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In [[population ecology]] and [[economics]], '''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 function|logistic growth]], 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]], 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 of 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]] and [[maximum economic yield]].

MSY is extensively used for [[fisheries management]]. Unlike the logistic (Schaefer) model,<ref name="Schaefer">Schaefer 1954</ref> 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 biology|conservation biologists]] it is widely regarded as dangerous and misused.<ref name=Epitaph>Larkin PA (1977) [http://docs.google.com/viewer?a=v&q=cache:0ZITmBnzlDUJ:fiesta.bren.ucsb.edu/~gsd/595e/docs/22.%2520Larkin_Epitaph_Max_Sust_Yield.pdf+Larkin+1977+%22An+epitaph+for+the+concept+of+maximum+sustained+yield%22&hl=en&gl=nz&sig=AHIEtbSBFh-w5xzWiBK7rnHSLBmolsfaWA "An epitaph for the concept of maximum sustained yield"] ''Transactions of the American Fisheries Society'', '''106''': 1–11.</ref><ref name=Lessons>[[Carl Walters|Walters C]] and Maguire J (1996) "Lessons for stock assessment from the northern cod collapse", ''Reviews in Fish Biology and Fisheries'', '''6''':125–137.</ref>

==History==
The concept of MSY as a fisheries management strategy developed in Belmare New Jersey in the early 1930s.<ref name="Russell">
Russell 1931</ref><ref>Hjort et al 1933</ref><ref>Graham 1935</ref>  It increased in popularity in the 1950s with the advent of surplus-production models with explicitly estimate MSY.<ref name="Schaefer"/>  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., [[International Whaling Commission|IWC]], [[Inter-American Tropical Tuna Commission|IATTC]],<ref>IATTC, Inter-American Tropical Tuna Commission</ref> [[International Commission for the Conservation of Atlantic Tunas|ICCAT]], [[Northwest Atlantic Fisheries Organization|ICNAF]]), and individual countries.<ref name="Mace">Mace 2001</ref>

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.<ref name="Botsford">Botsford et al 1997</ref>

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 level|trophic]] 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.<ref name="Larkin">Larkin 1977</ref> 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.<ref name="Botsford"/>

Some managers began to use more conservative quota recommendations, but the influence of the MSY model for [[fisheries management]] still prevailed. Even while the scientific community was beginning to question the appropriateness and effectiveness of MSY as a management goal,<ref name="Larkin"/><ref>Sissenwine 1978</ref> it was incorporated into the [[United Nations Convention on the Law of the Sea|1982 United Nations Convention for the Law of the Sea]], thus ensuring its integration into national and international fisheries acts and laws.<ref name="Mace"/> According to Walters and Maguire, an ‘‘institutional juggernaut had been set in motion’’, climaxing in the early 1990s with the collapse of northern cod.<ref>Walters and Maguire, 1996</ref>

== Modelling MSY ==
=== Population growth ===
{{see also|Population growth}}

The key assumption behind all [[sustainable harvest]]ing 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]],<ref name="Russell"/> 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 function|logistic growth]]).  At this equilibrium population size, called the [[carrying capacity]], the population remains at a stable size.<ref name="ReferenceA">Milner-Gulland and Mace 1998, pp. 14-17.</ref>

[[Image:logisticpopulationgrowth2.jpg|thumb|300px|right|Figure 1]]

The logistic model (or [[logistic function]]) is a function that is used to describe bounded [[population growth]] under the previous two assumptions.  The [[logistic function]] 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]]).  Under the logistic model, population growth rate between these two limits is most often assumed to be [[sigmoid function|sigmoidal]] (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]].

The equation describing logistic growth is:<ref name="ReferenceA"/>

:: <math> N_{t} = \frac{K}{1 + \frac{K-N_0}{N_0} e^{-rt}}</math> (equation 1.1)

The parameter values are:
:: <math> N_{t}</math>=The population size at time t

:: <math> K</math>=The carrying capacity of the population

:: <math> N_{0}</math>= The population size at time zero

:: <math> r</math>= 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 <math> r</math>, <math> K</math>, and <math> N_{0}</math> are known.

[[Image:growthratevs.populationsize.jpg|thumb|300px|right|Figure 2]]

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 ([[Pierre François Verhulst|Verhulst's]] 1838 [[Logistic function#In ecology: modeling population growth|growth model]]):<ref name="ReferenceA"/>

::<math> \frac{dN}{dt} = r N \left(1 - \frac {N}{K} \right)</math> (equation 1.2)

<math> \frac{dN}{dt}</math> 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 <math> \frac{N}{K}</math> is small, so the population growth rate is approximately equal to <math> rN</math>, 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 <math> \frac{N}{K}</math> 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.<ref name="ReferenceA"/>   As a result of these two extremes, the population growth rate is maximum at an intermediate population or half the carrying capacity (<math>N= \frac{K}{2}</math>).

=== MSY model ===

[[Image:harvestingrates.jpg|thumb|300px|right|Figure 3]]

The simplest way to model harvesting is to modify the logistic equation so that a certain number of individuals is continuously removed:<ref name="ReferenceA"/>

:: <math> \frac{dN}{dt} = rN\left(1-\frac{N}{K}\right) - H </math> (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 <math> \frac{dN}{dt} = 0</math>.  This occurs when the population growth rate is the same as the harvest rate:

:: <math> rN\left(1-\frac{N}{K}\right) = H </math>

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 (<math> N_{MSY}</math>).   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.<ref name="Jennings">Jennings et al 2001</ref>  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 <math>H_1</math>, there are two possible population equilibrium points: a low population size (<math>N_a</math>) and a high one (<math>N_b</math>).  At <math>H_2</math>, a slightly higher harvest rate, however there is only one equilibrium point (at <math> N_{MSY}</math>), 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 <math> N = \frac{K}{2}</math>). The maximum sustainable yield is the largest yield that can be taken from a population at equilibrium.
In figure 3, if <math>H</math> is higher than <math>H_2</math>, the harvesting would exceed the population’s capacity to replace itself at any population size (<math> H_3</math> in figure 3).  Because harvesting rate is higher than the population growth rate at all values of <math>N</math>, 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 <math>N_b</math> harvested at a constant harvest level <math>H_1</math>.  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 <math>N_b</math>, a stable equilibrium.  In this case, a negative feedback loop creates stability.  The lower equilibrium point for the constant harvest level <math>H_1</math> 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 <math>N_{MSY}</math> is also potentially unstable.  A small decrease in the population can lead to a positive feedback loop and extinction if the harvesting regime (<math>H_2</math>) is not reduced.  Thus, some consider harvesting at MSY to be unsafe on ecological and economic grounds.<ref name="Jennings"/><ref name="Milner-Gulland and Mace 1998">Milner-Gulland and Mace 1998.</ref>  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.<ref name="Jennings"/>

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.<ref name="Jennings"/><ref name="Milner-Gulland and Mace 1998"/>

The formula for maximum sustained harvest (<math>H</math>) is one-fourth the maximum population or carrying capacity (<math>K</math>) times the intrinsic rate of growth (<math>r</math>).<ref>Bolden and Robinson 1999</ref>

<math> H = \frac{Kr}{4}</math>

=== 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.<ref name="Jennings"/><ref name="Milner-Gulland and Mace 1998"/> 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.<ref name="Milner-Gulland and Mace 1998"/>

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.<ref>National Research Council (NRC). 1998. Improving Fish Stock Assessments. National Academy Press, Washington, D.C.</ref> 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.<ref>Clark 1990</ref><ref>National Marine Fisheries Service (NMFS). 1996. OUr Living Oceans: Report on the Status of U.S. Living Marine Resources 1995. NOAA Technical Memorandum NMFS0F/SPO-19. NMFS, Silver Springs, Md.</ref>  Like [[optimum sustainable yield]], 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.<ref name="Milner-Gulland and Mace 1998"/>  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.<ref name="Mace"/><ref name="Townsend">Townsend et al 2008</ref>  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.<ref name="Townsend"/>

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.<ref name="Townsend"/>  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.<ref name="Mace"/>

For a more contemporary description of MSY and its calculation see <ref name="Maunder">Maunder 2008</ref>  

==== Orange roughy ====
{{see also|Orange roughy}}
An example of errors in estimating the [[population dynamics]] of a species occurred within the New Zealand [[Orange roughy]] 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]] lived a long time and had [[k-selection|bred slowly]] (~30 years). By this stage stocks had been largely depleted.{{Citation needed|date=February 2011}}

== Overfishing ==
{{see also|Overfishing}}
All around the world, from the arctic to the tropics, there is a crisis in the world’s fisheries.{{http://www.sciencemag.org/content/314/5800/787.short}}  Until fairly recently it was assumed that our marine resources were limitless.{{Citation needed|date=February 2011}}

In recent years however, an accelerating decline has been observed in the productivity of many important fisheries.<ref>Christy and Scott 1965</ref>  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.<ref name="Clark 1973">Clark 1973</ref>  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.<ref>FAO, Review of the State of World Marine Fishery Resources, FAO Technical Paper 335 (1994).</ref>

In addition, the composition of global catches has changed.<ref>Roberts 2007)</ref>  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.{{Citation needed|date=February 2011}}

Overfishing is a classic example of the [[tragedy of the commons]].<ref name="Clark 1973"/>

== Optimum sustainable yield ==
{{see also|Optimum sustainable yield}}
In [[population ecology]] and [[economics]], '''optimum sustainable yield''' is the [[Level of Effort|level of effort]] (LOE) 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]], '''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]]
* [[Population dynamics of fisheries]]
* [[Population dynamics]]
* [[Fisheries management]]

== Notes ==
{{Page numbers needed|date=February 2011}}
{{reflist|2}}

==References==
{{refbegin|2}}
* Bolden, E.G., Robinson, W.L. (1999), [http://books.google.co.nz/books?id=PL6r1Ir_x_oC&printsec=frontcover&dq=%22Wildlife+ecology+and+management%22&ei=ugW6SpjREJCINMbR1L8P#v=onepage&q=&f=false ''Wildlife ecology and management''] 4th ed. Prentice-Hall, Inc. Upper Saddle River, NJ. ISBN 0-13-840422-4
* Botsford, L.W.,  Castilla, J.C., and Peterson, C.H. (1997), [http://mk.geog.uu.nl/homepages/Peter/teaching/Themes/Botsford.pdf "The management of fisheries and marine ecosystems"] ''Science'' '''277''': 509-515. {{doi|10.1126/science.22.5325.509}}
* Bousquet, N., Duchesne, T., Rivest, L.-P. (2008). [http://mat.ulaval.ca/pages/duchesne/BDR.pdf  "Redefining the maximum sustainable yield for the Schaefer population model including multiplicative environmental noise"] ''Journal of Theoretical Biology.'' '''254''': 65-75. {{doi|10.1016/j.jtbi.2008.04.025}}
* Christy, F.T., and Scott, A.D. (1965), ''The common Wealth in Ocean Fisheries'', Johns Hopkins Press, Baltimore
* Clark, C.W. (1973), [http://www.sciencemag.org/cgi/content/abstract/181/4100/630 "The Economics of Overexploitation"] ''Science'' '''118''': 630-634
* Clark, C.W. (1990), ''Mathematical Bioeconomics: The Optimal Management of Renewable Resources'', 2nd ed. Wiley-Interscience, New York
* Graham, M. (1935), "Modern theory of exploiting a fishery, an application to North Sea trawling", ''Journal de Conseil International pour l’Exploration de la Mer'' '''10'''" 264-274.
* Hjort, J., Jahn, G. and Ottestad, P. (1933), "The optimum catch." ''Hvalradets Skrifter'' '''7''': 92-127.
* Jennings, S., Kaiser, M.J. and Reynolds, J.D. (2001), [http://books.google.com/books?id=_J_E8O33E2gC&printsec=frontcover&dq=%22Marine+Fisheries+Ecology%22&ei=uge6SrG2H4vUNN361bwP#v=onepage&q=&f=false ''Marine Fisheries Ecology''] Blackwell Science Ltd. Malden, MA. ISBN 0-632-05098-5
* Larkin PA (1977) [http://docs.google.com/viewer?a=v&q=cache:0ZITmBnzlDUJ:fiesta.bren.ucsb.edu/~gsd/595e/docs/22.%2520Larkin_Epitaph_Max_Sust_Yield.pdf+Larkin+1977+%22An+epitaph+for+the+concept+of+maximum+sustained+yield%22&hl=en&gl=nz&sig=AHIEtbSBFh-w5xzWiBK7rnHSLBmolsfaWA "An epitaph for the concept of maximum sustained yield"] ''Transactions of the American Fisheries Society'', '''106''': 1–11.
* Mace, P.M. (2001), [http://courses.washington.edu/susfish/2002/p.mace.pdf "A new role for MSY in single-species and ecosystem approaches to fisheries stock assessment and management"] ''Fish and Fisheries'' '''2''': 2-32.
* Maunder, M.N. (2008) [http://www.sciencedirect.com/science/article/pii/B978008045405400522X "Maximum Sustainable Yield"] ''Encyclopedia of Ecology'' '''5''': 2292-2296
* Milner-Gulland, E.J., Mace, R. (1998), [http://books.google.co.nz/books?id=SfozQRjheekC&printsec=frontcover&dq=%22Conservation+of+biological+resources%22&ei=lsH5SfPYKoHgkwTh2dmyBA ''Conservation of biological resources''] Wiley-Blackwell. ISBN 978-0-86542-738-9.
* [[Callum Roberts|Roberts, C.]] (2007), ''The Unnatural History of the Sea'', Island Press. ISBN 978-1-59726-102-9
* Russell, E.S. (1931), [http://icesjms.oxfordjournals.org/cgi/pdf_extract/6/1/3 "Some theoretical considerations on the ‘overfishing’ problem"] ''Journal de Conseil International pour l’Explaration de la mer'' '''6''': 1-20.
* Schaefer, M.B. (1954), [http://www.springerlink.com/content/a428w467h7667564/ "Some aspects of the dynamics of populations important to the management of the commercial marine fisheries"] ''Bulletin of the Inter-American Tropical Tuna Commission'' '''1''': 25-56.
* Sissenwine, M.P. (1978), [http://afsjournals.org/doi/abs/10.1577/1548-8446%281978%29003%3C0022:IMAAFF%3E2.0.CO%3B2 "Is MSY an adequate foundation for optimum yield?"] ''Fisheries'' '''3''': 22-42.
* Townsend, C.R., Begon, M., and Harper, J.L. (2008), [http://books.google.com/books?id=pQTNFYPgDdEC&printsec=frontcover&dq=%22Essentials+of+Ecology%22&ei=vBC6SpG7IJeQMuGNmbAP#v=onepage&q=&f=false ''Essentials of Ecology''] Blackwell Publishing. ISBN 978-1-4051-5658-5
* [[Carl Walters|Walters C]] and Maguire J (1996) "Lessons for stock assessment from the northern cod collapse", ''Reviews in Fish Biology and Fisheries'', '''6''':125–137.</ref>
{{refend}}

{{fishery science topics|expanded=management}}
{{modelling ecosystems|expanded=none}}

[[Category:Fisheries science]]
[[Category:Conservation]]
[[Category:Economics terminology]]
[[Category:Renewable resources]]
[[Category:Ecological metrics]]
[[Category:Fishing and the environment]]