Ecology-Stimulation of different functional groups of bacteria by various plant residues
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TPO69阅读-1Why Snakes Have Forked Tongues原文 (1)译文 (3)题目 (4)答案 (8)背景知识 (9)原文Why Snakes Have Forked Tongues①The forked tongue of snakes has intrigued people for millennia,inspiring many hypotheses.In many cultures and religions,the forked tongue symbolizes malevolence and deceit.The first person known to inquire about the functional significance of the forked tongue was Aristotle;he suggested that it would double the pleasure of sensations of taste.By the beginning of the twentieth century,the consensus was that the snake’s tongue is a tactile organ;that is,the snake uses it to tap the ground much as a blind person uses a cane.②In1920Browman suggested what seemed to be a winning hypothesis:When the snake retracts its tongue,the tips(or tines)of the forked tongue are inserted into openings on both sides of the roof of the mouth;through these openings chemical stimuli reach special organs that help snakes detect smells—the vomeronasal organs(VNO).These organs are highly developed in snakes,lizards, and many mammals.They are a second system for detecting smells that appears to have evolved specifically to detect pheromones,the chemical signals that animals secrete as messages to other animals of their species.Browman suggested that the forked tongue flicks out,picking up chemical signals,and then delivers these to the VNO.This hypothesis was widely accepted into the1980s.Then X-ray movie studies of tongue flicks in snakes and lizards with forked tongues disproved the hypothesis;they showed that when the tongue is withdrawn into the mouth,it enters a sheath and the tips do not go into the openings to the VNO.Instead,the chemical molecules are deposited on pads at the bottom of the mouth,and closing the mouth presses the pads and molecules against the VNO openings.③If the tongue is not forked to fit into the VNO,then what function could the forked shape serve?Schwenk proposes a solution that encompasses observations from several fields—animal behavior,ecology,sensory physiology,and neuroanatomy.He hypothesizes that the forked tongue allows the snake to sense chemical stimuli at two points simultaneously,thereby giving it the ability to detect differences in an odor trail.Obtaining two simultaneous readings enhances the ability of the snake to detect the edges of odor trails,and thus to follow pheromone trails accurately.This ability is important in seeking both prey and mates.④This spatial chemical perception is like other systems for spatial perception that are based on simultaneous stimulation of two separated sense organs—for example,auditory localization,which depends on differential stimulation at the two ears.Similarly,the use of the two eyes permits stereovision.⑤Several kinds of evidence support the hypothesis that forked tongues evolved as chemosensory edge detectors to enhance the ability to follow odor trails:(1) Snakes and lizards spread the tines of their tongue apart when they retrieve odor molecules,then draw the tines together when retracting the tongue.The greater the distance between sampling points,the better the animals sample differences within an odor trail.(2)Lizards that forage widely have forked tongues,whereas lizard species without forked tongues tend not to forage widely.(3)Forked tongues have evolved independently at least twice in different families of reptiles, indicating their value as an adaptation.(4)In the snake nervous system,each tine of the tongue is linked to a nucleus in the other side of the brain,and the two nuclei are linked across the two hemispheres.This arrangement is similar to the anatomy of auditory centers in mammals and birds that permits the computation of differences between what one ear hears and what the other ear hears and thus mediates auditory localization.⑥Species in other orders have also evolved paired chemical receptors to guide individuals to mates or prey.For example,male gypsy moths have large,elaborate, odor-detecting antennae with which they track potential mates over large distances,and the ant nest beetle has spoon-shaped antennae extending from each side of the head with which it detects and follows the pheromones of the ants that are its food.译文为什么蛇有叉状舌头①蛇的叉状舌头吸引了人们数千年,激发了许多假设。
全国2023-2024学年英语高三上期末学业水平测试模拟试题注意事项:1.答题前,考生先将自己的姓名、准考证号码填写清楚,将条形码准确粘贴在条形码区域内。
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第一部分(共20小题,每小题1.5分,满分30分)1.She is ____________ being pleased about it ; she is very angry.A.free from B.free of C.out of D.far from2.Working hard is not a __________ of great success, but it is among the essential requirements.A.sign B.signalC.guarantee D.mark3.I ______ tell you with cer tainty that he won’t be able to finish it before the deadline.A.will B.must C.can D.may4.It really matters _______ he treated the latest failure, for the examination is around the corner.A.if B.thatC.why D.how5.When they first came to the city, my parents often went to neighbors for a talk, just as they ________ in the countryside.A.will do B.had doneC.have done D.were doing6.I’m most awfully grateful to you. I can’t think what I ________ without you. A.could do B.would doC.should have done D.must have done7.You can’t use the computer now, ________ the upgrade of the system is under way. A.until B.unlessC.as D.after8.Every four years Americans __________ the transfer of power.A.carry over B.carry offC.carry on D.carry out9.Nowadays, more and more young ladies, figures most are fine enough, are going on a diet.A.who B.whoseC.of whose D.of whom10.Justin a book about his adventures in Tibet.I hope he can find a good publisher when it is finished.A.was writing B.is currently writing C.has already written D.wrote11.Don’t forget to send ______ attended the conference a follow-up email. A.however B.whateverC.whoever D.wherever12.Painting from still images leads to a loss of sensitivity, which is _____ to an artist. A.absolute B.urgent C.especial D.vital13.The farmers are in bad need of rain,but a (an) large amount of rainfall will cause Floods.A.Eventually B.constantly C.gradually D.extremely14.He liked the lovely dog so much that he ________ his book and played with it immediately.A.gave off B.set asideC.took over D.turned down15.—You ought to have made an apology to Tom yesterday evening.—Yes, I know I __ __.A.ought to have B.have to C.should D.must have16.He couldn’t make his voi ce ______ above the noise of the traffic.A.to hear B.hearingC.heard D.hear17.The main issue at the APEC meeting was a climate-change plan _____ by Australia’s Howard and backed by Bush.A.put out B.put offC.put away D.put forward18.________ the danger of drunk driving, many drivers promised never to drive after drinking.A.Realizing B.To have realizedC.Realized D.Having realized19.Beijing’s new international airport into operation in 2019 will serve 72 million passengers annually.A.being put B.to be putC.put D.to put20.______ for the free tickets, I would not have gone to see films so often.A.If it is not B.Had it not beenC.Were it not D.If they were not第二部分阅读理解(满分40分)阅读下列短文,从每题所给的A、B、C、D四个选项中,选出最佳选项。
BMPs (GDF2)介绍骨形态发生蛋白( BMPs ) 属于转化生长因子-β 超家族,其在骨的再生和修复中起重要作用。
许多的研究还发现BMP 参与调节许多种细胞的增殖、分化和凋亡的生物学过程1BMPs家族的种类骨形态发生蛋白( bone morphogenetic protein, BMP) 属于转化生长因子- β( TGF- β) 超家族, 该家族已经发现40 多个成员,包括BMPs、生长和分化因子( GDFs)、活化素( activins)、抑制素( inhibins)、缪勒管抑制质( Muller ian inhibitor substance , MIS)和TGF -β[1-5]。
TGF -β 超家族成员是以前体大分子的形式合成,经过蛋白水解酶切割从前肽区释放出成熟蛋白[6]。
已经发现的BMPs有15 种, 根据它们氨基酸序列的类似分为 3 个亚群:BMP -2 和BMP -4 、BMP -5、BMP -8、BMP -3 和GDF -10。
除BMP -1外,每个成员都有骨诱导的功能[7]。
BMPs 对肿瘤细胞增殖的影响BMP2/4已被证实可以抑制肿瘤细胞的增殖。
Brubaker等[8]在前列腺肿瘤细胞中发现,采用BMP2/4处理的肿瘤细胞增殖能力较对照组明显降低(P<0.05),其作用机制主要是通过Smad1信号通路上调肿瘤抑制基因细胞周期蛋白依赖性激酶抑制剂p21的表达,并引起下游Rb蛋白失活,从而抑制增殖。
目前有关BMP2/4对肿瘤细胞的抑制作用主要表现在激素敏感型肿瘤细胞中(如LNCaP,MDA-MB-231),对非敏感型细胞则作用不明显,这可能是由于性激素在BMP2/4信号转导中起开关作用有关。
另一方面,Langenfeld等[9]在小鼠体内实验证实,BMP2蛋白含量与肺癌组织中肿瘤细胞的增殖成正比,其机制可能为Smad1/5通路上调Id-1和pERK-1/2的表达;而在无血清培养基中,BMP2则发挥相反作用。
动物园用于保护野生动物的必要性!英文作文全文共3篇示例,供读者参考篇1The Necessity of Zoos for Protecting WildlifeIntroductionZoos play a crucial role in conserving and protecting wildlife. They provide a safe haven for endangered species and facilitate breeding programs to ensure their survival. In this essay, we will explore the importance of zoos in protecting wildlife and discuss the various ways in which they contribute to conservation efforts.Preservation of Endangered SpeciesOne of the primary functions of zoos is to preserve endangered species. Many animals face the threat of extinction due to habitat loss, poaching, and other human activities. Zoos provide a secure environment for these animals to live and breed, thereby increasing their chances of survival. By maintaining healthy populations of endangered species in captivity, zoos help to prevent their extinction and ensure that future generations will be able to admire these magnificent creatures.Breeding ProgramsZoos play a vital role in breeding programs for endangered species. Many animals bred in zoos are released into the wild to bolster wild populations and restore balance to ecosystems. These programs are essential for maintaining genetic diversity and preventing inbreeding, which can lead to health issues and reduced fertility in animals. By carefully managing breeding programs, zoos can help to ensure the long-term survival of endangered species.Education and AwarenessZoos also serve as centers for education and awareness about wildlife conservation. They provide opportunities for visitors to learn about different species, their habitats, and the threats they face in the wild. Through educational programs, interpretive displays, and interactive exhibits, zoos help to raise awareness about the importance of protecting wildlife and inspire action to safeguard the environment. By connecting people with nature, zoos play a crucial role in fostering a sense of responsibility towards the planet and its inhabitants.Research and ConservationZoos are actively involved in research and conservation efforts to protect wildlife. They collaborate with scientists, conservationists, and government agencies to study endangered species, develop conservation strategies, and implement sustainable practices. Research conducted in zoos helps to advance our understanding of animal behavior, biology, and ecology, which in turn informs conservation initiatives in the wild. By supporting research and conservation projects, zoos contribute valuable knowledge and resources to protect endangered species and their habitats.Global ImpactThe work of zoos has a global impact on wildlife conservation. Many species are threatened by environmental degradation, climate change, and other global challenges that require coordinated efforts to address. Zoos play a crucial role in international conservation networks, sharing expertise, resources, and best practices to protect endangered species on a global scale. By working together, zoos can make a significant difference in preserving biodiversity and safeguarding the future of wildlife.ConclusionIn conclusion, zoos are essential for protecting wildlife and conserving biodiversity. They provide a safe haven for endangered species, facilitate breeding programs, educate the public about conservation, conduct research and conservation initiatives, and collaborate with global partners to address pressing environmental challenges. By supporting zoos and their conservation efforts, we can help to ensure a future where wildlife thrives and ecosystems are healthy. Let us all work together to protect and preserve the magnificent creatures that share our planet.篇2The Importance of Zoos in Protecting Wild AnimalsZoos have long been controversial institutions, with some arguing that they confine animals in unnatural environments and others arguing that they play a crucial role in wildlife conservation. While there are valid arguments on both sides, it is clear that zoos can be valuable tools in protecting and preserving endangered species.One of the primary ways in which zoos contribute to wildlife conservation is through breeding programs. Many zoos participate in Species Survival Plans (SSPs), which arecooperative programs designed to manage the breeding of endangered species in captivity. By carefully selecting breeding pairs and monitoring the health and genetic diversity of captive populations, zoos can help to ensure the long-term survival of endangered species. In some cases, zoos have even successfully reintroduced captive-bred animals into the wild, bolstering populations that are struggling in their natural habitats.In addition to breeding programs, zoos also play a key role in education and public awareness. Zoos provide an opportunity for people to see and learn about animals that they may never have the chance to encounter in the wild. This first-hand experience can foster a sense of connection and empathy towards animals, inspiring visitors to care about conservation issues and take action to protect wildlife. Zoos often offer educational programs and resources for schools and community groups, further spreading awareness about the importance of protecting endangered species.Furthermore, zoos can serve as important research centers for studying wildlife biology and behavior. By closely observing animals in captivity, researchers can gain valuable insights into the needs and behaviors of endangered species. This information can inform conservation efforts in the wild, helpingto develop effective strategies for protecting and managingat-risk populations. Zoo research can also contribute to our understanding of animal welfare and husbandry practices, ensuring that captive animals receive the best possible care.Despite these benefits, it is important to acknowledge that not all zoos are created equal. Some facilities prioritize animal welfare and conservation, while others prioritize profit and entertainment. It is crucial for zoos to adhere to high standards of animal care and conservation ethics, ensuring that they are contributing positively to wildlife conservation efforts. Accreditation by organizations such as the Association of Zoos and Aquariums can help to ensure that zoos are meeting these standards and operating responsibly.In conclusion, zoos can be valuable institutions for protecting and conserving wild animals. Through breeding programs, education initiatives, and research efforts, zoos can play a vital role in ensuring the survival of endangered species. However, it is essential for zoos to prioritize animal welfare and conservation ethics in order to maximize their positive impact on wildlife conservation. by working together with government agencies, conservation organizations, and the public, zoos canhelp to protect and preserve the incredible diversity of life on our planet.篇3The Necessity of Zoos in Conserving WildlifeZoos play a crucial role in conserving and protecting wildlife, particularly endangered species. Despite some criticisms about animal welfare and captivity, zoos serve as important institutions that contribute significantly to the preservation of species at risk of extinction. In this essay, we will explore the importance and necessity of zoos in wildlife conservation.One of the main reasons why zoos are essential for protecting wildlife is that they provide a safe and controlled environment for animals that are endangered in the wild. Many species face threats such as habitat loss, poaching, and climate change, which put them at risk of extinction. Zoos play a vital role in breeding programs and conservation efforts to ensure the survival of these species. Through careful management and breeding, zoos help to increase populations of endangered animals and reintroduce them into the wild when conditions are suitable.Moreover, zoos also serve as educational institutions that raise awareness about the importance of wildlife conservation. By showcasing a variety of species from around the world, zoos help to educate the public about the diversity of life on Earth and the threats facing many species. Visitors to zoos can learn about the habitats, behaviors, and conservation status of different animals, which can inspire them to take action to protect wildlife and the environment.Furthermore, zoos provide research opportunities for scientists and conservationists to study and understand endangered species. By studying animals in captivity, researchers can learn more about their biology, behavior, and reproduction, which can inform conservation strategies in the wild. Zoos also play a role in supporting field conservation projects by providing funding, expertise, and resources to protect endangered species and their habitats.In addition, zoos are important for promoting the welfare and well-being of individual animals. While some critics argue that animals in captivity may suffer from stress and confinement, modern zoos place a strong emphasis on providing enriching environments and care for their animals. Zoos invest in creating habitats that replicate the natural environments of animals,provide mental and physical stimulation, and ensure proper nutrition and veterinary care.In conclusion, the necessity of zoos in conserving wildlife cannot be overstated. Zoos play a crucial role in protecting endangered species, raising awareness about conservation issues, supporting research and education, and promoting the welfare of individual animals. While challenges exist in maintaining high standards of animal welfare in captivity, zoos continue to evolve and adapt to ensure the well-being of the animals in their care. By supporting zoos and their conservation efforts, we can help to safeguard the future of wildlife and advance our understanding of the natural world.。
Microbial Ecology of the GutThe gut microbiome is a complex ecosystem of microorganisms that play a crucial role in human health. Recent advances in sequencing technology have enabled researchers to identify and characterize the vast array of microorganisms that inhabit the gut. The gut microbiome has been linked to a wide range of health outcomes, including obesity, diabetes, and inflammatory bowel disease. In this essay, I will explore the microbial ecology of the gut from multiple perspectives, including the role of microbial diversity, the impact of diet and lifestyle, and the potential for therapeutic interventions.One of the key factors that influence the composition of the gut microbiome is microbial diversity. The gut microbiome is composed of a diverse array of microorganisms, including bacteria, viruses, fungi, and archaea. The diversity of the gut microbiome is important because it allows for the ecosystem to be resilient to perturbations. For example, if a particular species of bacteria is lost due to an antibiotic treatment, the ecosystem can recover by relying on other species to perform similar functions. In addition, a diverse microbiome can provide a range of metabolic functions that are important for human health, including the production of short-chain fatty acids, which are important for gut health and immune function.Another important factor that influences the gut microbiome is diet and lifestyle. The Western diet, which is high in fat and sugar, has been shown to reduce microbial diversity and alter the composition of the gut microbiome. In contrast, a diet high in fiber and plant-based foods has been shown to increase microbial diversity and promote a more diverse and healthy gut microbiome. In addition, lifestyle factors such as stress, exercise, and sleep have also been shown to influence the gut microbiome. For example, chronic stress has been shown to reduce microbial diversity and alter the composition of the gut microbiome.The gut microbiome has also been linked to a wide range of health outcomes, including obesity, diabetes, and inflammatory bowel disease. For example, studies have shown that obese individuals have a less diverse gut microbiome than lean individuals, and that the composition of the gut microbiome is altered in individuals with type 2 diabetes. In addition,the gut microbiome has been implicated in the development of inflammatory bowel disease, a chronic inflammatory condition of the gut.Given the importance of the gut microbiome for human health, there is growing interest in developing therapeutic interventions that target the gut microbiome. One approach is to use probiotics, which are live microorganisms that are intended to confer health benefits when consumed in adequate amounts. Probiotics have been shown to have a range of health benefits, including improving gut health and immune function. Another approach is to use prebiotics, which are non-digestible food ingredients that selectively stimulate the growth and activity of beneficial microorganisms in the gut. Prebiotics have been shown to increase microbial diversity and promote a more diverse and healthy gut microbiome.In conclusion, the gut microbiome is a complex ecosystem of microorganisms that plays a crucial role in human health. The diversity of the gut microbiome is important for resilience and metabolic function, and diet and lifestyle factors can influence the composition of the gut microbiome. The gut microbiome has been linked to a wide range of health outcomes, and there is growing interest in developing therapeutic interventions that target the gut microbiome. Probiotics and prebiotics are two approaches that show promise for promoting a healthy gut microbiome. Overall, a better understanding of the microbial ecology of the gut has the potential to improve human health and well-being.。
Ecology is the scientific study of the interactions of organisms and their environment.The biosphere is the part of Earth where life exists.All organisms live and interact in the biosphere.To understand relationships in biosphere, ecologists study events and organisms that range in complexity from a single individual to the entire biosphereBiotic factors that are living things that influence other organisms in an ecosystem.Abiotic factors are physical or nonliving factors that shape and ecosytem.Ecologists study many levels of organization.• individual organisms• species—a group of similar organisms that breed and produce fertile offspring• population—a group of individuals of the same species that live in the same area• community—a collection of different populations that live together in an area• ecosystem—all organisms living in a specific place, together with their physical environment• biome—a group of ecosystems with the same climate and similar dominant communities• biosphere—the part of the planet (land, water, and air) where all life exists.Scientists conduct ecological research using three basic approaches: \1.observing,2.experimenting,3.modeling.All of these approaches rely on the application of scientific methods to guide ecological inquiry.∙Observing is often the first step in asking ecological questions.∙Observations can also be used when designing experiments and making models.∙Experiments can be used to test hypotheses.∙They may be done in a laboratory or in the field.∙Modeling helps scientists understand complex processes.3–2 Energy FlowOrganisms use energy from the environment for life processes.Living things get energy in different ways.Sunlight is the main energy source for life on Earth.anisms that use the energy in sunlight or chemicals to make food are called autotrophs.Autotrophs, also called producers, make food in two ways.1.Some autotrophs use light energy to make food in a process called photosynthesis.o In photosynthesis, carbon dioxide and water are changed to carbohydrates and oxygen.o Plants, some algae, and certain bacteria carry out photosynthesis.o Some types of organisms rely on the energy stored in organic chemical compounds.2.Some autotrophs use chemical energy to make carbohydrates is called chemosynthesis.o Only certain types of bacteria carry out chemosynthesis.Organisms that rely on other organisms for energy and food are called heterotrophs.Heterotrophs also are called consumers.Types of heterotrophs.o Herbivores, such as cows, get energy by eating only plants.o Carnivores, such as snakes, get energy by eating only animals.o Omnivores, such as humans, get energy by eating both plants and animals.o Detritivores, such as earthworms, feed on the remains (dead matter) or wastes of other organisms.o Insectivores –such a anteaters, or some birds.o Scavengers an organism that feeds on dead or once living organismso Decomposers, such as fungi, break down organic matter.Energy flows through an ecosystem in one direction.It flows from the sun (or inorganic compounds) to autotrophs and then to heterotrophs.A food chain shows how living things transfer energy by eating and being eaten.For ex. a food chain might consist of grass (producer), an antelope (herbivore), and a coyote (carnivore).A food web links together all of the food chains in an ecosystem.For example, rabbits may also feed on the grass in the food chain above. These rabbits may be eaten by the coyotes. The feeding relationships of the grass, rabbits, antelopes, and coyotes make up a food web.Each step in a food chain or food web is called a trophic level.o Producers are at the first trophic levelo Consumers make up higher trophic levels.o Each consumer depends on the trophic level below it for energy.Ecological pyramids are diagrams that show the relative amounts of energy or matter at each trophic level.Only about 10 percent of the energy available at one trophic level is passed on to organisms at the next trophic level.Three types of ecological pyramids are1.Energy pyramids show how much energy is available at each trophic level.2.Biomass pyramidsshow the biomass, or total amount of living tissue, at each trophiclevel.3.Pyramid of numbers shows the relative number of individual organisms at each trophiclevel.3–3 Cycles of MatterEnergy and matter move through the biosphere very differently.Unlike the one-way flow of energy, matter is recycled within and between ecosystems. Matter, including water and nutrients, moves through organisms and parts of the biosphere through BIOCHEMICAL CYCLES.The Water Cycle.o All living things need water to survive.o Water cycles between the ocean, atmosphere, land, and living things.o Many processes part of the water cycle. Ex. during evaporation liquid water changes to gas.o Transpiration is the evaporation of water from the leaves of plants.o Water changes from a gas to a liquid through the process of condensation.o Water vapor in the atmosphere condenses into tiny droplets that form clouds.o When the droplets get large enough, they fall to Earth’s surface as precipitation.Nutrients are chemical substances that organisms need to survive.Lliving organism needs nutrients to build tissues and carry out essential life functions. Like water, nutrients are passed between organisms and the environment through biogeochemical cycles.The Carbon Cycle.o Carbon is a key part of living tissue.o Photosynthesis and cellular respiration are parts of the carbon cycle.o Human activities such as burning fossil fuels are also parts of the carbon cycle.The Nitrogen Cycle.o Organisms need nitrogen to build proteins.o Different forms of nitrogen cycle through the biosphere.o Nitrogen gas is the most abundant form of nitrogen on Earth.o However, only certain kinds of bacteria can use this form directly.o These bacteria change nitrogen gas into ammonia in a process called nitrogen fixation.o Other bacteria in the soil convert ammonia into nitrates and nitrites.o When organisms die, decomposers return nitrogen to the soil.o Other bacteria change nitrogen compounds called nitrates back into nitrogen gas.o This process is called denitrification.The Phosphorus Cycle.o Most phosphorus is stored in rocks and ocean sediments.o This phosphorus is slowly released into water and soil and then used by organisms.o Phosphorus is a key part of DNA and RNA.Primary productivityo It is the rate at which producers form organic matter in an ecosystem.o The availability of a nutrient affects primary productivity of a producer.o A nutrient that is scarce or cycles slowly through an ecosystem is a limiting nutrient.o A limiting nutrient can affect ecosystem health.。
Most of us are confused when it comes to the words ecosystem and biome. What's the difference? There is a slight difference between the two words. An ecosystem is much smaller than a biome. Conversely, a biome can be thought of many similar ecosystems throughout the world grouped together. An ecosystem can be as large as the Sahara Desert, or as small as a puddle or vernal pool.Ecosystems are dynamic interactions between plants, animals, and micro organisms and their environment working together as a functional unit. Ecosystems will fail if they do not remain in balance. No community can carry more organisms than its food, water, and shelter can accomodate. Food and territory are often balanced by natural phenomena such as fire, disease, and the number of predators. Each organism has its own niche, or role, to play.How have humans affected the ecosystems?We have affected ecosystems in almost every way imaginable! Every time we walk out in the wilderness or bulldoze land for a new parking lot we are drastically altering an ecosystem. We have disrupted the food chain, the carbon cycle, the nitrogen cycle, and the water cycle. Mining minerals also takes its toll on an ecosystem. We need to do our best to not interfere in these ecosystems and let nature take its toll.EcosystemFrom Wikipedia, the free encyclopediaJump to: navigation, searchThe High Peaks Wilderness Area in the 6,000,000-acre (2,400,000 ha) Adirondack Park is an example of a diverse ecosystem.An ecosystem is a system of interdependent organisms which share the same habitat, in an area functioning together with all of the physical (abiotic) factors of the environment. Ecosystems can be permanent or temporary. Ecosystems usually form a number of food webs. [1]Contents[hide]∙1Overview∙2Examples of ecosystems∙3Biomes∙4Ecosystem topicso 4.1Classificationo 4.2Ecosystem serviceso 4.3Ecosystem legal rightso 4.4Function and biodiversity∙5The study of ecosystemso 5.1Ecosystem dynamicso 5.2Ecosystem ecology∙6See also∙7References∙8Further reading9External links[edit] OverviewFlora of Baja California Desert, Cataviña region, Mexico.Savanna at Ngorongoro Conservation Area, Tanzania.The term ecosystem was coined in 1930 by Roy Clapham to mean the combined physical and biological components of an environment. British ecologist Arthur Tansley later refined the term, describing it as "The whole system,… including not only the organism-complex, but also the whole complex of physical factors forming what we call the environment".[2] Tansley regarded ecosystems not simply as natural units, but as mental isolates.[2]Tansley later[3]defined the spatial extent of ecosystems using the term ecotope.Central to the ecosystem concept is the idea that living organisms interact with every other element in their local environment. Eugene Odum, a founder of ecology, stated: "Any unit that includes all of the organisms (ie: the "community") in a given area interacting with the physical environment so that a flow of energy leads to clearly defined trophic structure, biotic diversity, and material cycles (ie: exchange of materials between living and nonliving parts) within the system is an ecosystem."[4] The human ecosystem concept is then grounded in the deconstruction of the human/nature dichotomy and the premise that allspecies are ecologically integrated with each other, as well as with the abiotic constituents of their biotope.[edit] Examples of ecosystems∙Agroecosystem∙Aquatic ecosystem ∙Chaparral∙Coral reef∙Desert∙Forest∙Greater Yellowstone Ecosystem ∙Human ecosystem∙Large marineecosystem∙Littoral zone∙Marine ecosystem∙Prairie∙Rainforest∙Savanna∙SubsurfaceLithoautotrophic MicrobialEcosystem∙Taiga∙Terrestrial Ecosystem∙Tundra∙Urban ecosystemA freshwater ecosystem in Gran Canaria, an island of the Canary Islands.[edit] BiomesMap of Terrestrial biomes classified by vegetation.Main article: BiomeBiomes are similar to ecosystems, and are climatically and geographically defined area of ecologically similar climatic conditions such as communities of plants, animals, and soil organisms, often referred to as ecosystems. Biomes are defined based on factors such as plant structures(such as trees, shrubs, and grasses), leaf types (such as broadleaf and needleleaf), plant spacing (forest, woodland, savanna), and climate. Unlike ecozones, biomes are not defined by genetic, taxonomic, or historical similarities. Biomes are often identified with particular patterns of ecological succession and climax vegetation.[edit] Ecosystem topics[edit] ClassificationThe Daintree Rainforest in Queensland, Australia.Ecosystems have become particularly important politically, since the Convention on Biological Diversity (CBD) - ratified by more than 175 countries - defines "the protection of ecosystems, natural habitats and the maintenance of viable populations of species in natural surroundings"[5]as a commitment of ratifying countries. This has created the political necessity to spatially identify ecosystems and somehow distinguish among them. The CBD defines an "ecosystem" as a "dynamic complex of plant, animal and micro-organism communities and theirnon-living environment interacting as a functional unit".With the need of protecting ecosystems, the political need arose to describe and identify them efficiently. Vreugdenhil et al. argued that this could be achieved most effectively by using aphysiognomic-ecological classification system, as ecosystems are easily recognizable in the field as well as on satellite images. They argued that the structure and seasonality of the associated vegetation, complemented with ecological data (such as elevation, humidity, and drainage), are each determining modifiers that separate partially distinct sets of species.This is true not only for plant species, but also for species of animals, fungi and bacteria. The degree of ecosystem distinction is subject to the physiognomic modifiers that can be identified on an image and/or in the field. Where necessary, specific fauna elements can be added, such as seasonal concentrations of animals and the distribution of coral reefs.Several physiognomic-ecological classification systems are available:∙Physiognomic-Ecological Classification of Plant Formations of the Earth: a system based on the 1974 work of Mueller-Dombois and Heinz Ellenberg[6], and developed byUNESCO. This classificatie "describes the above-ground or underwater vegetationstructures and cover as observed in the field, described as plant life forms. Thisclassification is fundamentally a species-independent physiognomic, hierarchicalvegetation classification system which also takes into account ecological factors such asclimate, elevation, human influences such as grazing, hydric regimes, and survivalstrategies such as seasonality. The system was expanded with a basic classification foropen water formations".[7]∙Land Cover Classification System (LCCS), developed by the Food and Agriculture Organization (FAO).[8]∙Forest-Range Environmental Study Ecosystems (FRES) developed by the United States Forest Service for use in the United States.[9]Several aquatic classification systems are available, and an effort is being made by the United States Geological Survey (USGS) and theInter-American Biodiversity Information Network (IABIN) to design a complete ecosystem classification system that will cover both terrestrial and aquatic ecosystems.From a philosophy of science perspective, ecosystems are not discrete units of nature that simply can be identified using "the right" classification approach. In agreement with the definition by Tansley ("mental isolates"), any attempt to delineate or classify ecosystems should be explicit about the observer/analyst input in the classification including its normative rationale.Summer field in Belgium(Hamois). The blue flower is Centaurea cyanus and the red one a Papaver rhoeas.[edit] Ecosystem servicesMain article: Ecosystem servicesEcosystem services are “fundamental life-support services upon which human civilization depends,”i and can be direct or indirect. Examples of direct ecosystem services are: pollination, wood, and erosion prevention. Indirect services could be considered climate moderation, nutrient cycles, and detoxifying natural substances.[edit] Ecosystem legal rightsEcuador's new constitution of 2008 is the first in the world to recognize legally enforceable Rights of Nature, or ecosystem rights.[10]The borough of Tamaqua, Pennsylvania passed a law giving ecosystems legal rights. The ordinance establishes that the municipal government or any Tamaqua resident can file a lawsuit on behalf of the local ecosystem.[11] Other townships, such as Rush, followed suit and passed their own laws.[12]This is part of a growing body of legal opinion proposing 'wild law'. Wild law, a term coined by Cormac Cullinan (a lawyer based in South Africa), would cover birds and animals, rivers and deserts.[13][14][edit] Function and biodiversitySee also: BiodiversityFrom an anthropological point of view, many people see ecosystems as production units similar to those that produce goods and services. Among some of the most common goods produced by ecosystems, is wood by forest ecosystems and grass for cattle by natural grasslands. Meat from wild animals, often referred to as bush meat in Africa, has proven to be extremely successful under well-controlled management schemes in South Africa and Kenya. Much less successful has been the discovery and commercialization of substances of wild organism for pharmaceutical purposes. Services derived from ecosystems are referred to as ecosystem services. They may include (1) facilitating the enjoyment of nature, which may generate many forms of income and employment in the tourism sector, often referred to as eco-tourisms, (2) water retention, thus facilitating a more evenly distributed release of water, (3) soil protection, open-air laboratory for scientific research, etc.A greater degree of species or biological diversity - popularly referred to as Biodiversity- of an ecosystem may contribute to greater resilience of an ecosystem, because there are more species present at a location to respond to change and thus "absorb" or reduce its effects. This reduces the effect before the ecosystem's structure is fundamentally changed to a different state. This is not universally the case and there is no proven relationship between the species diversity of an ecosystem and its ability to provide goods and services on a sustainable level: Humid tropical forests produce very few goods and direct services and are extremely vulnerable to change, while many temperate forests readily grow back to their previous state of development within a lifetime after felling or a forest fire. Some grasslands have been sustainably exploited for thousands of years (Mongolia, Africa, European peat and mooreland communities).[edit] The study of ecosystemsThe Blue Marble, taken in 1972 by the Apollo 17 crew. This image is the only photograph of its kind to date, showing a fully sunlit hemisphere of the Earth. Approximately 72% of the Earth's surface (an area of some 361 million square kilometers) consists of ocean.[edit] Ecosystem dynamicsSpiny forest at Ifaty, Madagascar, featuring various Adansonia(baobab) species, Alluaudia procera (Madagascar ocotillo) and other vegetation.Introduction of new elements, whether biotic or abiotic, into an ecosystem tend to have a disruptive effect. In some cases, this can lead to ecological collapse or "trophic cascading" and the death of many species within the ecosystem. Under this deterministic vision, the abstract notion of ecological health attempts to measure the robustness and recovery capacity for an ecosystem; i.e. how far the ecosystem is away from its steady state.Often, however, ecosystems have the ability to rebound from a disruptive agent. The difference between collapse or a gentle rebound is determined by two factors—the toxicity of the introduced element and the resiliency of the original ecosystem.Ecosystems are primarily governed by stochastic (chance) events, the reactions these events provoke on non-living materials, and the responsesby organisms to the conditions surrounding them. Thus, an ecosystem results from the sum of individual responses of organisms to stimuli from elements in the environment. The presence or absence of populations merely depends on reproductive and dispersal success, and population levels fluctuate in response to stochastic events. As the number of species in an ecosystem is higher, the number of stimuli is also higher. Since the beginning of life organisms have survived continuous change through natural selection of successful feeding, reproductive and dispersal behavior. Through natural selection the planet's species have continuously adapted to change through variation in their biological composition and distribution. Mathematically it can be demonstrated that greater numbers of different interacting factors tend to dampen fluctuations in each of the individual factors.Arctic tundra on Wrangel Island, Russia.Forest on San Juan IslandGiven the great diversity among organisms on earth, most ecosystems only changed very gradually, as some species would disappear while others would move in. Locally, sub-populations continuously go extinct, to be replaced later through dispersal of other sub-populations. Stochastists do recognize that certain intrinsic regulating mechanisms occur in nature. Feedback and response mechanisms at the species level regulate population levels, most notably through territorial behaviour. Andrewatha and Birch[15] suggest that territorial behaviour tends to keep populations at levels where food supply is not a limiting factor. Hence, stochastists see territorial behaviour as a regulatory mechanism at the species level but not at the ecosystem level. Thus, in their vision, ecosystems are notregulated by feedback and response mechanisms from the (eco)system itself and there is no such thing as a balance of nature.If ecosystems are indeed governed primarily by stochastic processes, they may be more resilient to sudden change than each species individually. In the absence of a balance of nature, the species composition of ecosystems would undergo shifts that would depend on the nature of the change, but entire ecological collapse would probably be infrequent events.The theoretical ecologist Robert Ulanowicz has used information theory tools to describe the structure of ecosystems, emphasizing mutual information (correlations) in studied systems. Drawing on this methodology and prior observations of complex ecosystems, Ulanowicz depicts approaches to determining the stress levels on ecosystems and predicting system reactions to defined types of alteration in their settings (such as increased or reduced energy flow, and eutrophication.[16]See also: Relational order theories, as to fundamentals of life organization[edit] Ecosystem ecologyEcosystem ecology is the integrated study of biotic and abiotic components of ecosystems and their interactions within an ecosystem framework. This science examines how ecosystems work and relates this to their components such as chemicals, bedrock, soil, plants, and animals. Ecosystem ecology examines physical and biological structure and examines how these ecosystem characteristics interact.。
高三生物生态系统英语阅读理解25题1<背景文章>The tropical rainforest ecosystem is one of the most diverse and fascinating ecosystems on Earth. It is characterized by its lush vegetation, high humidity, and abundant rainfall. The rainforest is home to a wide variety of plants and animals, many of which are found nowhere else in the world.The biodiversity of the tropical rainforest is truly remarkable. There are thousands of different species of plants, including towering trees, delicate orchids, and colorful bromeliads. The animal life is equally diverse, with monkeys, sloths, toucans, and countless other species calling the rainforest home.However, the tropical rainforest is facing many threats. Deforestation is one of the most serious problems, as large areas of rainforest are being cleared for agriculture, logging, and urban development. This not only destroys the habitat of countless species but also contributes to climate change.To protect the tropical rainforest, various measures are being taken. Conservation organizations are working to raise awareness about the importance of the rainforest and to promote sustainable development.Governments are also implementing policies to limit deforestation and to protect the remaining rainforests.1. The tropical rainforest ecosystem is characterized by ___.A. low humidity and little rainfallB. sparse vegetation and dry climateC. lush vegetation, high humidity, and abundant rainfallD. cold temperature and few plants答案:C。
Unit 3Text A The Case Against Man(状告人类)Teaching Objectives(教学目的)Students will be able to1. grasp the main idea and structure of the text2. master the key language points and grammatical structures3. conduct a series of reading, listening, speaking and writing activities related to the theme of the unit.Teaching procedures(教学步骤)(1) Pre-reading activity (Period1)a. Information related to the textb. Introduction to the Textc. Pre-reading questions(2)While-reading activity(Periods2-3)a. The organization of the textb. Detailed reading---study of the language points(3) Post-reading activity (Period4)a. Speaking Activityb. Writing Practicec. Have students finish the exercisesPeriod 1Pre-reading activitya. Background Information1. Introduction to the AuthorIsaac Asimov (1920-1992) was a Russian-born American writer and scientist. Though he published several serious scientific works, he is best known for his science fiction stories. His prolific output contains some of the finest novels in science fiction, including Robot (1950) and The Foundation Trilogy (1951-1953).2. Introduction to the TextEcology is the scientific study of the relationship of plants and animals to their physical and biological environment. The physical environment covers factors such as the Earth‟s land, sea, light, temperature, atmosphere etc. The biological environment refers to living organisms (also known as biotic organisms) such as plants, microorganisms, and animals. The ecology of a particular place is the balance of relation between plants, animals, people, and the environment there.Ecosystem refers to the interacting community of organisms and their pysical environment. It includes all organic life in a given area along with the soil, water, and other inorganic components of their habitat, and all the ecological interactions that takeplace within and between the organic and inorganic. It is a complete ecosystem only if it can incorporate energy into organic compounds and pass it from organism to organism, and if it recycles elements for re-use. What will happen to the ecosystem if we allow the present rate of population increase to continue? Can we count on science and technology to develop methods for finding food and materials for the increasing number of population? What will be the effect on the Earth‟s environment, and eventually on the human race itself? These are the questions the present article “The Case Against Man”discusses. After reading the article, the reader will realize what exactly the case against man is.3. Pre-reading questionsLead-in questionsPurposes:·To present key words and phrases related to environment and ecology·To encourage students to think about the theme of the coming text, i.e.the rapidly increasing human population is destroying the environment·To arouse students‟ interest in the topic of the coming text·To develop students‟ skill in predicting the main idea of the textMethodology:1. The purpose of activities 1, 2 and 3 is to arouse students’interests in the topic ofthe text and prepare them to read the text actively. It may be very easy for your students to give a list of environmental problems and the harmful effects of population explosion on the ecological system and human life as a whole, but to find the way out is more difficult. If you encourage students to think carefully about the solution to the population explosion, they will probably hit upon the central idea and the main purpose of the coming article.2.In order to get the whole class involved, pair-work is recommended for activity 1, 2 and 3. If time permits, you can find on or two students to report to the class what they have discussed.3. In activity 4, some key words and phrases along with their definitions arepresented.Although many students are already familiar with these words and phrases, they cannot always give accurate definitions of them in English. To make sure that the whole class participate in the discussion, let them work in pairs first and then encourage them to compare their answers with other neighboring classmates. By working with a partner first and then more classmates, students should be able to match the terms and definitions correctly. There is no need to check the answers in class, but be ready to provide any help they need while they are working.Key to lead-in questions1.The environment is all the natural conditions, such as air, water, and land in whichman lives. Environmental problems have recently become bad to worse in the world scope.Y es. For example, with the development of industry, the pollution of air and wateris found everywhere. Forests and vegetation have been destoryed, thus causing a large number of environmental problems, ranging from global warming to ozone depletion. What‟s the worse, thousands of species are becoming extinct every year, while human population is increasing at an unprecedented rate…2.Ecology is a science that studies the intricate interrelated and interdependentsystem of the Earth plus solar-energy.The ecological system in which we live is of great significance to mankind. If one species was removed from the system, there would occur the knock-on effects on the ecosystem. If we lost a key species, we might cause a whole flood of other extinctions. So it is up to everyone in the world to preserve the ecosystem.3.At the present rate of growth, the population explosion will do great harm to theEarth‟s environment. Every body born means an additional burden to the Earth‟s supporting capability, a challenge to other species on the planet. In my opinion, the best way out is to have birth controlling, so as to keep the world population withina limited number.4.1) c 2) l 3) d 4) i 5) b 6) k 7) h 8) e 9) a 10) g 11)j 12) fPeriod 2-3While-reading activitya. The organization of the textThe Gist of the ArticleThe author of this article mainly argues that if the number of population continues to grow at the present rate, without birth control, even if science serves us in an absolutely ideal way, we will reach the planetary high-rise with no animals but man, with no plants but algae, with no room for even one more person by the year 2430. If science serves us in less than an ideal way, the end will come sooner, and mankind will start fading long before he is forced to construct buildings that will cover all the Earth‟s surface. In the end, man, like murderer, will destroy life and the whole planet. For this, he will have to stand at the bar of justice. Therefore, the author concludes, if life is to be saved, birth control should come right now, when there are still oak trees in the world and daisies and tigers and butterflies, and when there is still open land and space.In order to convince the reader of the serious situation humankind is facing, the author in the first seven paragraphs tries to define the nature of mank ind, life, the planet, and their relationship. Mankind is not an independent existence. It is part of an intricate web of life. Nor is life an independent existence. It is part of the intricate structure of the Earth. Life takes many complex forms. Each of these forms is an inseparable part of a whole. All life forms are interrelated to each other. The entire planet, plus solar energy, is one enormous intricately interrelated system. It is made up of a variety of nonliving portions and living portions.Form paragraph 8 to paragraph 18, the author uses the analogy of human body, which consists of 50 trillion cells of many different types, to explain the interrelationship between different life forms on the planet. The structure of the planet is similar to that of the human body in many ways. For example, all the cells of humanbody are interrelated and interdependent. The loss of one cell will seriously handicap the rest of the organism. In the same way, the loss of one animal or a forest will seriously change the life forms and geological structure of a region. On the other hand, if a particular group of cell grows and multiplies without limit, it will become the fatal disease cancer, which will kill the whole human organism completely. Similarly, if one type of organism on the planet grows and multiplies without limit, killing its competitors, it will threaten the existence of a larger system, and eventually destroy the organism itself.Unfortunately, this is what is happening at present. The number of human population is growing at a cancerous rate, which is threatening the balance of ecosystem. This will eventually destroy the environment and humankind itself.Therefore, it is of utmost importance now to stop the population from growing too fast. Since we cannot increase death rate, we have to reduce birthrate, and reduce it right now.b. Detailed reading---study of the language pointsDetailed reading of the textPurposes:·To help students draw important facts of each paragraph ‟·To help students identify and remove the source of difficulty in understanding the text·To help students actively interact with the text and develop strategies to work out the meaning of the text·To give students practice in note-takingMethodology:When working through the text, you can, besides explanation (refer to “Detailed E xplanation of the Text‟‟ in the student’s book), use the technique of asking questions to develop students‟understanding. Make sure students always keep their text open when they answer the questions. The purpose of asking questions is to make students actively interact with the text and develop their strategies to derive the meaning from the text. At the same time, they can become aware of the way language is used to convey meaning. There is a close connection between answering questions and developing understanding.At the start, three types of questions should be designed for this purpose. First, straightforward questions ne ed to be asked to draw students‟attention to important facts in the text and to ensure they understand the literal meaning of the facts.The second type of questions should be used to develo p students‟ skills of tackling difficult sentences.A third type should make students recognize grammatical and lexical features that play a crucial part in the interpretation of the text.…When students answer your questions they should also practice taking notes.Y ou can ask the following questions orally.1. How were ocean, atmosphere, and solid crust formed in the process of evolution? In what way did they interact with each other? What were the noticeable results of theirinteraction?2. What happened to the portion of water between 2 billion and 3 billion years ago?3. In what sense is life an inseparable part of the strcture of the earth?4. What analogy is used to explain that the entire planet, plus solar energy, is oneenormous intricately interrelated sysyem?5. In what sense are the 50 trillion cells in a human body interrelated andinterdependent ?6. What is the author‟s purpose of mentioning the chopping down of an entire forestand a serious decline in the bee population in paragrapg 9?7. How many types of cells are grouped in paragraph 10? What point is the authormaking by distinguishing these types?8. What is the author‟s purpose of explaining the relationship between cougars anddeer? What is the importance for an animal type to maintain the proper number? 9. What will happen to human body if a group of cells multiplies without limit? Whatwill happen to the ecological system if one particular type of organism multiplies without limit? What does this particular type of organism refer to according to paragraph 14?10. For what purpose does the author compare the increase rate of population inCaesar‟s time and the present increase rate?11. What figure of speech does the author use to describe the present rate of populationincrease?12. Does the author suggest a cure for the cancerous growth of population? If yes, whatis the solution he suggests?13. With what crime would mankind be charged if birthrate was not effectivelycontrolled?14. What is that “Case Against Man”suggested in the title?Period 4Post-reading activitya. Speaking ActivityAsk Ss to discuss the “effect of population explosion on the earth‟s environment”in groups and then present their opinions in class.b. Writing PracticeWith the help of writing strategies on P76, Ss write an essay based upon one of the topics on page 81.c. Have students finish the exercises。
考研英语二练习题考研英语二阅读理解练习题Passage 1: The Impact of Technology on EducationIn recent years, the rapid advancement of technology has dramatically transformed the educational landscape. This passage will discuss how the integration of digital tools and online platforms has reshaped the way students learn and teachers teach.Questions:1. What has been the primary driver of changes in education?2. How have digital tools affected the learning process?3. In what ways have online platforms influenced teaching methods?Passage 2: The Benefits of a Balanced DietA balanced diet is essential for maintaining good health and well-being. This article explores the nutritional components necessary for a balanced diet and the benefits they provideto the body.Questions:1. Why is a balanced diet important for health?2. What are the key nutritional components of a balanced diet?3. What benefits can be derived from consuming a balanceddiet?Passage 3: The Role of Volunteering in Community DevelopmentVolunteering plays a crucial role in the development of communities. This passage examines the various ways in which volunteers contribute to the betterment of society and the impact of their efforts.Questions:1. What is the significance of volunteering in community development?2. How do volunteers contribute to societal improvement?3. What are the effects of volunteering on both thevolunteers and the community?Passage 4: The Effects of Climate Change on BiodiversityClimate change poses a significant threat to biodiversity around the world. This article delves into the effects of climate change on various ecosystems and the species that inhabit them.Questions:1. What is the relationship between climate change and biodiversity?2. How do ecosystems respond to the impacts of climate change?3. What are the consequences for species as a result ofclimate change?Passage 5: The Evolution of Urban PlanningUrban planning has evolved significantly over time in response to changing societal needs and technological advancements. This passage outlines the historical development of urban planning and its current trends.Questions:1. How has urban planning evolved over time?2. What factors have influenced the development of urban planning?3. What are the current trends in urban planning and their implications for future cities?Passage 6: The Importance of Sleep for Cognitive FunctionSleep is vital for cognitive function, affecting memory, learning, and decision-making. This article discusses the importance of sleep and the consequences of sleep deprivation on cognitive abilities.Questions:1. Why is sleep important for cognitive function?2. What are the effects of sleep deprivation on cognitive abilities?3. How can individuals improve their sleep quality to enhance cognitive performance?Passage 7: The Challenges of Sustainable DevelopmentSustainable development aims to meet the needs of the present without compromising the ability of future generations tomeet their own needs. This passage addresses the challenges faced in achieving sustainable development and potential solutions.Questions:1. What are the main challenges in achieving sustainable development?2. How can these challenges be overcome to ensure a sustainable future?3. What role do individuals, communities, and governments play in sustainable development?。
Stimulation of Different Functional Groups of Bacteria by Various Plant Residues as a Driver of Soil PrimingEffectNoe´mie Pascault,1Lionel Ranjard,1,2Aurore Kaisermann,1Dipankar Bachar,3Richard Christen,3Se´bastien Terrat,1,2Olivier Mathieu,4Jean Le´ve ˆque,4Christophe Mougel,1Catherine Henault,1Philippe Lemanceau,1Michel Pe´an,5Se ´verine Boiry,5Se ´bastien Fontaine,6and Pierre-Alain Maron 1,2*1UMR Agroecology,INRA/Universite´de Bourgogne,CMSE,BP 86510,17rue de Sully,21065Dijon Cedex,France;2INRA,Plateforme GenoSol,UMR1347Agroe´cologie,BP 86510,21000Dijon,France;3Laboratoire de Biologie Virtuelle,Centre de Biochimie,Parc Valose,Universite´de Nice et CNRS UMR 6543,06108Nice,France;4UMR 5561Bioge ´osciences,CNRS/Universite ´de Bourgogne,Universite´de Bourgogne,6bd Gabriel,21000Dijon,France;5CEA,DSV,IBEB,Group Rech Appl Phytotechol,UMR 6191CEA/CNRS/Universite´Aix-Marseille,13108Saint-Paul-lez-Durance,France;6UR874Unite ´de Recherche sur l’Ecosyste `me Prairial,INRA,234avenue du Bre´zet,63100Clermont-Ferrand,France A BSTRACTThe turnover of organic matter in soil depends onthe activity of microbial decomposers.However,little is known about how modifications of the diversity of soil microbial communities induced by fresh organic matter (FOM)inputs can regulate carbon cycling.Here,we investigated the decom-position of two 13C labeled crop residues (wheat and alfalfa)and the dynamics of the genetic structure and taxonomic composition of the soilbacterial communities decomposing 13C labeled FOM and native unlabeled soil organic matter (SOM),respectively.It was achieved by combining the stable isotope probing method with molecular tools (DNA genotyping and pyrosequencing of 16S rDNA).Although a priming effect (PE)was always induced by residue addition,its intensity increased with the degradability of the plant residue.The input of both wheat and alfalfa residues induced a rapid dynamics of FOM-degrading communities,corresponding to the stimulation of bacterial phyla which have been previously described as copio-trophic organisms.However,the dynamics and the identity of the bacterial groups stimulated depended on the residue added,with Firmicutes dominating in the wheat treatment and Proteobac-teria dominating in the alfalfa treatment after 3days of incubation.In both treatments,SOM-degrading communities were dominated by Acido-bacteria ,Verrucomicrobia ,and Gemmatimonadetes phyla which have been previously described as oligotrophic organisms.An early stimulation of SOM-degrading populations mainly belonging toReceived 26September 2012;accepted 20December 2012;published online 22February 2013Electronic supplementary material:The online version of this article (doi:10.1007/s10021-013-9650-7)contains supplementary material,which is available to authorized users.Author Contributions:NP performed the research;NP and AK jointly conducted the incubations;DB,RC,and CM performed bioinformatic analysis;OM and JL performed 13C analysis;CH performed PLFA analysis;MP labeled the plants;PL and SF commented on the manuscript,the modeling and the statistical analyses;PAM and LR designed the study;PAM wrote the first version of the manuscript.All the authors contrib-uted substantially to revisions.*Corresponding author;e-mail:pamaron@dijon.inra.frEcosystems (2013)16:810–822DOI:10.1007/s10021-013-9650-7Ó2013Springer Science+Business Media New York810Firmicutes and Bacteroidetes groups was observed in the alfalfa treatment whereas no change occurred in the wheat treatment.Ourfindings support the hypothesis that the succession of bacterial taxo-nomic groups occurring in SOM-and FOM-degrading communities during the degradation process may be an important driver of the PE,and consequently of carbon dynamics in soil.Key words:bacterial diversity;soil;stable isotope probing;pyrosequencing;carbon cycle;soil organic matter.I NTRODUCTIONSoil organic matter(SOM)is a major component controlling the ability of soil to deliver many ser-vices on which human well-being is dependent. Most SOM transformations rely on the activity of soil decomposers(mainly soil microorganisms) which simultaneously determine(i)the recycling of nutrients essential to the dynamics of agro-ecosystems,(ii)the storage of SOM necessary for the maintenance of soil structure and fertility(Le Guillou and others2012),and(iii)the release of greenhouse gases(Schlesinger and Andrews2000). Despite the importance of the microbial compo-nent,its qualitative integration into carbon cycle models is still hampered by a knowledge gap regarding the diversity of the populations involved and the functional significance of microbial diver-sity in SOM turnover(Ingwersen and others2008). In most of these models,and on the basis of Bei-jerinck’s postulate(1913)that‘‘everything is everywhere’’,soil microorganisms are considered as ubiquitous organisms harboring a high functional redundancy.Thus,indigenous microbial commu-nities are still regarded as a functional‘‘black box’’generatingfluxes of different intensities which are solely dependent on abiotic factors such as tem-perature,moisture,and pH.The hypothesis that the diversity and composition of microbial communi-ties,as well as trophic interactions between popu-lations,may play a functional role is therefore excluded(McGill1996;Gignoux and others2001). New models have recently been proposed that evaluate the interest of using more qualitative parameters(that is,taxonomic or functional diversity)than global microbial biomass to consider the soil microbial component(Schimel and Wein-traub2003;Fontaine and others2003;Fontaine and Barot2005;Neill and Gignoux2006;Lawrence and others2009).This changing of the conceptual structure of the models has shown particular promise for predicting the priming effect(PE), which is the stimulation of the decomposition of native SOM following the supply of energy-rich fresh organic matter(FOM)to soil(Fontaine and others2007;Kuzyakov2010).It suggests that thePE may depend on interactions between two microbial functional groups:the copiotrophs and the oligotrophs,which may correspond to the populations decomposing FOM and native SOM, respectively(Fontaine and others2003;Fontaine and Barot2005;Neill and Gignoux2006).How-ever,these models are mainly conceptual and need to be tested by experimental observations.In this context,experiments involving simultaneous assessment of the PE and of the diversity of the microbial communities involved in FOM and SOM degradation,respectively,would be of particular interest.The recent development of stable isotope probing (SIP)provides an opportunity for such studies be-cause it allows specific tracking of the microbial groups/populations actively involved in the decomposition of C-substrates labeled with stable isotopes(for example,13C)(Radajewski and others 2000;Chen and Murrell2010).It has been suc-cessfully used to identify soil microorganisms involved in the degradation of labeled lignin,ace-tate,glucose,and cellulose(Padmanabhan and others2003;Haichar and others2007);plant root exudates(Haichar and others2008);or labeled plant residues(Malosso and others2004;Bernard and others2007,2009).These results showed overall that the biochemical quality of the C-substrate added strongly influenced the diversity of the FOM-and SOM-degrading communities.This then raised the question as to whether these modifications of soil microbial diversity could explain the induction of different PE.In this work,we hypothesized that the dynamics of the structure and composition of the FOM-and SOM-degrading communities depend on the qual-ity of FOM added,and that differences in the stimulated communities result in differences in PE intensity.A microcosm experiment was set up to test this hypothesis.Two13C labeled crop residues (wheat and alfalfa)with contrasting biochemical compositions were incorporated into soil.The subsequent kinetics of FOM and SOM mineraliza-tion,and the dynamics of FOM-and SOM-degrading communities,were monitored. Linking Microbial Diversity to Priming Effect811M ATERIALS AND M ETHODSPlant Culture and13C-labelingSeeds of wheat(Triticum aestivum cv Caphorn)were germinated at4°C in darkness and seeds of alfalfa (Medicago truncatula ec.J5)were germinated on 0.7%(w/v)water agar plates at25°C in darkness. After germination,each plantlet was grown in a mix of sand(1/4)and perlite(3/4)in an airtight chamber which allowed accurate regulation of atmospheric gas composition and environmental parameters(Groupe de Recherches Applique´es en Phytotechnologie,CEA Cadarache,France).The day–night period was set at8/16and14/10for the wheat and alfalfa plants,respectively,the photon flux rate was350l mol m-2s-1for wheat and 300l mol m-2s-1for alfalfa,and the relative humidity was65%day/80%night and70%day/ 80%night for wheat and alfalfa,respectively.Both plant cultures were watered with half-diluted Hoagland’s nutrient solution(wheat plants were watered6times a day and alfalfa plants were wa-tered13times a day to avoid water stress)and CO2 concentration was maintained at380l l l-1.The partial pressures of13CO2and12CO2in the cham-ber were both continuously monitored by near infrared spectroscopy to determine13C enrichment of the CO2.Regulation was achieved by automatic injection of pure(>99%atom13C)13CO2(pur-chased from CortecNet,Paris,France).The isotope excess in the chamber wasfixed at greater than 80%atom13C during thefirst10days and greater than90%atom13C thereafter.After sowing,wheat and alfalfa were grown for196and80days, respectively,to obtain mature plants.The roots were separated from the shoots which were then oven-dried at65°C for48h before further use. Wheat was labeled with13C at96%and charac-terized by a C/N ratio of78.2(determined using an elementar analyser Euro EA(EUROVECTOR, Milano,Italia)),and alfalfa was labeled with13C at 89%and characterized by a C/N ratio of7.5. Soil,Microcosms,and Sampling Strategy The soil used was a calcareous silty-clay from the experimental farm of INRA Epoisse(47.14°N,5.06°E,Burgundy,France).The soil had a pH H2Oof6.7and contained37.9%clay(<2l m);32.5%silt (2–20l m);23.7%silt(20–50l m);3.8%sand(50–200l m);2.1%sand(200–2000l m);CaCO3less than1g kg-1;1.47%organic C and0.15%total N. The C/N ratio was11.1.The13C content of the soil was1.078%.The soil was collected from the sur-face layer(0–5cm)and sieved to4mm.Microcosms were set up by placing30g of dry soil in150-ml hermetically sealed plasmaflasks. Sterile water was added to the microcosms to attain 60%of the maximum water holding capacity.After a pre-incubation of1week at20°C,the microcosms were either amended with13C-residues of wheat or alfalfa(5mg g-1of dry weight soil)which were finely ground in a bead beater to obtain a powder, or not amended(control).This represented a total of63microcosms(3treatments,7sampling dates,3 replicates per treatment and sampling date).The soil microcosms were then incubated at20°C in the dark for120days.After3,7,10,14,21,28,44,60, 75,90,105,and120days of incubation,micro-cosms were aerated byflushing with air and soil water content was verified.For each treatment, CO2measurements(total and13C-CO2)were per-formed on triplicate microcosms just before the aeration of the microcosms and biological analyses were performed on triplicate microcosms just after application of the treatments(0)and after3,7,14, 28,60,and120days of incubation.At each sampling point,triplicate microcosms from every treatment(enriched with residue and control microcosms)were sacrificed and soil was stored at-80°C in50ml Falcon tubes before DNA extraction.Total and13C-CO2MeasurementsThe gaseous phases of the microcosms were sam-pled at3,7,10,14,21,28,44,60,75,90,105,and 120days of incubation in3ml Terumo evacuated blood collection tubes for measurement of the CO2 concentration and in37ml airtightflasks for determination of carbon isotopic abundances. Because the microcosms were hermetically sealed and aerated byflushing with air at each sampling date,the concentrations measured corresponded to the CO2accumulated in the microcosm headspace between two sampling dates.CO2concentration was determined on a MT1Analytical Instrument micro GC equipped with a pora-PLOT Q capillary column and a Solid State Detector.Carbon isotopic enrichment was determined by gas chromatogra-phy isotope ratio mass spectrometry(GC-IRMS) using a trace gas interface coupled in continuous flow with a mass spectrometer VG Isochrom (Micromass,Manchester,England).For isotopic CO2,a pure bottle(CO2>99.999%,N48,Air Liquide,France),previously calibrated against a certified isotopic standard(d13C=-25.5±0.2& vs.PDB,ISO-TOP,Air Liquide),was selected as internal standard.Gas samples were manually injected into the trace gas with an air gas syringe.812N.Pascault and others13C labeling of the plant residues allowed separa-tion of soil C(R s)and plant residue(R r)respiration (mg C-CO2kg-1soil)using mass balance equations:R sþR r¼R tR sÂA s13þR rÂA r13¼R tÂA t13where A s13is the13C abundance(dimensionless)of soil carbon,A r13the13C abundance of plant resi-due,R t the total CO2emitted by soil with plant residue and A t13its13C abundance.The priming effect(PE,mg C-CO2kg-1soil)induced by the addition of plant residue was calculated as:PE¼R s soil with plant residueðÞÀðR s control soilÞwhere(R s control soil)is the CO2emitted by con-trol soil.DNA Extraction from SoilFor each treatment,DNA was extracted from triplicate microcosms from every treatment(enriched with residue and control microcosms)just after application of the treatments(0)and after3,7,14,28,60,and 120days of incubation.DNA was extracted from 391g of dry soil per microcosm following the DNA extraction procedure of Bernard and others(2007). The threefiltrates resulting from the391g nucleic acids extraction for each microcosm were pooled.Total DNA concentrations in each sample werefluoromet-rically quantified with the Quant-iT PicoGreenÒdsDNA Assay Kit(Invitrogen,Cergy-Pontoise,France) following the manufacturer’s instructions. Isopycnic Centrifugation and FractionationAn isopycnic density gradient centrifugation of all DNA extracts was performed to obtain a heavy DNA fraction containing13C-DNA and a light DNA fraction con-taining12C-DNA,as previously described(Bernard and others2007).The DNA concentration of each fraction of the gradient wasfluorometrically quantified with Quant-iT PicoGreenÒdsDNA Assay Kit(Invitrogen, Cergy-Pontoise,France)following the manufacturer’s instructions.Heavy and light DNA fractions were purified using the GeneClean kit(QBiogene,Montreal, Quebec)following the manufacturer’s protocol.Bacterial-Automated Ribosomal Intergenic Spacer Analysis of CommunitiesARISA was performed to characterize the diversity of bacterial communities at each sampling date(3,7,14,28,60,and120days of incubation)and for each treatment.The bacterial ribosomal intergenic spacer(IGS)of heavy and light DNA fractions was amplified by PCR using the primers L-D-Bact-132-a-A-18/S-D-Bact-1522-b-S-20.The S-D-Bact-1522-b-S-20was labeled at its5¢end with the IRD800fluorochrome for the detection of the resulted amplicons by a LiCorÒDNA sequencer(ScienceTec, Les Ulis,France)(Ranjard and others2006).The principle and procedure of the technique have been fully detailed in a previous article(Ranjard and others2006).PyrosequencingPyrosequencing of the16S rRNA gene was used to assess the bacterial diversity in the heavy and light DNA fractions extracted and fractionated from wheat-and alfalfa-amended microcosms after3,14, and60days of incubation,and from DNA extracted from control microcosms after3days of incubation.The selection of those particular samples was based on the results of the ARISA genotyping of the com-munities.Primers530F(5¢-GTGCCAGCMGCNGC GG-3¢)(Acosta-Martinez and others2008)and 803R(5¢-CTACCNGGGTATCTAAT-3¢)were used to amplify a16S rRNA gene fragment of the appropriate size(average amplicon length:290bases without tags)and sequence variability for454pyrosequenc-ing.Ten base pair tags at5¢position(ACGAGTGCGT, ACGCTCGACA,AGACGCACTC,AGCACTGTAG,A TCAGACACG,ATATCGCGAG,CGTGTCTCTA,CTC GCGTGTC,TAGTATCAGC,TCTCTATGCG,TGAT ACGTCT,and TACTGAGCTA)were used to identify each sample as recommended by the manufacturer (Beckman Coulter Genomics).For each treatment and sampling date analyzed,DNA extracted from the three replicates was pooled to perform the PCR.This pooling of the three replicates was performed on the basis of B-ARISA profiles that showed little variability between the three replicate microcosms for each treatment and sampling date analyzed by pyrose-quencing.Two ng aliquots of DNA from each pool were used for a50l l PCR under the following con-ditions:95°C for3min,30cycles of1min at95°C (denaturation),55°C for30s(annealing),and72°C for30s(extension),followed by5min at72°C.For each sample,amplicons offive replicated PCRs were purified using a MinElute gel extraction kit(Qiagen, Courtaboeuf,France)following the manufacturer’s protocol.Adapter sequences were added to300ng DNA per sample,as recommended by the manufac-turer and pyrosequencing was carried out on a Genome Sequencer FLX454(Beckman Coulter Linking Microbial Diversity to Priming Effect813Genomics).The16S rRNA gene fragments ranged between250and300bp.Taxonomic Assignments and Clustering of16S rRNA Gene FragmentsAfter pyrosequencing,943,184raw reads were obtained and have been analyzed using two different bioinformatic pipelines,as described in Stoeck and others(2009).Thefirst pipeline was assigned to the cleaning,denoising and clustering of tags as described previously(Stoeck and others2009;Behnke and others2011).Sequences without any ambiguities (without Ns)werefirst strictly dereplicated and sorted according to decreasing number of occurrences.They were then clustered when they had three differences or less with a seed sequence,ignoring differences due to homopolymers(which are the major454 sequencing errors).Finally,reads detected only once and not clustered(named singletons)were deleted,to eliminate potential errors and to avoid an overesti-mated number of clusters(deleted singletons repre-sented only5%of all reads,but more than55%of detected clusters)(Pawlowski and others2011).Our simplefiltering method resulted in a higher rate of deleted sequences compared to the method of Quince and others(2009),($5%to$3%in their example), suggesting that our approach is probably more strin-gent(Quince and others2009;Stecher and others 2010).In fact,a majority of the singletons were gen-erally defined as artifacts(for example,PCR chimeras and/or containing large sequencing errors)resulting from noise in high-throughput pyrosequencing of the most abundant clusters(Fodor and others2012; Stearns and others2011;Reeder and Knight2009). After these different qualityfilters616,881high-quality reads were kept and rarefaction curves and diversity indexes(that is,Shannon)werefinally computed using cleaned clusters.The second pipeline was for taxo-nomic assignment of16S rRNA gene fragments.The 16S rRNA gene fragments were compared with se-quences from the Silva reference database(release104: http://www.arb-silva.de/documentation/background/ release-104/)(Pruesse and others2007).Briefly,each quality-checked read was compared with these refer-ence sequences by a similarity search using global alignments and requesting an output containing any hit at70%or more similarity,with penalties of1for mismatch,gap open and gap extended.Reference sequences with a similarity above a given threshold percentage(here at97%,details below)were retained and a consensus was built for the taxonomies found, which was then assigned to the tag.Various thresholds were tested(99,98.5,98,97.5,97,96,95,94,93,92,91, 90,85,and80%),and after manual examination of the relationships between threshold and qualities of taxo-nomic assignment,the97%similarity threshold was chosen for assigning a given tag to a phylum. Statistical AnalysisTo study the significance of the variation in total CO2release,plant residue mineralization and PE with time between wheat,alfalfa,and control treatments,we used a linear mixed effects(LME) model on CO2release data with log transformation for time parameter,followed by an ANOVA to evaluate significant differences determined at P<0.01.These statistical calculations were carried out with R software version2.11.1with the nlme package(Pinheiro and others2012).For B-ARISA results,the data obtained from the 1D-Scan software were converted into a table summarizing the band presence and intensity using the PrepRISA program(Ranjard and others2006) using R software version2.11.1(the R Foundation for Statistical Computing,Vienna,Austria).This program allowed us to choose the number of peaks, the profile resolution and the method of evaluating peak intensity.As described in a previous study,we used100peaks,2bp resolution and Gaussian peak area for a robust analysis of bacterial communities (Ranjard and others2006).The B-ARISA data matrix(bacterial communities as rows and bands as columns),was subjected to principal component analysis(PCA)on a covariance matrix.This provided an ordination of bacterial communities in a factorial map based on the scores of thefirst two principal components.The significance of the discrimination of the genetic structures of the bacterial communities revealed by PCA was con-firmed by performing the Monte Carlo test with 1000permutations.PCA and Monte Carlo tests were performed using the ADE-4package(Thioulouse and others1997;Chessel and others2004)using R software version2.11.1(the R Foundation for Sta-tistical Computing,Vienna,Austria). Nucleotide Sequence Accession NumbersThe nucleotide sequences determined in this study have been deposited to the SRA of EBI database with the accession number:ERP001023.R ESULTSSoil and Residue C MineralizationThe soil respiration rate was rapidly increased by the addition of plant residues compared to the814N.Pascault and otherscontrol (Figure 1A),with a greater stimulation for alfalfa (P <0.01).At the end of incubation,1050and 760l g g -1soil C-CO 2had been emitted for alfalfa and wheat treatments,respectively.Mineralization of alfalfa and wheat residues fol-lowed the same dynamics,with an initial exponen-tial phase during the first 2weeks of incubation,a slowdown between 14and 28days,and finally a weak 13C-CO 2release until the end of the incubation (Figure 1B).Alfalfa residues decomposed signifi-cantly faster than wheat residues (P <0.01)because we estimated that about 21and 9%of alfalfa and wheat residues,respectively,had been miner-alized at 28days,and about 28and 17%at 120days.The release of 12C-CO 2was also significantly stimulated in both alfalfa and wheat treatments during the incubation,as compared to the control.This pointed to a PE of native SOM,with the degree of priming being significantly related to the qualityof FOM (P <0.01)(Figure 1B).More precisely,PE occurred mainly during the first month of incuba-tion,representing 205.2l g C g -1soil and 404.5l g C g -1soil 28days after wheat and alfalfa addition,and 293.5l g C g -1soil and 458.1l g C g -1soil after 120days,corresponding to an increase of 220and 343%,respectively,in SOM decomposition.Genetic Structure of Bacterial CommunitiesBacterial-ARISA was performed on heavy and light DNA fractions to characterize the dynamics of popu-lations involved in the decomposition of 13C-labeled residues and SOM,respectively.Visual analysis of the B-ARISA profiles highlighted differences between the light and the heavy fractions in terms of relative intensity,but also of presence/absence of specific bands (Figure 1S).This suggested that the input of both wheat and alfalfa residues stimulated particular active bacterial communities.PCA of the profiles confirmed visual observations because,for each sampling date,the heavy fraction was significantly discriminated from the light fraction on the factorial maps (Monte Carlo test,1000permutations)(Fig-ure 2).Examination of the profile patterns of the bacterial communities actively involved in degrada-tion of the plant residues (FOM-degrading commu-nities),revealed a similar pattern of community dynamics for both residues (Figure 2),characterized by a highly dynamic phase from 3to 14days followed by a slow down of community modifications from 28to parison of active bacterial commu-nities between wheat and alfalfa treatments showed that different communities were stimulated depend-ing on the biochemical quality of the residue (Fig-ure 2S).Those bacterial communities that were consid-ered as not involved in the degradation of residues did not show any significant dynamics in the wheat treatment,with all the sampling dates grouping together on the factorial map (Figure 2A).In con-trast,significant dynamics were observed in the alfalfa treatment,mainly represented by the dis-crimination between an early (from 3to 14days)and a late (from 28to 120days)phase (confirmed by Monte Carlo test)(Figure 2B).Taxonomic Composition of Bacterial CommunitiesExamination of the taxonomic affiliation of the 616,881classifiable bacterial sequences obtained revealed a broad diversity,with 266different gen-era detected.Most of these genera (87%of all theB10001200 g o f s o i l )Alfalfa Wheat6080100120alfalfa wheate m i n e r a l i z a t i o n C O 2/ g of s o i l )ALinking Microbial Diversity to Priming Effect815identified sequences)were distributed across nine major bacterial phyla that were present in all samples(Figure3).Shannon diversity(H)indices as well as rare-faction curves calculated for each library showed that bacterial diversity retrieved from the light DNA fraction was higher compared to the heavy DNA fraction for each sampling date and each treatment (Figures3,3S).This discrepancy between the heavy and light fractions was particularly large at day3for both wheat(H=5.0vs.H=6.0)and alfalfa(H=4.5vs.H=5.6),mainly due to a con-siderably lower species richness in the heavy frac-tion compared to the light fraction for both wheat (2012vs.1179molecular operational taxonomic units(MOTU)detected)and alfalfa(1840vs.909 MOTU detected).At each date and for each residue,the phyloge-neticfingerprints of the communities in the light fractions were distinct from those of the corre-sponding heavy fractions(Figure4A).The differ-ence between the heavy and light fractions was the main parameter discriminating all the libraries, mainly due to an increase in the relative proportion of Proteobacteria and Firmicutes in the heavy frac-tions,and of Acidobacteria,and to a lesser extent Verrucomicrobia and Gemmatimonadetes in the light fractions(Figures3,4B).At3days,Firmicutes dominated the‘‘heavy’’library for wheat(44%of all sequences)whereas most of the sequences retrieved were affiliated to the Proteobacteria phylum for alfalfa(64%of all sequences)(Figures3,4B).At14and60days, the FOM-degrading communities for wheat and alfalfa showed more similar compositions,and were dominated by bacteria belonging to the Proteobacteria phylum(mainly the a-,b-,and c-subgroups).The phylogeneticfingerprints obtained from the light fractions of the wheat treatment were close to those of the control and did not show strong dynamics between the three sampling dates(Fig-ure4A).Contrastingly,changes were observed in the light fraction of the alfalfa treatment.After 3days of incubation,compared to the other sampling dates,most of the discrimination was explained by an increase in sequences affiliated to the Firmicutes and Bacteroidetes groups(Figures 3,4B).D ISCUSSIONSoil and Residue C MineralizationThe patterns of soil respiration observed in our study were in agreement with other authors (Xu and others2006;Cayuela and others2009), that is,decomposition occurring during the 120days of incubation with a peak during thefirst 2weeks.This dynamic may be explained by the decomposition of easily degradable compounds during thefirst steps of the decomposition process,816N.Pascault and others。