ice storm made roads impassable and knocked down power lines, leaving hundreds of thousands of people without electricity, some for several days. In addition, the storm acted as a density-independent limiting factor, causing declines in the populations of maple, birch, and cedar trees, among other species. Chapter 14 Population Ecology • MHR 481 small enough to be far below its carrying capacity, density-dependent factors have no effect and population growth is rapid. Eventually, however, the population reaches a density at which these factors start to have an effect; after this point the population growth slows, and it eventually stops when the carrying capacity is reached. Later in this chapter, Investigation 14-A will give you an opportunity to examine the effect of both density-dependent and density-independent regulating factors on a species of micro-organisms. The remainder of this section describes some specific types of density-dependent factors in more detail. Competition In contrast to density-independent factors, densitydependent factors are typically biotic — they involve living things. As has been described, these living things are often members of the population itself. For example, the carrying capacity of a population’s environment may depend chiefly on the availability of food. When the population reaches the inflection point in its logistic growth curve, there is no longer an abundance of food for each member of the population. Members must now compete with each other for the limited food supply, which becomes even more limited as the population size increases. The result is that the birth rate decreases or the death rate increases, or both (see Figure 14.19), and the population growth slows more and more as the density increases. This type of competition among the members of a population is referred to as intraspecific competition. You have encountered its effects already, since it leads to evolutionary change as a result of natural selection. A higher proportion of successful competitors survive longer and have greater reproductive success; they are said to have higher fitness. This allows them to pass on more copies of their alleles to subsequent generations. The result is a change in allele frequencies within a population — which results in evolution. 10 000 Average clutch size Average number of seeds per reproducing individual 12 1000 100 11 10 9 8 0 10 0 100 Seeds planted per m2 A B 10 20 30 40 50 60 70 80 90 Number of breeding pairs 100 Survivors (%) 80 60 40 20 0 C 482 20 60 100 Density (beetles/0.5 g flour) MHR • Unit 5 Population Dynamics Figure 14.19 (A) A varying number of plantain seeds (a common Ontario weed) were planted in experimental plots. The average number of seeds produced by an adult plant decreased as the density of the plants in the plots increased. (B) Many songbird populations are limited by the amount of food available in their habitat. In some species, the average number of eggs laid by females declines as the density of breeding pairs increases. (C) In some cases, increased population density reduces the ability of individuals to survive rather than (or in addition to) reducing reproduction. This is true for laboratory populations of flour beetles. This graph shows the percentage of beetles that survive from egg to adult, in relation to the population density of the beetle. In addition to its role in evolution, intraspecific competition is an important density-dependent factor limiting the growth of many populations. Members of a population may compete for a variety of needed resources, including food, water, sunlight, soil nutrients, shelter, or breeding sites (see Figure 14.20). The effect is always the same — a reduction in the population’s growth rate. Density-dependent limiting factors can involve interactions between two or more populations as well as interactions within a single population. For instance, two species with similar habitat requirements (for example, see Figure 14.21) may compete with each other for soil nutrients, food, or other resources found in that habitat. This type of interaction between two or more populations is referred to as interspecific competition. In some cases, one species may eventually out-compete the other or others, and the “losing” species disappears from the area (see Figure 14.22 on the next page). A result like this often indicates that the interacting species had very similar ecological niches. Ecologists explain the effects of interspecific competition by referring to the competitive exclusion principle. Essentially, this theory states that species with niches that are exactly the same cannot co-exist; if two species have completely overlapping niches, one will always exclude the other, as shown by the dashed line in Figure 14.22B. However, if the niches of competing species are sufficiently different, they can both live in a particular area, although the density of one or more of the populations may be lowered by the presence of the other (as shown by the dashed line in Figure 14.22A). Figure 14.20 For gannets, which nest on rocky islands such as this one off Cape St. Mary’s in Newfoundland, the limited number of nesting sites determines the carrying capacity of the environment. As a result, only a certain number of pairs can nest and reproduce at any given time. When N is low, all birds can find a suitable nest site and reproduce, so the population grows. Above a certain N, however, many pairs fail to breed successfully and, as a result, population growth slows and eventually levels off. BIO FACT For some species, internal rather than external cues (such as a reduced food supply) seem to limit population size. Small rodents kept in experimental conditions with abundant food and shelter will reproduce quickly until their population reaches a certain density. At this point, even though the resources they need most are still unlimited, reproduction decreases. High density seems to produce a stress response in which hormonal changes delay sexual maturation, cause shrinkage of reproductive organs, reduce the effectiveness of the immune system, and often produce aggressive behaviour (sometimes including cannibalism). Exactly what cues trigger these hormonal changes is unclear, but similar effects of crowding have been noticed in some animal populations in nature. Figure 14.21 In Ontario, garlic mustard (Alliaria petiolata, a plant introduced from Europe) invades moist woodland habitats and crowds out many native species. It is a particular threat to some endangered plant species, such as the wood poppy (Stylophorum diphyllum). Chapter 14 Population Ecology • MHR 483 Number of paramecia (per mL) A 800 grown alone 400 mixed culture 0 Time Paramecium caudatum B Number of paramecia (per mL) driving force of evolutionary change. In each competing species, the individuals that are most different from their competitors will be best able to avoid competitive interactions and will therefore obtain the most resources. For example, if two species of birds compete for seeds of roughly equal sizes, those individuals of both species that can eat larger or smaller seeds will be able to find more food. They will, therefore, be more likely to survive and reproduce than will members of their own species that cannot avoid interspecific competition. As a result, their alleles — coding for the characteristics (such as beaks that can handle different seeds) that distinguish them from their competitors — will increase in frequency in subsequent generations. In this way, natural selection can produce increased divergence between competing species. In Figure 14.23, natural selection may take two different routes to lead to reduced competition. The total range of prey sizes taken by the two species may increase, with one species extending its preferences to smaller items than were taken previously, while the other may include larger foods than were eaten before. Alternatively, the total range of prey eaten may remain the same, but the niche of each species may shrink; one species may become a specialist on small prey while the other takes mainly large prey. Over time, this can increase the diversity of species living in a community. Paramecium aurelia 200 grown alone 100 mixed culture 0 Time Figure 14.22 When two species of paramecia (Paramecium aurelia and P. caudatum) are grown in separate cultures in a laboratory, their populations follow a logistic curve to reach their own carrying capacities. This is shown by the solid lines. However, when they are grown together, P. caudatum goes extinct in the mixed culture, shown by the dashed line in (B), since P. aurelia is a much better competitor. What this means is that differences between species can be increased as a result of natural selection, with interspecific competition as the species B Number of individuals species A Prey size Number of individuals or Prey size Figure 14.23 Within a species, individuals vary in many of their features. As a result, they use a range of whatever resources are necessary for their survival and reproduction. In this example, the members of two species, A and B, vary 484 MHR • Unit 5 Population Dynamics Prey size with respect to the size of their prey, but some individuals of both species eat intermediate-sized prey. This overlap produces interspecific competition, which may lead to evolutionary changes in both species. Interaction of Predator and Prey Populations Some populations, especially those of certain insects, birds, and mammals, fluctuate regularly in density. These alternating periods of high and low populations are often referred to as population cycles. While some small herbivorous mammals, such as voles, have 3- to 5-year cycles, larger herbivorous mammals, such as snowshoe hares (Lepus americanus) and muskrats (Ondatra zibethica), have 9- to 11-year cycles. These longer cycles are also typical of some birds, including ruffed grouse (Bonasa umbellus). The causes of cycles vary with the species and from population to population within a single species. Some may be due to a lag in response to density-dependent factors, as discussed in section 14.2. If such a lag is fairly constant, a more or less regular cycle of fluctuation above and below the population’s carrying capacity could result. Some of the mammal species that display fluctuations in population density are predators that have cycles overlapping those of their prey (such as some of the herbivores already described). One explanation for these cycles is the densitydependent effect of each population on the other. For example, some populations of Canada lynx prey almost exclusively on snowshoe hare (see Figure 14.24). An increase in the hare population would reduce competition for food among the lynx (Lynx canadensis), allowing them to increase their reproduction rate and survive longer. The result would be an increase in the population density of lynx. However, the presence of a large number of these predators would eventually cause the hare population to decrease. This, in turn, would increase competition among lynx for food, causing a decline in the predator population and permitting the prey population to expand once again. BIO FACT Some of the most remarkable population cycles in the world are those displayed by various species of cicadas (insects in the order Homoptera). The life cycle of these species takes place mostly underground, and requires 13 to 17 years to complete. At the end of this period, they emerge as adults in extremely high densities (as high as 600 per m2 ). When the adults lay their eggs and die, the aboveground population shrinks to virtually nothing. The long life cycle may be an adaptation to reduce predation. Since these species are around for such a short time, few predators have learned how to prey on them efficiently. However, there is more to this story than just the relationship between predator and prey. Hare populations on arctic islands where there are no lynx also undergo a cycle, indicating that it is not simply the effect of predators that causes hare populations to increase or decrease. An alternative hypothesis for fluctuations in the size of snowshoe hare populations is that the grazing activity of a large number of these herbivores causes serious damage to the plants (especially willows) that they eat. When the hare population is small, only a small portion of each plant is consumed. The plants can maintain high survival and reproduction rates in this situation, resulting in high plant density. This, in turn, allows the hare population to increase, perhaps to a point where their grazing damages the plants, thereby lowering plant survival and reproduction. The hare population will then decline because of a decrease in their food supply. A long-term research Continued on page 488 Number of pelts (in thousands) 150 ➥ lynx hare 100 50 0 1845 1855 1865 1875 1885 1895 1905 1915 1925 1935 Figure 14.24 This graph shows the number of Canada lynx and snowshoe hare pelts traded annually to the Hudson’s Bay Company over a 100-year period in Canada’s arctic (data which allowed biologists to estimate the population sizes for both species). Note that approximately every 10 years, populations of both species become very large, decline, and then become large again. These cycles seem much too regular to be explained by randomly occurring abiotic factors. Chapter 14 Population Ecology • MHR 485 Investigation SKILL FOCUS 1 4 • A Initiating and planning Paramecium Populations Predicting Cultures of small organisms, such as microscopic paramecia, can be used to study how various abiotic and biotic factors affect population growth patterns. These unicellular protozoa are ideal for studying changes in population size because they have a relatively short reproductive cycle and cultures can be easily maintained, measured, and manipulated in a laboratory environment. This investigation focusses on how density-dependent factors and density-independent factors affect the growth rate of paramecium populations. These observations will allow you to suggest how abnormal pH levels in natural ecosystems, resulting from acid precipitation or other types of contamination, might have negative effects on organisms living in aquatic environments. Pre-lab Questions What is the natural food supply of micro-organisms such as paramecium? Describe the life cycle of paramecium. How would lower pH levels affect populations of organisms that inhabit aquatic ecosystems? How would changes in food availability affect populations of organisms that inhabit aquatic ecosystems? Problem How can you determine the impact of density-dependent factors (such as food supply) and density-independent factors (such as pH) on the growth rate of paramecium populations? Prediction Predict the effect of changes in pH level and food supply on the growth rate of paramecium populations. CAUTION: Handle hydrochloric acid with care. Wash your hands after each observation. Materials glass jars of uniform size culture medium (skim milk powder) distilled water paramecium cultures droppers heating sources thermometers dilute hydrochloric acid pH hydrion papers (or other indicators that provide accurate pH readings) microscopes methyl cellulose Procedure 1. For this investigation, your class should be split into several groups. One half of the class should focus on testing the effects of a density-dependent limiting factor (food supply), while the other should investigate the effects of a density-independent 486 MHR • Unit 5 Population Dynamics Identifying variables Performing and recording Analyzing and interpreting Conducting research limiting factor (pH). Each student group will run one control and one or more experimental set-ups. 2. Identify the specific factor or variable (such as pH or food supply) to be investigated. This will be your independent (manipulated) variable. Select the range of factors to be tested. (For example, one group could maintain one control culture at neutral pH and an experimental culture at a slightly lower pH while the other groups maintain cultures at different pH levels. That way, a broad range of pH levels can be analyzed.) 3. Maintain all cultures at constant temperature and at uniform, medium light conditions. Avoid direct sunlight, drafts, and contamination of cultures. Ensure that all glassware is clean and uncontaminated by soap or other chemical residue. Leave each culture open to the air, but ensure that water levels remain constant by adding roomtemperature distilled water as required. 4. To start, add about 5 g of skim milk powder to 250 mL of distilled water. Inoculate each set-up with an identical volume of paramecium culture (about three or four full pipettes or droppers). The control set-up for each group should be kept at neutral pH and be provided with the same food supply and starter paramecium culture. In fact, you may wish to maintain each set-up for three to four days with a standard food supply and neutral pH, to stabilize each paramecium population before you begin to manipulate the environments. 5. If you are investigating food supply as a variable, add different amounts of food to your experimental set-ups (the class should work with a range of 0–20 g of food per experimental culture). 6. If you are investigating pH as a variable, add varying quantities of dilute HCl (start with 0.1 mol/L HCl) to produce a range of pH, from neutral to about pH 3. Maintain a constant pH in each set-up throughout the experiment. 7. Use a microscope to view samples of your cultures each day. Estimate the paramecium population of each culture. Try the following procedure to view and count the paramecia in your samples: (a) Squeeze all the air out of the dropper bulb. Use the dropper to remove a few drops of your paramecium culture. (b) Place a drop of methylcellulose and one or two drops of water on the centre of a glass microscope slide. (You can also add a few grains of very fine, washed sand to slow down the paramecia and prevent the cover slip from crushing the specimens in your sample.) (c) Carefully squeeze the bulb to deposit your culture sample into this mixture. (d) Gently lower a cover slip over the sample. (e) View your sample with the low-power objective lens of your microscope. Scan the sample and count the number of paramecia in five different fields of view (view the centre and the area near each edge of the cover slip). Add up the total number of individuals counted in all of the samples and divide by five to determine an average estimate of the current population size. Enter your data in a summary data table. (f) Use the same procedure to observe samples daily from each set-up and record your estimates of population size in the appropriate data table. 8. Ensure that your sampling technique can provide an accurate estimate of the population size without unduly disturbing the integrity of the ecosystem (and, if possible, without causing any changes to the appearance of individual paramecia in each culture). 9. After the data are entered in your summary table, graph the results. If available, you may wish to use a computer spreadsheet program to record your experimental data and generate graphs. Combine data from other student groups to generate graphs depicting the class results for each experimental variable tested. Post-lab Questions 1. Describe any observed changes in the paramecium population size. Graph the control and the experimental population sizes over the duration of your experimental procedure. 3. How did your results relate to your original prediction? 4. Discuss possible sources of error in your procedure (relating to such factors as the sampling procedures used and the ability to control any extraneous variables) that may influence your results. 5. Were your results consistent with the observations of other groups? Conclude and Apply 6. What can you conclude about the accuracy of your original prediction? 7. How could your procedure be improved? Exploring Further 8. Devise a supervised investigation to study populations of micro-organisms in actual aquatic ecosystems, such as ponds or streams. Have your teacher approve your experimental design and safety precautions in advance. Test the pH of each sample. Identify other significant density-dependent and density-independent regulating factors that may have an impact on certain species of microscopic organisms in local aquatic ecosystems (such as rivers or ponds). Indicate your sample locations on a local map. 9. Conduct research to assess population changes through the seasons in the ecosystems you have chosen. 10. What do you think is the mechanism by which changes in pH increase or decrease the rate of population growth? That is, why does the pH of its environment matter to a paramecium, or to any other organism? How would an effect on an individual organism translate into changes in population growth? 11. Which general type of regulating factor — densitydependent or density-independent — is most important in regulating the size of paramecium populations in nature? Or do you think both might be important, but under different circumstances? Explain your answer in full. 2. How do the size and growth rate of the control culture compare with the populations of your experimental paramecium cultures? Specifically, what do your results and the class results suggest about the impact of a density-dependent factor and a density-independent factor on the rate of growth of paramecium populations? Chapter 14 Population Ecology • MHR 487 project in the Yukon, led by Charles Krebs of the University of British Columbia, suggests that both predation and fluctuations in food supply contribute to the cycling of hare populations. Symbiotic Relationships In some cases, the interactions that occur between members of two different species are so close that they are said to have a symbiotic relationship. One member of the pair in such an association is referred to as the host and the other as the symbiont. The host is typically larger and more independent (that is, it can live on its own) than the symbiont (which often depends on the host and the symbiotic relationship for its survival). There are three general types of symbiotic relationships. In parasitism, the symbiont is referred to as a parasite; it benefits from the relationship, whereas the host is harmed. In terms of population growth, this relationship is much like that between a predator and its prey. An increase in host population density makes it possible for the parasites to increase in number. The increase in the number of parasites decreases host parasite 800 Table 14.1 Interspecific interactions 400 Population density 10 800 20 40 30 host parasite 400 50 800 host survival or reproductive ability, and may subsequently lead to a decrease in host density. With fewer hosts available, parasite density then declines. The result may be cyclical fluctuations of host and parasite population densities, as shown in Figure 14.25. A second type of symbiotic relationship is mutualism, in which both partners benefit from the relationship and growth in one population typically causes an increase in population density of the associated population. This is shown in Table 14.1, which summarizes the effects of different symbiotic relationships on population growth. In this table, a plus sign (+) indicates an increase in population density and a minus sign (−) indicates a decrease in density. In mutualism, for example, both species benefit from the relationship; positive growth in one population causes positive growth in the other. Mutualism is common in nature (as shown in Figure 14.26). For example, termites and various mammalian herbivores (such as cattle and sheep) depend on certain types of bacteria living in their digestive systems to help them digest the plant material they eat. The bacteria also benefit, since they are able to live in a protected, nutrient-rich environment. 60 70 80 Nature of relationship between populations Effect of growth in one population on the other population competitive – / – (both are negatively affected) predator/prey (includes herbivore/plant) + / – (one population gains at the expense of the other) host/parasite – / + (one population gains at the expense of the other) mutualistic + / + (both are positively affected) host parasite 400 90 100 110 Number of generations Figure 14.25 In a laboratory, populations of an Adzuki bean weevil (Callosobruchus chinensis) and its wasp parasitoid (Heterospilus prosopidis) cycled for 112 generations or six years (until the experiment ended). In this case the wasp is called a parasitoid because it feeds on living tissue (like a parasite), but kills the larvae of its host, in the process making it like a predator in some ways. 488 MHR • Unit 5 Population Dynamics Figure 14.26 Examples of mutualism include the relationships that exist between many types of plant and animal species. In this case, the animal gets food (pollen or nectar) while the plant gets help moving its pollen from plant to plant for reproduction. The third type of symbiotic relationship is commensalism, in which one partner benefits and the other is unaffected. There are few (if any) examples of true commensalism in nature, since it is unlikely that one participant in any type of ecological interaction will be unaffected. Barnacles that “hitchhike” by attaching themselves to whales are sometimes considered to be the partner that benefits in a commensal relationship, while the whale is thought to be unaffected. However, because barnacles may slightly decrease a whale’s ability to find food or escape from predators, they may actually lower the whale’s reproductive success or chance of survival (and thus reduce the density of whale populations). Other often-cited examples of commensalism are the relationships between cowbirds or cattle egrets and the cattle they associate with (see Figure 14.27). ELECTRONIC LEARNING PARTNER Use the Electronic Learning Partner to enhance your understanding of symbiotic relationships, including parasitism, mutualism, and commensalism. As you might predict, populations that are regulated by density-dependent factors tend to be more predictable than those regulated by weather, fire, and other abiotic mechanisms. Ecologists and wildlife managers make use of this predictability in various ways, as demonstrated in the Thinking Lab on page 490. BIO FACT Forestry and farming depend on a mutualistic relationship between fungi and plant roots. A fungus’s body consists of many thread-like extensions, which maximize the amount of surface area exposed to its food source. This is important since a fungus absorbs the nutrients it needs over its body surface. When the threads of certain fungal species become intertwined with a plant’s roots (these associations of roots and fungal threads are called mycorrhizae), they help the plant absorb far more water and nutrients than it could on its own. The fungus also benefits, getting a small portion of the photosynthetic products the plant makes with these materials. One of the problems arising from clearcutting is that removal of all the trees from an area often causes the death of the soil fungi. This removal of soil fungi, in turn, may reduce the ability of newly planted seedlings to survive and grow. Population Regulation: Many Factors at Work One way to think about the many factors involved in the growth and regulation of a population’s size is shown in Figure 14.28. The line labelled (a) shows the biotic potential of a population — how it would grow if it could realize its full potential for growth. The jagged line labelled (b) shows the regulating effects that the environment has on the population’s growth. In this case, abiotic factors produce the sudden dramatic declines in the curve, while biotic factors produce less dramatic declines and hold the population roughly to the environment’s carrying capacity. Since both the (b) environmental resistance Population size (a) biotic potential Figure 14.27 Is this relationship commensal? The birds benefit because they feed on insects flushed out of the grass by the cows while they graze. Although the cattle are mostly unaffected by the presence of the birds, they may obtain some benefit since the birds sometimes eat ticks and other parasites they find on the cows’ skin. Time Figure 14.28 The relationship between a population’s biotic potential and the environmental resistance to its growth Chapter 14 Population Ecology • MHR 489 abiotic and biotic factors are part of the population’s environment, the combination of their effects is sometimes referred to as environmental resistance to population growth. The examples discussed under “Symbiotic Relationships” might suggest that populations of some kinds of organisms (such as insects) are regulated by abiotic, density-independent factors, whereas populations of other kinds of organisms (including birds and mammals) are regulated by biotic, density-dependent mechanisms. This apparent contrast led early ecologists to argue about which form of population regulation was THINKING LAB Sustainable Harvesting You Try It Background Theoretically, populations regulated by density-dependent factors can be harvested. In other words, individuals can be removed for commercial and/or recreational purposes in such a way that the population is not depleted. This will be the case if the number of individuals being harvested (referred to as the yield of the population) is just equal to the number being added as a result of reproduction. One of the tasks of those who manage populations of organisms that humans use as resources (for example, forests; fish stocks; and deer, moose, and bear populations) is to try to determine the Maximum Sustainable Yield (MSY). MSY is the largest number of individuals that can be removed from the population each year (or other harvest period) without reducing the population’s growth over the long term. In this lab, you will see how the MSY can be assessed — in theory. 500 N0 = 5, K = 500, r = 0.2 Population size (N) 400 200 100 10 20 30 40 50 Time Growth curve for a theoretical, density-dependent population 490 1. The graph shows the growth curve for a theoretical, density-dependent population living in an environment in which the carrying capacity is 500 (K = 500). The population started out with five individuals (N0 = 5) and grew at a rate of 0.2 offspring per individual per time period (r = 0.2). Using the curve, estimate the number of individuals added to the population in the first five time units. Estimate the size of the population in the middle of this time interval (that is, at time = 2.5 units). Follow the same procedure for intervals 5–10, 10–15, 15–20, and so on up to 45–50. Record your data in a table, or use a spreadsheet program if available. 2. Plot the data you obtained in Step 1 for each of the 10 time intervals on a graph of “Number of individuals added to the population” (y-axis) versus “Population size” (x-axis). How big is the population during the interval when the maximum number of individuals is being added? Is N at this time near K, far below it, or halfway to K? Explain the shape of the curve you have plotted. Why does the curve increase to a maximum and then decrease? 3. Suppose you are the manager of this population and you are trying to determine its MSY. At what size would you want to maintain the population, and why? How would an increase in the population’s size above the MSY point affect yield? How would a decrease affect yield? 300 0 more important, or at least more common. These discussions, along with more research, eventually led to the understanding that for most populations, both types of regulating factors play a role. For example, most of the time a population of birds may be regulated by density-dependent factors. However, occasional catastrophes may reduce the population to a point where density has little influence on the growth rate. Similarly, severe weather may regularly cause the size of an insect population to decline. If the environment contains a limited number of protective sites where members of this population can hide to avoid the weather, MHR • Unit 5 Population Dynamics 4. Although MSY values have often been used to set harvest quotas (such as the number of fish that can be caught or deer that can be hunted), the system has not worked well in some situations. Many populations have been seriously depleted as a result of overfishing or excessive hunting, in part because quotas were set too high. Why do you think this might have happened? What problems can you see occurring as a result of using this method to set quotas? density does play a role. The amount by which N exceeds the number of hiding places will affect the proportion of the population killed. In fact, most populations are probably regulated by a combination of density-dependent and densityindependent factors. SECTION The abiotic and biotic factors that regulate population size also influence other characteristics of populations. These characteristics will be described in the last section of this chapter. REVIEW affected? In what way might your answer to this last question change your interpretation of the type of relationship exhibited here? 1. MC What factors might eventually limit the exponential growth rates of a particular insect species found in such regions as the Canadian prairies? 2. K/U What might allow some animal populations to grow beyond the theoretical carrying capacity of their environment and then crash to abnormally low levels? 10. K/U What is the difference between the way in which abiotic and biotic factors typically produce environmental resistance to population growth? 3. K/U What factors might limit the population size of animals (such as vultures) that live as scavengers? 11. 4. K/U In a density-dependent population, why does population growth decline as population density increases? 5. MC Provide a real-life example of a situation in which interspecific competition limits the population size and growth of a particular species. 6. K/U What combination of factors might produce regular population cycles typical of small herbivore species such as mice and squirrels? MC Roosevelt elk (Cervus elaphus roosevelti) were re-introduced to the Sunshine Coast region of British Columbia because overhunting eliminated the original elk population many years ago. Their population has now grown to about 250, and some local residents are complaining that the elk are devouring plants in gardens, parks, and golf courses. Limited hunting of this population is now permitted in an attempt to control the problem. Do you think hunting of this elk population should be permitted? How do you think the resumption of hunting might affect the long-term growth patterns of this population and other species that share the same environment as the elk? 7. I Rat populations in many urban areas in Canada have increased dramatically in recent years. Use the Internet to research public health records and other web sites for data about the extent of this problem. Describe the factors that seem to be contributing to the growth in local rat populations and the factors that may ultimately limit their population size. 12. MC In ecological terms, what measures would be most effective in controlling populations of pest species such as rats, mice, or certain types of insects? 13. I Having been recently appointed regional wildlife manager, you must set a quota on the number of moose that can be hunted during the next hunting season. What information do you need? Design a study that will allow you to obtain the necessary data. 14. C You have just been hired to teach Grade 6 at a local school. Design a lesson plan, including an activity, that you might use to teach your students about the factors that affect population growth in different environments. 15. C Although all populations eventually face environmental resistance to continued growth, the contribution of abiotic and biotic factors to this resistance may vary from species to species. Compare a micro-organism, such as E. coli, a plant (such as a type of tree), and a mammal, such as the snowshoe hare or the black bear, with respect to the type of factors that typically limit the growth of populations of each species. 8. 9. MC Biologists have observed increasing instances of coral bleaching in recent years. This phenomenon involves the death of small green algae that live in the tissues of coral polyps (the tiny animals whose bodies secrete much of the material that forms coral reefs). Following the death of the algae, the coral organisms that hosted them also die. Describe the probable type of relationship that exists between the algae and coral organisms. K/U Some species of sharks are often seen with small fish called remoras attached to their bellies. The remoras seem to feed off the scraps of food dropped by the sharks when they are feeding on prey. The sharks seem to derive no benefit from the presence of the remoras. What type of symbiotic relationship might the association of the shark and remora illustrate? Do you think it likely that the shark is totally unaffected by the remora? In what way might it be Chapter 14 Population Ecology • MHR 491