Maximum sustainable yield
Encyclopedia
In population ecology
Population ecology
Population ecology is a sub-field of ecology that deals with the dynamics of species populations and how these populations interact with the environment. It is the study of how the population sizes of species living together in groups change over time and space....

 and economics
Economics
Economics is the social science that analyzes the production, distribution, and consumption of goods and services. The term economics comes from the Ancient Greek from + , hence "rules of the house"...

, maximum sustainable yield or MSY is, theoretically, the largest yield (or catch) that can be taken from a species' stock over an indefinite period. Fundamental to the notion of sustainable harvest, the concept of MSY aims to maintain the population size at the point of maximum growth rate by harvesting the individuals that would normally be added to the population, allowing the population to continue to be productive indefinitely. Under the assumption of logistic growth
Logistic function
A logistic function or logistic curve is a common sigmoid curve, given its name in 1844 or 1845 by Pierre François Verhulst who studied it in relation to population growth. It can model the "S-shaped" curve of growth of some population P...

, resource limitation does not constrain individuals’ reproductive rates when populations are small, but because there are few individuals, the overall yield is small. At intermediate population densities, also represented by half the carrying capacity
Carrying capacity
The carrying capacity of a biological species in an environment is the maximum population size of the species that the environment can sustain indefinitely, given the food, habitat, water and other necessities available in the environment...

, individuals are able to breed to their maximum rate. At this point, called the maximum sustainable yield, there is a surplus of individuals that can be harvested because growth of the population is at its maximum point due to the large number reproducing individuals. Above this point, density dependent factors increasingly limit breeding until the population reaches carrying capacity. At this point, there are no surplus individuals to be harvested and yield drops to zero. The maximum sustainable yield is usually higher than the optimum sustainable yield
Optimum sustainable yield
In population ecology and economics, optimum sustainable yield is the level of effort that maximizes the difference between total revenue and total cost. Or, where marginal revenue equals marginal cost. This level of effort maximizes the economic profit, or rent, of the resource being utilized...

 and maximum economic yield.

MSY is extensively used for fisheries management
Fisheries management
Fisheries management draws on fisheries science in order to find ways to protect fishery resources so sustainable exploitation is possible. Modern fisheries management is often referred to as a governmental system of appropriate management rules based on defined objectives and a mix of management...

. Unlike the logistic (Schaefer) model, MSY has been refined in most modern fisheries models and occurs at around 30% of the unexploited population size. This fraction differs among populations depending on the life history of the species and the age-specific selectivity of the fishing method.

However, the approach has been widely criticized as ignoring several key factors involved in fisheries management and has led to the devastating collapse of many fisheries. As a simple calculation, it ignores the size and age of the animal being taken, its reproductive status, and it focuses solely on the species in question, ignoring the damage to the ecosystem caused by the designated level of exploitation and the issue of bycatch. Among conservation biologists
Conservation biology
Conservation biology is the scientific study of the nature and status of Earth's biodiversity with the aim of protecting species, their habitats, and ecosystems from excessive rates of extinction...

 it is widely regarded as dangerous and misused.

History

The concept of MSY as a fisheries management strategy developed in the early 1930s. It increased in popularity in the 1950s with the advent of surplus-production models with explicitly estimate MSY. As an apparently simple and logical management goal, combined with the lack of other simple management goals of the time, MSY was adopted as the primary management goal by several international organizations (e.g., IWC
International Whaling Commission
The International Whaling Commission is an international body set up by the terms of the International Convention for the Regulation of Whaling , which was signed in Washington, D.C...

, IATTC
Inter-American Tropical Tuna Commission
The Inter-American Tropical Tuna Commission is an international commission that is responsible for the conservation and management of tuna and other marine resources in the eastern Pacific Ocean.-History:The Inter-American Tropical Tuna Commission was created by the Convention for the...

, ICCAT
International Commission for the Conservation of Atlantic Tunas
The International Commission for the Conservation of Atlantic Tunas is an intergovernmental organisation responsible for the management and conservation of tunas and tuna-like species in the Atlantic Ocean and adjacent seas. The organization was established in 1969, at a conference in Rio de...

, ICNAF
Northwest Atlantic Fisheries Organization
The is an intergovernmental organization with a mandate to provide scientific advice and management of fisheries in the northwestern part of the Atlantic Ocean...

), and individual countries.

Between 1949 and 1955, the U.S. maneuvered to have MSY declared the goal of international fisheries management (Johnson 2007). The international MSY treaty that was eventually adopted in 1955 gave foreign fleets the right to fish off any coast. Nations that wanted to exclude foreign boats had to first prove that its fish were overfished.

As experience was gained with the model, it became apparent to some researchers that it lacked the capability to deal with the real world operational complexities and the influence of trophic
Trophic level
The trophic level of an organism is the position it occupies in a food chain. The word trophic derives from the Greek τροφή referring to food or feeding. A food chain represents a succession of organisms that eat another organism and are, in turn, eaten themselves. The number of steps an organism...

 and other interactions. In 1977, Larkin wrote its epitaph, challenging the goal of maximum sustained yield on several grounds: It put populations at too much risk; it did not account for spatial variability in productivity; it did not account for species other than the focus of the fishery; it considered only the benefits, not the costs, of fishing; and it was sensitive to political pressure. In fact, none of these criticisms was aimed at sustainability as a goal. The first one noted that seeking the absolute MSY with uncertain parameters was risky. The rest point out that the goal of MSY was not holistic; it left out too many relevant features.

Some managers began to use more conservative quota recommendations, but the influence of the MSY model for fisheries management
Fisheries management
Fisheries management draws on fisheries science in order to find ways to protect fishery resources so sustainable exploitation is possible. Modern fisheries management is often referred to as a governmental system of appropriate management rules based on defined objectives and a mix of management...

 still prevailed. Even while the scientific community was beginning to question the appropriateness and effectiveness of MSY as a management goal, it was incorporated into the 1982 United Nations Convention for the Law of the Sea
United Nations Convention on the Law of the Sea
The United Nations Convention on the Law of the Sea , also called the Law of the Sea Convention or the Law of the Sea treaty, is the international agreement that resulted from the third United Nations Conference on the Law of the Sea , which took place from 1973 through 1982...

, thus ensuring its integration into national and international fisheries acts and laws. According to Walters and Maguire, an ‘‘institutional juggernaut had been set in motion’’, climaxing in the early 1990s with the collapse of northern cod.

Population growth

The key assumption behind all sustainable harvesting models such as MSY is that populations of organisms grow and replace themselves – that is, they are renewable resources. Additionally it is assumed that because the growth rates, survival rates, and reproductive rates increase when harvesting reduces population density
Population density
Population density is a measurement of population per unit area or unit volume. It is frequently applied to living organisms, and particularly to humans...

, they produce a surplus of biomass that can be harvested. Otherwise, sustainable harvest would not be possible.

Another assumption of renewable resource harvesting is that populations of organisms do not continue to grow indefinitely; they reach an equilibrium population size, which occurs when the number of individuals matches the resources available to the population (i.e., assume classic logistic growth
Logistic function
A logistic function or logistic curve is a common sigmoid curve, given its name in 1844 or 1845 by Pierre François Verhulst who studied it in relation to population growth. It can model the "S-shaped" curve of growth of some population P...

). At this equilibrium population size, called the carrying capacity
Carrying capacity
The carrying capacity of a biological species in an environment is the maximum population size of the species that the environment can sustain indefinitely, given the food, habitat, water and other necessities available in the environment...

, the population remains at a stable size.

The logistic model (or logistic function
Logistic function
A logistic function or logistic curve is a common sigmoid curve, given its name in 1844 or 1845 by Pierre François Verhulst who studied it in relation to population growth. It can model the "S-shaped" curve of growth of some population P...

) is a function that is used to describe bounded population growth
Population growth
Population growth is the change in a population over time, and can be quantified as the change in the number of individuals of any species in a population using "per unit time" for measurement....

 under the previous two assumptions. The logistic function
Logistic function
A logistic function or logistic curve is a common sigmoid curve, given its name in 1844 or 1845 by Pierre François Verhulst who studied it in relation to population growth. It can model the "S-shaped" curve of growth of some population P...

 is bounded at both extremes: when there are not individuals to reproduce, and when there is an equilibrium number of individuals (i.e., at carrying capacity
Carrying capacity
The carrying capacity of a biological species in an environment is the maximum population size of the species that the environment can sustain indefinitely, given the food, habitat, water and other necessities available in the environment...

). Under the logistic model, population growth rate between these two limits is most often assumed to be sigmoidal
Sigmoid function
Many natural processes, including those of complex system learning curves, exhibit a progression from small beginnings that accelerates and approaches a climax over time. When a detailed description is lacking, a sigmoid function is often used. A sigmoid curve is produced by a mathematical...

 (Figure 1). There is scientific evidence that some populations do grow in a logistic fashion towards a stable equilibrium – a commonly cited example is the logistic growth of yeast
Yeast
Yeasts are eukaryotic micro-organisms classified in the kingdom Fungi, with 1,500 species currently described estimated to be only 1% of all fungal species. Most reproduce asexually by mitosis, and many do so by an asymmetric division process called budding...

.

The equation describing logistic growth is:
(equation 1.1)


The parameter values are:
=The population size at time t

=The carrying capacity of the population

= The population size at time zero

= the intrinsic rate of population increase (the rate at which the population grows when it is very small)


From the logistic function, the population size at any point can be calculated as long as , , and are known.

Differentiating equation 1.1 give an expression for how the rate of population increases as t increases. At first, the population growth rate is fast, but it begins to slow as times goes on until it levels off to the maximum growth rate, after which it begins to decrease (figure 2).

The equation for figure 2 is the differential of equation 1.1 (Verhulst's
Pierre François Verhulst
Pierre François Verhulst was a mathematician and a doctor in number theory from the University of Ghent in 1825...

 1838 growth model):
(equation 1.2)


can be understood as the change in population (N) with respect to a change in time (t). Equation 1.2 is the usual way in which logistic growth is represented mathematically and has several important features. First, at very low population sizes, the value of is small, so the population growth rate is approximately equal to , meaning the population is growing exponentially at a rate r (the intrinsic rate of population increase). Despite this, the population growth rate is very low (low values on the y-axis of figure 2) because, even though each individual is reproducing at a high rate, there are few reproducing individuals present. Conversely, when the population is large the value of approaches 1 effectively reducing the terms inside the brackets of equation 1.2 to zero. The effect is that the population growth rate is again very low, because either each individual is hardly reproducing or mortality rates are high. As a result of these two extremes, the population growth rate is maximum at an intermediate population or half the carrying capacity ().

MSY model

The simplest way to model harvesting is to modify the logistic equation so that a certain number of individuals is continuously removed:
(equation 1.3)


Where H represents the number of individuals being removed from the population – that is, the harvesting rate. When H is constant, the population will be at equilibrium when the number of individuals being removed is equal to the population growth rate (figure 3). The equilibrium population size under a particular harvesting regime can be found when the population is not growing – that is, when . This occurs when the population growth rate is the same as the harvest rate:


Figure 3 shows how growth rate varies with population density. For low densities (far from carrying capacity), there is little addition (or “recruitment”) to the population, simply because there are few organisms to give birth. At high densities, though, there is intense competition for resources, and growth rate is again low because the death rate is high. In between these two extremes, the population growth rate rises to a maximum value (). This maximum point represents the maximum number of individuals that can be added to a population by natural processes. If more individuals than this are removed from the population, the population is at risk for decline to extinction. The maximum number that can be harvested in a sustainable manner, called the maximum sustainable yield, is given by this maximum point.

Figure 3 also shows several possible values for the harvesting rate, H. At , there are two possible population equilibrium points: a low population size () and a high one (). At , a slightly higher harvest rate, however there is only one equilibrium point (at ), which is the population size that produces the maximum growth rate. With logistic growth, this point, called the maximum sustainable yield, is where the population size is half the carrying capacity (or ). The maximum sustainable yield is the largest yield that can be taken from a population at equilibrium.
In figure 3, if is higher than , the harvesting would exceed the population’s capacity to replace itself at any population size ( in figure 3). Because harvesting rate is higher than the population growth rate at all values of , this rate of harvesting is not sustainable.
An important feature of the MSY model is how harvested populations respond to environmental fluctuations or illegal offtake. Consider a population at harvested at a constant harvest level . If the population falls (due to a bad winter or illegal harvest) this will ease density-dependent population regulation and increase yield, moving the population back to , a stable equilibrium. In this case, a negative feedback loop creates stability. The lower equilibrium point for the constant harvest level is not stable however; a population crash or illegal harvesting will decrease population yield farther below the current harvest level, creating a positive feedback loop leading to extinction. Harvesting at is also potentially unstable. A small decrease in the population can lead to a positive feedback loop and extinction if the harvesting regime () is not reduced. Thus, some consider harvesting at MSY to be unsafe on ecological and economic grounds. The MSY model itself can be modified to harvest a certain percentage of the population or with constant effort constraints rather than an actual number, thereby avoiding some of its instabilities.

The MSY equilibrium point is semi-stable – a small increase in population size is compensated for, a small decrease to extinction if H is not decreased. Harvesting at MSY is therefore dangerous because it is on a knife-edge – any small population decline leads to a positive feedback, with the population declining rapidly to extinction if the number of harvested stays the same.

The formula for maximum sustained harvest () is one-fourth the maximum population or carrying capacity () times the intrinsic rate of growth ().


Implications of MSY model

Starting to harvest a previously unharvested population will always lead to a decrease in the population size. That is, it is impossible for a harvested population to remain at its original carrying capacity. Instead, the population will either stabilize at a new lower equilibrium size or, if the harvesting rate is too high, decline to zero.

The reason why populations can be sustainably harvested is that they exhibit a density-dependent response. This means that at any population size below K, the population is producing a surplus yield that is available for harvesting without reducing population size. Density dependence is the regulator process that allows the population to return to equilibrium after a perturbation. The logistic equation assumes that density dependence takes the form of negative feedback.

If a constant number of individuals is harvested from a population at a level greater than the MSY, the population will decline to extinction. Harvesting below the MSY level leads to a stable equilibrium population if the starting population is above the unstable equilibrium population size.

Uses of MSY

MSY has been especially influential in the management of renewable biological resources such as commercially important fish and wildlife. In fisheries terms, maximum sustainable yield (MSY) is the largest average catch that can be captured from a stock under existing environmental conditions. MSY aims at a balance between too much and too little harvest to keep the population at some intermediate abundance with a maximum replacement rate.

Relating to MSY, the maximum economic yield (MEY) is the level of catch that provides the maximum net economic benefits or profits to society. Like optimum sustainable yield
Optimum sustainable yield
In population ecology and economics, optimum sustainable yield is the level of effort that maximizes the difference between total revenue and total cost. Or, where marginal revenue equals marginal cost. This level of effort maximizes the economic profit, or rent, of the resource being utilized...

, MEY is usually less than MSY.

Limitations of MSY approach

Although it is widely practiced by state and federal government agencies regulating wildlife, forests, and fishing, MSY has come under heavy criticism by ecologists and others from both theoretical and practical reasons. The concept of maximum sustainable yield is not always easy to apply in practice. Estimation problems arise due to poor assumptions in some models and lack of reliability of the data. Biologists, for example, do not always have enough data to make a clear determination of the population’s size and growth rate. Calculating the point at which a population begins to slow from competition is also very difficult. The concept of MSY also tends to treat all individuals in the population as identical, thereby ignoring all aspects of population structure such as size or age classes and their differential rates of growth, survival, and reproduction.

As a management goal, the static interpretation of MSY (i.e., MSY as a fixed catch that can be taken year after year) is generally not appropriate because it ignores the fact that fish populations undergo natural fluctuations (i.e., MSY treats the environment as unvarying) in abundance and will usually ultimately become severely depleted under a constant-catch strategy. Thus, most fisheries scientists now interpret MSY in a more dynamic sense as the maximum average yield (MAY) obtained by applying a specific harvesting strategy to a fluctuating resource.

Orange roughy

An example of errors in estimating the population dynamics
Population dynamics
Population dynamics is the branch of life sciences that studies short-term and long-term changes in the size and age composition of populations, and the biological and environmental processes influencing those changes...

 of a species occurred with in the New Zealand Orange roughy
Orange roughy
The orange roughy, red roughy, or deep sea perch, Hoplostethus atlanticus, is a relatively large deep-sea fish belonging to the slimehead family . The Marine Conservation Society has categorized orange roughy as vulnerable to exploitation...

 fishery. Early quotas were based on an assumption that the orange roughy had a fairly short lifespan and bred relatively quickly. However, it was later discovered that the orange roughy
Orange roughy
The orange roughy, red roughy, or deep sea perch, Hoplostethus atlanticus, is a relatively large deep-sea fish belonging to the slimehead family . The Marine Conservation Society has categorized orange roughy as vulnerable to exploitation...

 lived a long time and had bred slowly (~30 years). By this stage stocks had been largely depleted.

Overfishing

All around the world, from the arctic to the tropics, there is a crisis in the world’s fisheries. Until fairly recently it was assumed that our marine resources were limitless.

In recent years however, an accelerating decline has been observed in the productivity of many important fisheries. Fisheries which have been devastated in recent times include (but are not limited too) the great whale fisheries, the Grand Bank fisheries of the western Atlantic, and the Peruvian anchovy fishery. Recent assessments by the United Nations Food and Agriculture Organization (FAO) of the state of the world’s fisheries indicate a levelling off of landings in the 1990s, at about 100 million tons.

In addition, the composition of global catches has changed. As fishers deplete larger, long-lived predatory fish species such as cod, tuna, shark, and snapper, they move down to the next level – to species that tend to be smaller, shorter-lived, and less valuable.

Overfishing is a classic example of the tragedy of the commons
Tragedy of the commons
The tragedy of the commons is a dilemma arising from the situation in which multiple individuals, acting independently and rationally consulting their own self-interest, will ultimately deplete a shared limited resource, even when it is clear that it is not in anyone's long-term interest for this...

.

Optimum sustainable yield

In population ecology
Population ecology
Population ecology is a sub-field of ecology that deals with the dynamics of species populations and how these populations interact with the environment. It is the study of how the population sizes of species living together in groups change over time and space....

 and economics
Economics
Economics is the social science that analyzes the production, distribution, and consumption of goods and services. The term economics comes from the Ancient Greek from + , hence "rules of the house"...

, optimum sustainable yield is the level of effort (LOE
Loe
Loe or LOE may refer to:People:*King Loe, semi-legendary king of Maui*Erlend Loe, Norwegian novelist*Kameron Loe, American baseball pitcherPlaces:*Loe, Estonia, a village in Estonia*The Loe, also known as Loe Pool, Cornwall, UK...

) that maximizes the difference between total revenue and total cost. Or, where marginal revenue equals marginal cost. This level of effort maximizes the economic profit, or rent, of the resource being utilized. It usually corresponds to an effort level lower than that of maximum sustainable yield.
In environmental science
Environmental science
Environmental science is an interdisciplinary academic field that integrates physical and biological sciences, to the study of the environment, and the solution of environmental problems...

, optimum sustainable yield is the largest economical yield of a renewable resource achievable over a long time period without decreasing the ability of the population or its environment to support the continuation of this level of yield.

See also

  • Maximum economic yield (MEY)
  • Ecological yield
    Ecological yield
    Ecological yield is the harvestable population growth of an ecosystem. It is most commonly measured in forestry;sustainable forestry is defined as that which does not harvest more wood in a year than has grown in that year, within a given patch of forest....

  • Population dynamics of fisheries
    Population dynamics of fisheries
    A fishery is an area with an associated fish or aquatic population which is harvested for its commercial or recreational value. Fisheries can be wild or farmed. Population dynamics describes the ways in which a given population grows and shrinks over time, as controlled by birth, death, and...

  • Population dynamics
    Population dynamics
    Population dynamics is the branch of life sciences that studies short-term and long-term changes in the size and age composition of populations, and the biological and environmental processes influencing those changes...

  • Fisheries management
    Fisheries management
    Fisheries management draws on fisheries science in order to find ways to protect fishery resources so sustainable exploitation is possible. Modern fisheries management is often referred to as a governmental system of appropriate management rules based on defined objectives and a mix of management...

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