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Theoretical production ecology

From Wikipedia, the free encyclopedia

Theoretical production ecology tries to quantitatively study the growth of crops. The plant is treated as a kind of biological factory, which processes light, carbon dioxide, water, and nutrients into harvestable parts. Main parameters kept into consideration are temperature, sunlight, standing crop biomass, plant production distribution, nutrient and water supply.

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Transcription

There's a lot of ideas that we just assume we know a lot about because we hear about them all the time. For instance, I know what Pop music is, but if you were to corner me at a party and say, "HANK, What is Pop Music?", I'd be like, "It's uh... it's like, uh... the music that plays on the pop station?" Just because we're familiar with a concept does not mean that we actually understand it. Ecology's kind of the same way: even though it's a common, everyday concept, and ecosystem is a word that we hear a lot, I think most of us would be little stumped if somebody actually asked us what an ecosystem is or how one works, or why they're important, etc. I find it helps to think of an ecosystem, a collection of living and nonliving things interacting in a specific place, as one of those Magic Eye posters, for those of you who were sentient back in 1994. An ecosystem is just a jumble of organisms, weather patterns, geology and other stuff that don't make a lot of sense together until you stare at them long enough, from far enough away, and then suddenly a picture emerges. And just like with Magic Eye posters, it helps if you're listening to Jamiroquai while you're doing it. So, the discipline of ecosystem ecology, just like other types of ecology we've been exploring lately, looks at a particular level of biological interaction on Earth. But unlike population ecology, which looks at interactions between individuals of one species, or community ecology, which looks at how bunches of living things interact with each other, ecosystem ecology looks at how energy and materials come into an ecosystem, move around in it, and then get spat back out. In the end, ecosystem ecology is mostly about eating, who's eating whom, and how energy, nutrients and other materials are getting shuffled around within the system. So today, we're setting the record straight! No more not understanding how an ecosystem works! Starting NOW! So, ecosystems may be a lot like Magic Eye posters, but the way that they're not like a Magic Eye poster is in the way that posters have edges. Ecosystems... I'll just come out and say it: No edge. Only fuzzy, ill-defined gradients that bleed into the ecosystems next door. So actually defining an ecosystem can be kind of hard. Mostly it depends on what you want to study. Say you're looking at a stream in the mountains. This stream gets very little sunlight because it's so small that the trees on its banks totally cover it with shade. As a result, very few plants or algae live in it, and if there's one thing that we know about planet Earth it's that plants are king. Without plants, there are no animals. But somehow there's a whole community of animals living in and around this mountain stream, even though there are few plants in it. So what are the animals doing there, and how are they making their living? From the land, of course! From the ecosystems around it. Because no stream is an island. It isn't there all by itself. All kinds of food, nutrients and other materials drop into the stream from the trees or are washed into it when it rains, leaves and bugs, you name it, flow down from neighboring terrestrial ecosystems. And that stuff gets eaten by bigger bugs, which get eaten by fish, which in turn are eaten by raccoons and birds and bears. So, even though the stream's got its own thing going on, without the rest of the watershed, the animals there wouldn't survive. And without the stream, plants would be thirsty and terrestrial animals wouldn't have as many fish to eat. So where does the ecosystem of the stream start and where does it end? This is a perennial problem for ecologists. Because the way it works, energy and nutrients are imported in from someplace, they're absorbed by the residents of an ecosystem, and then passed around within it for a little while, and then finally passed out, sometimes into another ecosystem. This is most obvious in aquatic systems, where little streams eventually join bigger and bigger waterways until they finally reach the ocean, this flow is a fundamental property of ecosystems. So, at the end of the day, how you define an ecosystem just depends on what you want to know. If you want to know how energy and materials come in, move through, and are pooped out of a knot in a tree that has a very specific community of insects and protists living in it, you can call that an ecosystem. If you want to know how energy and materials are introduced to, used and expelled by the North Pacific Gyre, you can call that an ecosystem. If you want to know how energy and materials move around a cardboard box that has a rabbit and a piece of lettuce in it, you can call that an ecosystem. I might tell you that your ecosystem is stupid, but go ahead! Do whatever you want! The picture you see in an ecosystem's Magic Eye is actually dictated by the organisms that live there and how they use what comes into it. An ecosystem can be measured through figuring out things like its biomass, that is, the total weight of living things in the ecosystem, and its productivity, how much stuff is produced, and how quickly stuff grows back, how good the ecosystem is at retaining stuff. And of course, all these parameters matter to neighboring ecosystems as well because if one ecosystem's really productive, the ones next door are going to benefit. So first things first, where do the energy and materials come from? And to be clear, when I talk about "materials," I'm talking about water or nutrients like phosphorous or nitrogen, or even toxins like mercury or DDT. Let's start out by talking about energy, because nothing lives without energy, and where organisms get their energy tells the story of an ecosystem. You remember physics, right? The laws of conservation state that energy and matter can neither be destroyed or created. They can only get transferred from place to place to place. The same is true of an ecosystem. Organisms in an ecosystem organize themselves into a trophic structure, with each organism situating itself in a certain place in the food chain. All of the energy in an ecosystem moves around within this structure, because when I say energy, of course I mean food. For most ecosystems, the primary source of energy is the sun, and the organisms that do most of the conversion of solar energy into chemical energy...you know this one. Who rules the world? The plants rule the world. Autotrophs like plants are able to gather up the sun's energy, and through photosynthesis, make something awesome out of it: little stored packets of chemical energy. So whether it's plants, bacteria or protists that use photosynthesis, autotrophs are always the lynchpin of every ecosystem, the foundation upon which all other organisms in the system get their energy and nutrients. For this reason, ecologists refer to plants as primary producers. Now, obviously, the way that energy gets transferred from plants to animals is by an animal eating the plant. For this reason, herbivores are known as primary consumers, the first heterotrophs to get their grubby paws on that sweet, sweet energy. After this stage in the trophic structure, the only way to wrestle the solar energy that was in the plants that the herbivore ate is to, you guessed it, eat the herbivore, which carnivores, known as secondary consumers, are very happy to do. And assuming that the ecosystem is big enough and productive enough, there might even be a higher level of carnivore that eats other carnivores, like an owl that eats hawks, and these guys are called tertiary consumers. And then there are the -vores that decompose all the dead animal and plant matter, as well as the animal poop: detritivores. These include earthworms, sea stars, fiddler crabs, dung beetles, fungi, and anything else that eats the stuff that none of the rest of us would touch with a 3-meter pole. So, that's a nice, hierarchical look at who's getting energy from what or whom within an ecosystem. But of course, organisms within an ecosystem don't usually abide by these rules very closely, which is why these days, we usually talk about food webs, rather than food chains. A food web takes into consideration that sometimes a fungus is going to be eating nutrients from a dead squirrel, and other times squirrels are going to be eating the fungi. Sometimes a bear likes to munch on primary producers, blueberry bushes, and other times it's going to be snacking on a secondary consumer, a salmon. And even the tippy tippy top, predators get eaten by stuff like bacteria in the end, which might or might not be the same bacteria that ate the top predator's poopies. Circle of Life! It's also worth noting that the size and scope of the food web in an ecosystem has a lot to do with things like water and temperature, because water and temperature are what plants like, right? And without plants, there isn't going to be a whole lot of trophic action going on. Take, for example, the Sonoran desert, which we've talked about before. There aren't very many plants there, compared to, say, the Amazon rainforest. So the primary producers are limited by the lack of water, which means that primary consumers are limited by lack of primary producers, and that leaves precious few secondary consumers, a few snakes, some coyotes and hawks. All this adds up to the Sonoran not being a terribly productive place, compared to the Amazon at least, so you might only get to the level of tertiary consumer occasionally. Now, all this conversation about productivity leads me to another point about ecosystem efficiency. When I talk about energy getting passed along from one place to another within an ecosystem, I mean that in a general sense, organisms are sustaining each other, but not in a particularly efficient way. In fact, when energy transfers from one place to another, from a plant or a bunny or from a bunny to a snake, the vast majority of that energy is lost along the way. So, let's take a cricket. That cricket has about 1 calorie of energy in it. And in order to get that 1 calorie of energy it had to eat about 10 calories of lettuce. Where did the other 9 calories go? It is not turned into cricket flesh. Most of it is used just to live, like to power its muscles, or run the sodium potassium pumps in its neurons, it's just used up. So only the 1 calorie of the original 10 calories of food is left over as actual cricket stuff. And then, right after his last meal, the cricket jumps into a spider web and is eaten by a spider, who converts only 10% of the cricket's energy into actual spider stuff. And don't get me started on the bird that eats the spider. This is not an efficient world that we live in. But you want to know what's scary-efficient? The accumulation of toxins in an ecosystem. Elements like mercury, which are puffed out the smokestacks of coal-fired power plants, end up getting absorbed in the ocean by green algae and marine plants. While the tiny animal that eats the algae only stores 10% of the energy it got, it keeps 100% of the mercury. So as we move up the chain, each trophic level consumes ten times more mercury than the last, and that's what we call bioaccumulation. Concentrations get much higher at each trophic level, until a human gets a hold of a giant tuna that's at the top of the marine food chain, and none of that mercury has been lost. It's all right there in that delicious tuna flesh. Because organisms only hold on to 10% of the energy they ingest, each trophic level has to eat about 10 times its biomass to sustain itself. And because 100% of the mercury moves up the food chain, that means that it becomes 10 times more concentrated with each trophic level it enters. That's why we need to take the seafood advisories seriously: as somebody who could eat anything you wanted, it's probably safest to eat lower on the food chain, primary producers or primary consumers. The older, bigger, higher in the food chain, the more toxic it's going to be. And that's not just my opinion, that's ecosystem ecology! Thank you for watching this episode of Crash Course Ecology. And thank you for everyone who helped us put this episode together. If you want to reviews any of the topics we went over today, there's a table of contents over there that you can click on. And if you have any questions or comments for us we're on Facebook or Twitter, or of course, down in the comments below. We'll see you next time.

Modelling

Modelling is essential in theoretical production ecology. Unit of modelling usually is the crop, the assembly of plants per standard surface unit. Analysis results for an individual plant are generalised to the standard surface, e.g. the leaf area index is the projected surface area of all crop leaves above a unit area of ground.

Processes

The usual system of describing plant production divides the plant production process into at least five separate processes, which are influenced by several external parameters.

Two cycles of biochemical reactions constitute the basis of plant production, the light reaction and the dark reaction.[1]

  • In the light reaction, sunlight photons are absorbed by chloroplasts which split water into an electron, proton and oxygen radical which is recombined with another radical and released as molecular oxygen. The recombination of the electron with the proton yields the energy carriers NADH and ATP. The rate of this reaction often depends on sunlight intensity, leaf area index, leaf angle and amount of chloroplasts per leaf surface unit. The maximum theoretical gross production rate under optimum growth conditions is approximately 250 kg per hectare per day.
  • The dark reaction or Calvin cycle ties atmospheric carbon dioxide and uses NADH and ATP to convert it into sucrose. The available NADH and ATP, as well as temperature and carbon dioxide levels determine the rate of this reaction. Together those two reactions are termed photosynthesis. The rate of photosynthesis is determined by the interaction of a number of factors including temperature, light intensity and carbon dioxide.
  • The produced carbohydrates are transported to other plant parts, such as storage organs and converted into secondary products, such as amino acids, lipids, cellulose and other chemicals needed by the plant or used for respiration. Lipids, sugars, cellulose and starch can be produced without extra elements. The conversion of carbohydrates into amino acids and nucleic acids requires nitrogen, phosphorus and sulfur. Chlorophyll production requires magnesium, while several enzymes and coenzymes require trace elements. This means, nutrient supply influences this part of the production chain. Water supply is essential for transport, hence limits this too.
  • The production centers, i.e. the leaves, are sources, the storage organs, growth tips or other destinations for the photosynthetic production are sinks. The lack of sinks can be a limiting factor for production too, as happens e.g. in apple orchards where insects or night frost have destroyed the blossoms and the produced assimilates cannot be converted into apples. Biennial and perennial plants employ the stored starch and fats in their storage organs to produce new leaves and shoots the next year.
  • The amount of crop biomass and the relative distribution of biomass over leaves, stems, roots and storage organs determines the respiration rate. The amount of biomass in leaves determines the leaf area index, which is important in calculating the gross photosynthetic production.
  • extensions to this basic model can include insect and pest damage, intercropping, climatic changes, etc.

Parameters

Important parameters in theoretical production models thus are:

Climate
  • Temperature – The temperature determines the speed of respiration and the dark reaction. A high temperature combined with a low intensity of sunlight means a high loss by respiration. A low temperature combined with a high intensity of sunlight means that NADH and ATP heap up but cannot be converted into glucose because the dark reaction cannot process them swiftly enough.
  • Light – Light, also called photosynthetic Active Radiation (PAR) is the energy source for green plant growth. PAR powers the light reaction, which provides ATP and NADPH for the conversion of carbon dioxide and water into carbohydrates and molecular oxygen. When temperature, moisture, carbon dioxide and nutrient levels are optimal, light intensity determines maximum production level.
  • Carbon dioxide levels – Atmospheric carbon dioxide is the sole carbon source for plants. About half of all proteins in green leaves have the sole purpose of capturing carbon dioxide.
Although CO2 levels are constant under natural circumstances [on the contrary, CO2 concentration in the atmosphere has been increasing steadily for 200 years], CO2 fertilization is common in greenhouses and is known to increase yields by on average 24% [a specific value, e.g., 24%, is meaningless without specification of the "low" and "high" CO2 levels being compared].[2]
C4 plants like maize and sorghum can achieve a higher yield at high solar radiation intensities, because they prevent the leaking of captured carbon dioxide due to the spatial separation of carbon dioxide capture and carbon dioxide use in the dark reaction. This means that their photorespiration is almost zero. This advantage is sometimes offset by a higher rate of maintenance respiration. In most models for natural crops, carbon dioxide levels are assumed to be constant.
Crop
  • Standing crop biomass – Unlimited growth is an exponential process, which means that the amount of biomass determines the production. Because an increased biomass implies higher respiration per surface unit and a limited increase in intercepted light, crop growth is a sigmoid function of crop biomass.
  • Plant production distribution – Usually only a fraction of the total plant biomass consists of useful products, e.g. the seeds in pulses and cereals, the tubers in potato and cassava, the leaves in sisal and spinach etc. The yield of usable plant portions will increase when the plant allocates more nutrients to this parts, e.g. the high-yielding varieties of wheat and rice allocate 40% of their biomass into wheat and rice grains, while the traditional varieties achieve only 20%, thus doubling the effective yield.
Different plant organs have a different respiration rate, e.g. a young leaf has a much higher respiration rate than roots, storage tissues or stems do. There is a distinction between "growth respiration" and "maintenance respiration".
Sinks, such as developing fruits, need to be present. They are usually represented by a discrete switch, which is turned on after a certain condition, e.g. critical daylength has been met.
Care
  • Water supply – Because plants use passive transport to transfer water and nutrients from their roots to the leaves, water supply is essential to growth, even so that water efficiency rates are known for different crops, e.g. 5000 for sugar cane, meaning that each kilogram of produced sugar requires up to 5000 liters of water.
  • Nutrient supply – Nutrient supply has a twofold effect on plant growth. A limitation in nutrient supply will limit biomass production as per Liebig's Law of the Minimum. With some crops, several nutrients influence the distribution of plant products in the plants. A nitrogen gift is known to stimulate leaf growth and therefore can work adversely on the yield of crops which are accumulating photosynthesis products in storage organs, such as ripening cereals or fruit-bearing fruit trees.

Phases in crop growth

Theoretical production ecology assumes that the growth of common agricultural crops, such as cereals and tubers, usually consists of four (or five) phases:

  • Germination – Agronomical research has indicated a temperature dependence of germination time (GT, in days). Each crop has a unique critical temperature (CT, dimension temperature) and temperature sum (dimensions temperature times time), which are related as follows.
When a crop has a temperature sum of e.g. 150 °C·d and a critical temperature of 10 °C, it will germinate in 15 days when temperature is 20 °C, but in 10 days when temperature is 25 °C. When the temperature sum exceeds the threshold value, the germination process is complete.
  • Initial spread – In this phase, the crop does not cover the field yet. The growth of the crop is linearly dependent on leaf area index, which in its turn is linearly dependent on crop biomass. As a result, crop growth in this phase is exponential.
  • Total coverage of field – in this phase, growth is assumed to be linearly dependent on incident light and respiration rate, as nearly 100% of all incident light is intercepted. Typically, the Leaf Area Index (LAI) is above two to three in this phase. This phase of vegetative growth ends when the plant gets a certain environmental or internal signal and starts generative growth (as in cereals and pulses) or the storage phase (as in tubers).
  • Allocation to storage organs – in this phase, up to 100% of all production is directed to the storage organs. Generally, the leaves are still intact and as a result, gross primary production stays the same. Prolonging this phase, e.g. by careful fertilization, water and pest management results directly in a higher harvest.
  • Ripening – in this phase, leaves and other production structures slowly die off. Their carbohydrates and proteins are transported to the storage organs. As a result, the LAI and, hence, the primary production decreases.

Existing plant production models

Plant production models exist in varying levels of scope (cell, physiological, individual plant, crop, geographical region, global) and of generality: the model can be crop-specific or be more generally applicable. In this section the emphasis will be on crop-level based models as the crop is the main area of interest from an agronomical point of view.

As of 2005, several crop production models are in use. The crop growth model SUCROS has been developed during more than 20 years and is based on earlier models. Its latest revision known dates from 1997. The IRRI and Wageningen University more recently developed the rice growth model ORYZA2000. This model is used for modeling rice growth. Both crop growth models are open source. Other more crop-specific plant growth models exist as well.

SUCROS

SUCROS is programmed in the Fortran computer programming language. The model can and has been applied to a variety of weather regimes and crops. Because the source code of Sucros is open source, the model is open to modifications of users with FORTRAN programming experience. The official maintained version of SUCROS comes into two flavours: SUCROS I, which has non-inhibited unlimited crop growth (which means that only solar radiation and temperature determine growth) and SUCROS II, in which crop growth is limited only by water shortage.

ORYZA2000

The ORYZA2000 rice growth model has been developed at the IRRI in cooperation with Wageningen University. This model, too, is programmed in FORTRAN. The scope of this model is limited to rice, which is the main food crop for Asia.

Other models

The United States Department of Agriculture has sponsored a number of applicable crop growth models for various major US crops, such as cotton, soy bean, wheat and rice.[3] Other widely used models are the precursor of SUCROS (SWATR), CERES, several incarnations of PLANTGRO, SUBSTOR, the FAO-sponsored CROPWAT, AGWATER, the erosion-specific model EPIC,[4] and the cropping system CropSyst.[5]

A less mechanistic growth and competition model, called the conductance model, has been developed, mainly at Warwick-HRI, Wellesbourne, UK. This model simulates light interception and growth of individual plants based on the lateral expansion of their crown zone areas. Competition between plants is simulated by a set algorithms related to competition for space and resultant light intercept as the canopy closes. Some versions of the model assume overtopping of some species by others. Although the model cannot take account of water or mineral nutrients, it can simulate individual plant growth, variability in growth within plant communities and inter-species competition. This model was written in Matlab. See Benjamin and Park (2007) Weed Research 47, 284–298 for a recent review.

References

  • Theoretical Production Ecology, college notes, Wageningen Agricultural University, 1990
  1. ^ Amthor JS (2010) From sunlight to phytomass: on the potential efficiency of converting solar radiation to phyto-energy. New Phytologist 188:939-959
  2. ^ "Carbon dioxide fertilization is neither boon nor bust". EurekAlert!.
  3. ^ "Available Crop Models : USDA ARS". www.ars.usda.gov.
  4. ^ "Crop growth models". Archived from the original on 2005-12-20. Retrieved 2005-07-30.
  5. ^ "CS_Suite - Dr. Claudio Stöckle WSU". Archived from the original on 2010-05-31. Retrieved 2014-01-05.

Further reading

This page was last edited on 11 June 2024, at 17:40
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