Do bacterial and fungal communities assemble differently during primary succession?
|
|
- Phillip Lester
- 5 years ago
- Views:
Transcription
1 Molecular Ecology (2014) 23, NEWS AND VIEWS PERSPECTIVE Do bacterial and fungal communities assemble differently during primary succession? S. K. SCHMIDT,* D. R. NEMERGUT, J. L. DARCY* and R. LYNCH* *Department of Ecology and Evolutionary Biology, University of Colorado, Boulder, CO, 80309, USA; Institute of Arctic and Alpine Research, University of Colorado, Boulder, CO 80309, USA; Environmental Studies Program, University of Colorado, Boulder, CO 80309, USA High-throughput sequencing technologies are now allowing us to study patterns of community assembly for diverse microbial assemblages across environmental gradients and during succession. Here we discuss potential explanations for similarities and differences in bacterial and fungal community assembly patterns along a soil chronosequence in the foreland of a receding glacier. Although the data are not entirely conclusive, they do indicate that successional trajectories for bacteria and fungi may be quite different. Recent empirical and theoretical studies indicate that smaller microbes (like most bacteria) are less likely to be dispersal limited than are larger microbes which could result in a more deterministic community assembly pattern for bacteria during primary succession. Many bacteria are also better adapted (than are fungi) to life in barren, early-successional sediments in that some can fix nitrogen and carbon from the atmosphere traits not possessed by any fungi. Other differences between bacteria and fungi are discussed, but it is apparent from this and other recent studies of microbial succession that we are a long way from understanding the mechanistic underpinnings of microbial community assembly during ecosystem succession. We especially need a better understanding of global and regional patterns of microbial dispersal and what environmental factors control the development of microbial communities in complex natural systems. Keywords: bacteria, climate change, community ecology, fungi, landscape genetics, phylogeography Received 24 October 2013; revised xxxx; accepted 12 November 2013 The conceptual framework for understanding primary succession was initially developed from the study of plant Correspondence: Steve Schmidt, Fax: ; Steve.Schmidt@Colorado.edu systems, with the earliest work dating to 1685 (Clements 1916), but also grew out of studies of pedogenesis (soil development, e.g. Jenny 1980) and more recently from generalized community assembly theory (Vellend 2010; Nemergut et al. 2013). Plants, microbes and soils share obvious connections, but it was not until the rise of high-throughput sequencing technologies that phylogenetic community profiles of multiple microbial communities could be rapidly inventoried. We now have the opportunity to test ecological hypotheses developed from the study of macroscopic plants and animals, as well as emerging hypotheses specific to microorganisms (Nemergut et al. 2013). Studies of microbial succession are also becoming more important because global warming is causing unprecedented rates of glacial retreat especially in high-elevation and high-latitude environments where the rate of plant colonization is quite slow compared with that of microbes (Fig. 1). Brown & Jumpponen (2014) studied bacterial and fungal community succession and assembly along a chronosequence of soils created by the receding Lyman Glacier in the northern Cascade Range. By comparing community assembly of bacteria and the fungi in these new soils, they address several specific questions, including: Do bacterial and fungal communities follow similar assembly trajectories along the chronosequence? Does the presence of plants alter the assembly trajectories for bacteria and fungi? The study of Brown & Jumpponen (2014) can also be viewed in light of the debate about whether historical or deterministic factors are more important in controlling community assembly during succession. This debate originated from the deterministic view of Clements (e.g. 1916, 1936) and the more historical view of Gleason (e.g. 1926) and has recently been addressed in studies of microbial systems (e.g. Peay et al. 2012; Ferrenberg et al. 2013). Here, we discuss the results of Brown & Jumpponen (2014) from the perspective of this debate and with respect to the growing body of knowledge about microbial dispersal, biogeography and biogeochemistry. Perhaps the most interesting finding of Brown & Jumpponen (2014) was that a greater fraction (19%) of fungal OTUs displays nonrandom patterns of occurrence along the chronosequence compared with bacterial OTUs (9.5%). However, when examined as a whole, bacterial communities were more affected by both distance along the chronosequence and vegetation cover than were fungal communities. In addition, bacterial communities converged along the chronosequence, whereas fungal community assembly appeared to be more stochastic (less deterministic) and showed no evidence of convergence towards one community type (Brown & Jumpponen 2014). Although the data of Brown & Jumpponen (2014) are by no means conclusive, they do hint at different trajectories of
2 NEWS AND VIEWS: PERSPECTIVE 255 Fig. 1 Repeat photography of a site in the High Andes of Peru where rapid glacial retreat is exposing large tracts of land that are rapidly colonized by microbes (months to years) but only slowly colonized by plants (decades to centuries). The top photograph was taken in 2005 and the bottom in 2010 (the person in each photograph is standing in approximately the same location). The photographer was standing at about 5200 m above sea level near the 100-m site viewable in an aerial photograph of the site previously published in Schmidt et al. (2009). The distance from the person to the closest edge of the ice is approximately 20 and 200 meters in the top and bottom photos, respectively. Note the edge of a lake that formed between 2005 and 2010 (lower left corner of bottom photo). Photograph credits S.K. Schmidt and J.L. Darcy. succession for bacteria and fungi at this site. If bacterial communities converge on one community type during succession, and fungi do not, this may indicate that bacterial communities assemble in a more deterministic fashion than do fungi. One likely explanation for this pattern is that fungi may be more dispersal limited than bacteria and therefore more prone to historical (stochastic) effects at this site (Fig. 2). Some recent studies indicate that early-successional Betaproteobacteria (e.g. Polaromonas spp.) are not dispersal limited at the global scale (Darcy et al. 2011), whereas larger microbes like algae and zoosporic fungi (both common in periglacial environments) show more divergent biogeographic patterns (and therefore perhaps dispersal limitation) at global and regional scales (De Wever et al. 2009; Schmidt et al. 2011; Naff et al. 2013). Recent modelling studies also support a difference in dispersal capabilities between smaller (e.g. bacteria) and larger (e.g. fungi) microbes. Wilkinson et al. (2012) showed that there is a very low probability that microbes greater than 20 lm in diameter can undergo passive dispersal between continents and that the successful dispersal of small microbes is due to their greater abundance and their longer residence times in the atmosphere compared with larger microbes. Thus, both empirical and theoretical studies point to a more consistent propagule rain (sensu Brown & Jumpponen 2014) of bacteria at early-successional sites (Fig. 2). A more consistent propagule rain for bacteria could reduce or eliminate priority effects for bacterial communities resulting in more deterministic community assembly across the landscape compared with fungi. Other research
3 256 NEWS AND VIEWS: PERSPECTIVE Time Propagules Fig. 2 The influence of the timing and contents of propagule rain is depicted. In some cases, dispersal and establishment are successful (solid blue arrows). In other cases, dispersal is not successful and the microorganisms perish in transit or on arrival (dashed lines) or may be poorly adapted to the new environment (teal hexagons). Priority effects may also play an important role in microbial community assembly during succession. For example, the red ovals and yellow squares arrive first, gaining a competitive advantage as they monopolize resources. Later migrants (small green circles) may not be competitive due to their lower relative abundance and the sequestration of limiting nutrients by earlier colonists (e.g. the red ovals). Had the green circles arrived first instead of the red ovals, they may have been more successful and the community would have assembled differently. has shown that priority effects can lead to greater divergence in fungal community assembly (Peay et al. 2012) and are more likely to occur when the rate of propagule input is low (Chase 2003). For example, priority effects determined the ultimate fungal community structure of two ectomycorrhizal species on pine roots (Kennedy & Bruns 2005). Likewise, complex wood decomposing fungal communities can be driven to significantly different successional outcomes by the order of addition of community members (Fukami et al. 2010). Another potential reason that bacteria and fungi have different early-successional trajectories is that bacteria exhibit a broader range of physiologies than do fungi and thus are more likely to be successful colonists of the oligotrophic, plant-free soils near the glacier terminus. Earlysuccessional bacteria can be photoautotrophs, heterotrophs or chemoautotrophs and many can fix atmospheric nitrogen (Nemergut et al. 2007; Schmidt et al. 2008b; Duc et al. 2009), whereas fungi are all heterotrophs and none can fix nitrogen. Thus, fungi are more dependent than bacteria on fixed sources of carbon and nitrogen and may not have as many available niches before there is significant organic matter build-up during succession. Indeed, it may be that many of the fungi present in recently deglaciated sites are actually dormant (due to lack of organic matter) as originally suggested by Jumpponen (2003), in which case their distribution across the landscape would probably be more stochastic due to the dispersal constraints discussed above. Alternatively, wind-blown particles tend to accumulate in protected pockets, for example next to rocks (Swan 1992), which would cause a very nonuniform (seemingly stochastic) accumulation of organic matter (and therefore fungi) across the early-successional landscape. Brown & Jumpponen (2014) also point out that many of the fungal OTUs at their sites are related to fungi known to be associated with insects. This observation supports the hypothesis of Hodkinson et al. (2002) and Swan (1992) that wind-blown arthropods can be important sources of organic matter in recently deglaciated environments. As highlighted by Brown & Jumpponen (2014) and other authors, one of the big mysteries surrounding the early stages of microbial succession is determining the source of carbon and energy for the early colonists. In addition to allochthonous (mostly wind-blown) carbon (C) inputs discussed above, the two other major sources of C to early-successional soils are ancient C (Welker et al. 2002; Bardgett et al. 2007; Sattin et al. 2009) and C fixed autochthonously by photo- and chemoautotrophs (Nemergut et al. 2007; Schmidt et al. 2008b). The relative contribution of each of these C sources to the overall pool of C in earlysuccessional soils probably has a major effect on the structure of the resulting microbial communities (Fierer et al. 2010). For example, sites with relatively high levels of ancient C, or high inputs of wind-blown C, might be dominated by heterotrophs early in succession. By contrast, sites with low levels of wind-blown and ancient C would probably be dominated by autotrophs early in microbial succession. As sunlight is abundant at the soil surface before plants colonize periglacial soils, it is logical to assume that photoautotrophs will be important players in most newly deglaciated soils especially in areas with low levels of soil carbon. However, other factors may stunt the development of microbes early in succession, such as limitations of essential nutrients (Yoshitake et al. 2007; G oransson et al. 2011; Schmidt et al. 2012). Given the importance of C and other elements for the development of microbial communities, more information is needed on the geochemistry and aeolian inputs of nutrients to all of the earlysuccessional sites presently being studied by microbial ecologists (Mladenov et al. 2012). Finally, the relationship between plant and soil microbial communities in early-successional sites should be highly correlated (Blaalid et al. 2012; Knelman et al. 2012) as the nutrient cycling cascades catalysed by microorganisms in plant-free soils are likely to strongly influence the success of plant colonization and because plants add substantial inputs of carbon to the soils through both litter and root exudates. Most importantly, plants form symbiotic relationships with many soil bacteria and fungi and should therefore skew microbial community assembly towards symbiotic heterotrophs. Therefore, it was somewhat surprising that Brown & Jumpponen (2014) found that the presence of plants played a very minor role in the distribution of fungal OTUs relative to bacterial OTUs at the community level. This difference could be explained by dispersal limitations of fungi relative to bacteria as discussed above. Previous studies have shown that inoculum levels for the most common type of plant
4 NEWS AND VIEWS: PERSPECTIVE 257 symbionts, arbuscular mycorrhizal fungi, are often below detection limits in early-successional soils favouring weedy, nonmycorrhizal plants early in succession (Miller 1979; Schmidt et al. 2008a). However, it is notable that, when examined at the community level, the presence of specific plant taxa had a significant influence on fungal, as well as bacterial composition (Brown & Jumpponen 2014). It would be interesting to re-examine the data of Brown & Jumpponen (2014) using a nested, multivariate approach to partition the relative influence of distance along the chronosequence vs. plant cover (presence or absence of vegetation as well as plant species type) on bacterial and fungal community composition as well as the relationships between these two groups. Likewise, a deeper exploration of soil physiochemical parameters and/or the use of null deviation analyses could help identify the cause of the difference in patterns of bacterial distribution along the chronosequence. Brown & Jumpponen (2014) posit that bacterial assembly processes are more stochastic very early in succession, but not later in succession, a pattern that has been documented in other systems (e.g. Ferrenberg et al. 2013). However, it is also possible that these seemingly stochastic patterns are actually due to heterogeneity in environmental filters (e.g. soil organic matter as discussed above) and that plant invasion serves to homogenize the landscape, making assembly appear to be more deterministic in older soils. Overall, the study of Brown & Jumpponen (2014) and other recent studies (e.g. Zumsteg et al. 2012) illuminate the importance of studying more than just one component of the microbial community and point the way to future work needed to understand the patterns they report. Especially needed are studies of the relative dispersal abilities of microbial groups and studies of nutrient levels, inputs and limitations in early-successional systems. New, highthroughput sequencing technologies have allowed us to finally describe the spatial and temporal patterns of microbial communities, but mechanistic studies that isolate the relative importance of the individual drivers of microbial community assembly lag far behind. References Bardgett RD, Richter A, Bol R et al. (2007) Heterotrophic microbial communities use ancient carbon following glacial retreat. Biology Letters, 3, Blaalid R, Carlsen T, Kumar S et al. (2012) Changes in root-associated fungal communities along a primary succession gradient analyzed by 454 pyrosequencing. Molecular Ecology, 21, Brown SP, Jumpponen A (2014) Contrasting primary successional trajectories of fungi and bacteria in retreating glacier soils. Molecular Ecology, 23, Chase JM (2003) Community assembly: when should history matter? Oecologia, 136, Clements FE (1916) Plant Succession: An Analysis of the Development of Vegetation (No. 242). Carnegie Institution of Washington, Washington, DC. Clements FE (1936) Nature and structure of the climax. Journal of Ecology, 24, Darcy JL, Lynch RC, King AJ, Robeson MS, Schmidt SK (2011) Global distribution of Polaromonas phylotypes - evidence for a highly successful dispersal capacity. PLoS ONE, 6, e doi: / journal.pone De Wever A, Leliaert F, Verleyen E et al. (2009) Hidden levels of phylodiversity in Antarctic green algae: further evidence for the existence of glacial refugia. Proceedings. Biological sciences/the Royal Society, 276, Duc L, Noll M, Meier BE, B urgmann H, Zeyer J (2009) High diversity of diazotrophs in the forefield of a receding Alpine glacier. Microbial Ecology, 57, Ferrenberg S, O Neill SP, Knelman JE et al. (2013) Changes in assembly processes in soil bacterial communities following a wildfire disturbance. The ISME Journal, 7, Fierer N, Nemergut D, Knight R, Craine JM (2010) Changes through time: integrating microorganisms into the study of succession. Research in Microbiology, 161, Fukami T, Dickie IA, Wilkie JP et al. (2010) Assembly history dictates ecosystem functioning: evidence from wood decomposer communities. Ecology Letters, 13, Gleason HA (1926) The individualistic concept of the plant association. Bulletin of the Torrey Botanical Club, 53, G oransson H, Venterink HO, Baath E (2011) Soil bacterial growth and nutrient limitation along a chronosequence from a glacier forefield. Soil Biology and Biochemistry, 43, Hodkinson ID, Webb NR, Coulson SJ (2002) Primary community assembly on land the missing stages: why are the heterotrophic organisms always first? Journal of Ecology, 90, Jenny H (1980) Soil Genesis with Ecological Perspectives. Springer-Verlag, New York. Jumpponen A (2003) Soil fungal community assembly in a primary successional glacier forefront ecosystem as inferred from rdna sequence analyses. New Phytologist, 158, Kennedy PG, Bruns TD (2005) Priority effects determine the outcome of ectomycorrhizal competition between two Rhizopogon species colonizing Pinus muricata seedlings. New Phytologist, 166, Knelman JE, Legg TM, O Neill SP et al. (2012) Bacterial community structure and function change in association with colonizer plants during early primary succession in a glacier forefield. Soil Biology & Biochemistry, 46, Miller RM (1979) Some occurrences of vesicular arbuscular mycorrhiza in natural and disturbed ecosystems of the Red Desert. Canadian Journal of Botany, 57, Mladenov N, Williams MW, Schmidt SK, Cawley K (2012) Atmospheric deposition as a source of carbon and nutrients to an alpine catchment of the Colorado Rocky Mountains. Biogeosciences, 9, Naff CS, Darcy JL, Schmidt SK (2013) Phylogeny and biogeography of an uncultured clade of Snow Chytrids. Environmental Microbiology, 15, Nemergut DR, Anderson SP, Cleveland CC et al. (2007) Microbial community succession in unvegetated, recently-deglaciated soils. Microbial Ecology, 53, Nemergut DR, Schmidt SK, Fukami T et al. (2013) Patterns and processes of microbial community assembly. Microbiology and Molecular Biology Reviews, 77, Peay KG, Belisle M, Fukami T (2012) Phylogenetic relatedness predicts priority effects in nectar yeast communities. Proceedings. Biological sciences/the Royal Society, 279, Sattin SR, Cleveland CC, Hood E et al. (2009) Functional shifts in unvegetated, perhumid, recently deglaciated soils do not
5 258 NEWS AND VIEWS: PERSPECTIVE correlate with shifts in soil bacterial community composition. Journal of Microbiology, 47, Schmidt SK, Sobieniak-Wiseman LC, Kageyama SA et al. (2008a) Mycorrhizal and dark-septate fungi in plant roots above 4270 meters elevation in the Andes and Rocky Mountains. Arctic, Antarctic, and Alpine Research, 40, Schmidt SK, Reed SC, Nemergut DR et al. (2008b) The earliest stages of ecosystem succession in high-elevation, recently de-glaciated soils. Proceedings. Biological sciences/the Royal Society, 275, Schmidt SK, Nemergut DR, Miller AE et al. (2009) Microbial activity and diversity during extreme freeze-thaw cycles in periglacial soils, 5400 m elevation, Cordillera Vilcanota, Peru. Extremophiles, 13, Schmidt SK, Lynch RC, King AJ et al. (2011) Phylogeography of microbial phototrophs in the dry valleys of the high Himalayas and Antarctica. Proceedings. Biological sciences/the Royal Society, 278, Schmidt SK, Nemergut DR, Todd BT et al. (2012) A simple method for determining limiting nutrients for photosynthetic crusts. Plant Ecology & Diversity, 5, Swan LW (1992) The aeolian biome, ecosystems of the Earth s extremes. BioScience, 42, Vellend M (2010) Conceptual synthesis in community ecology. Review Literature and Arts of the Americas, 85, Welker JM, Fahnestock JT, Henry GHR, O Dea KW, Piper RE (2002) Microbial activity discovered in previously ice-entombed Arctic ecosystems. EOS Transactions American Geophysical Union, 83, Wilkinson DM, Koumoutsaris S, Mitchell EAD, Bey I (2012) Modeling the effect of size on the aerial dispersal of microorganisms. Journal of Biogeography, 39, Yoshitake S, Uchida M, Koizumi H, Nakatsubo T (2007) Carbon and nitrogen limitation of soil microbial respiration in high Arctic successional glacier foreland near Ny- Alesund, Svalbard. Polar Research, 26, Zumsteg A, Luster J, G oransson H et al. (2012) Bacterial, archaeal and fungal succession in the forefield of a receding glacier. Microbial Ecology, 63, S.S., D.N., J.D. and R.L. wrote the paper. doi: /mec.12589
Role of mycorrhizal fungi in belowground C and N cycling
Role of mycorrhizal fungi in belowground C and N cycling Doc. Jussi Heinonsalo Department of Forest Sciences, University of Helsinki Finnish Meteorological Institute Finland The aim and learning goals
More informationNature: a harmonious & peaceful place?! What disturbs the peace?
Nature: a harmonious & peaceful place?! What disturbs the peace? Disturbance Disturbance: a relatively discrete event in time that causes abrupt change in ecosystem, community, or population structure,
More informationSuccession: A Closer Look
Succession: A Closer Look By: Sarah M. Emery (Department of Biology, University of Louisville) 2010 Nature Education Citation: Emery, S. (2010) Succession: A Closer Look. Nature Education Knowledge 3(10):45
More informationPeter Gault Kennedy CURRICULUM VITAE. 321 Koshland Hall phone: University of California, Berkeley fax: Berkeley, CA 94720
Peter Gault Kennedy CURRICULUM VITAE Department of Plant and Microbial Biology pkennedy@berkeley.edu 321 Koshland Hall phone: 510-643-5483 University of California, fax: 510-642-4995, CA 94720 Professional
More informationCommunity phylogenetics review/quiz
Community phylogenetics review/quiz A. This pattern represents and is a consequent of. Most likely to observe this at phylogenetic scales. B. This pattern represents and is a consequent of. Most likely
More informationMCMURDO DRY VALLEYS ANTARCTICA
POLAR WEEKEND MCMURDO DRY VALLEYS ANTARCTICA TOPIC: The McMurdo Dry Valleys, An Ecosystem of microbial life PROJECT PERSONNEL: Dr. Andrew G. Fountain, Geologist, Portland State University, Portland Oregon
More informationSuccession. Lesson Overview. Lesson Overview. 4.3 Succession
Lesson Overview 4.3 THINK ABOUT IT In 1883, the volcanic island of Krakatau in the Indian Ocean was blown to pieces by an eruption. The tiny island that remained was completely barren. Within two years,
More informationPOPULATIONS and COMMUNITIES
POPULATIONS and COMMUNITIES Ecology is the study of organisms and the nonliving world they inhabit. Central to ecology is the complex set of interactions between organisms, both intraspecific (between
More informationComputational Ecology Introduction to Ecological Science. Sonny Bleicher Ph.D.
Computational Ecology Introduction to Ecological Science Sonny Bleicher Ph.D. Ecos Logos Defining Ecology Interactions: Organisms: Plants Animals: Bacteria Fungi Invertebrates Vertebrates The physical
More informationGeorgia Performance Standards for Urban Watch Restoration Field Trips
Georgia Performance Standards for Field Trips 6 th grade S6E3. Students will recognize the significant role of water in earth processes. a. Explain that a large portion of the Earth s surface is water,
More informationSoil Biology. Chapter 10
Soil Biology Chapter 10 The Sounds of Soil Soil as a Transition Between Aquatic and Aerial System Bacteria in a Drying Environment Wet (open structure) Dry (dense) Holden P.A., J.R. Hunt, and M. K. Firestone,
More informationChapter 8. Biogeographic Processes. Upon completion of this chapter the student will be able to:
Chapter 8 Biogeographic Processes Chapter Objectives Upon completion of this chapter the student will be able to: 1. Define the terms ecosystem, habitat, ecological niche, and community. 2. Outline how
More information1 Soil Factors Affecting Nutrient Bioavailability... 1 N.B. Comerford
Contents 1 Soil Factors Affecting Nutrient Bioavailability........ 1 N.B. Comerford 1.1 Introduction........................... 1 1.2 Release of Nutrients from the Soil Solid Phase........ 2 1.3 Nutrient
More information1 Towards Ecological Relevance Progress and Pitfalls in the Path Towards an Understanding of Mycorrhizal Functions in Nature... 3 D.J.
Contents Section A: Introduction 1 Towards Ecological Relevance Progress and Pitfalls in the Path Towards an Understanding of Mycorrhizal Functions in Nature... 3 D.J. Read 1.1 Summary.............................
More informationTHE CHANGING SURFACE OF THE EARTH
THE CHANGING SURFACE OF THE EARTH Key words Drain geological agent weathering erosion Sediment deposition transport The landscape is a consequence of the action of two types of geological processes; internal
More informationRank-abundance. Geometric series: found in very communities such as the
Rank-abundance Geometric series: found in very communities such as the Log series: group of species that occur _ time are the most frequent. Useful for calculating a diversity metric (Fisher s alpha) Most
More informationAssociations between an invasive plant (Taeniatherum caputmedusae, Medusahead) and soil microbial communities
Associations between an invasive plant (Taeniatherum caputmedusae, Medusahead) and soil microbial communities Elise S. Gornish¹, Noah Fierer², Albert Barberán¹ ¹University of California, Davis; ²University
More informationSecondary Succession and its Effects on Soil Nutrients and Fungal Communities. Amanda Cayo
Cayo 1 Secondary Succession and its Effects on Soil Nutrients and Fungal Communities Amanda Cayo Abstract Fungi serve many purposes in ecosystems from fixing nitrogen for plants to decomposing detritus.
More informationLast Time. Biology of FUNgi. Lecture 24 Community ecology - what rules fungal communities. ! We wrapped up the good, the bad and the ugly.
Biology of FUNgi Lecture 24 Community ecology - what rules fungal communities Last Time! We wrapped up the good, the bad and the ugly.! Systemic fungal infections: blastomycosis, coccidiomycosis, cryptococcosis,
More informationDigital ESF. SUNY College of Environmental Science and Forestry. Max Hermanson. Silus Weckel. Alex Kozisky.
SUNY College of Environmental Science and Forestry Digital Commons @ ESF Cranberry Lake Biological Station Environmental and Forest Biology 2017 Session D, 2017 First Place: Under the Sphagnum: An Observational
More informationName: Characteristics of Life and Ecology Guided Notes (PAP)
Name: Characteristics of Life and Ecology Guided Notes (PAP) I. What is Biology? a. Biology is the study of II. The Eight Characteristics of Life a. Organization & the presence of or more cells b. Response
More informationUnit 3 Study Guide -- Greenberg science, 6C
Unit 3 Study Guide -- Greenberg science, 6C Name Pd. Date / / 2018 + +5 extra credit points on the test if submitted complete and correct ON THE TEST DATE. + A copy can be found on my website mgreenberg.weebly.com
More informationLesson Overview. Niches and Community Interactions. Lesson Overview. 4.2 Niches and Community Interactions
Lesson Overview 4.2 Niches and Community Interactions The Niche What is a niche? A niche is the range of physical and biological conditions in which a species lives and the way the species obtains what
More informationOrbital-Scale Interactions in the Climate System. Speaker:
Orbital-Scale Interactions in the Climate System Speaker: Introduction First, many orbital-scale response are examined.then return to the problem of interactions between atmospheric CO 2 and the ice sheets
More informationThe Tempo of Macroevolution: Patterns of Diversification and Extinction
The Tempo of Macroevolution: Patterns of Diversification and Extinction During the semester we have been consider various aspects parameters associated with biodiversity. Current usage stems from 1980's
More informationEcosystem change: an example Ecosystem change: an example
5/13/13 Community = An assemblage of populations (species) in a particular area or habitat. Here is part of a community in the grassland of the Serengetti. Trophic downgrading of planet Earth: What escapes
More informationHonors Biology Unit 5 Chapter 34 THE BIOSPHERE: AN INTRODUCTION TO EARTH S DIVERSE ENVIRONMENTS
Honors Biology Unit 5 Chapter 34 THE BIOSPHERE: AN INTRODUCTION TO EARTH S DIVERSE ENVIRONMENTS 1. aquatic biomes photic zone aphotic zone 2. 9 terrestrial (land) biomes tropical rain forest savannah (tropical
More informationSpheres of Life. Ecology. Chapter 52. Impact of Ecology as a Science. Ecology. Biotic Factors Competitors Predators / Parasites Food sources
"Look again at that dot... That's here. That's home. That's us. On it everyone you love, everyone you know, everyone you ever heard of, every human being who ever was, lived out their lives. Ecology Chapter
More informationThe World of Lichens Part of: Joint Science Education Project at Dartmouth Developed by: Ruth Heindel, Earth Sciences Department, Dartmouth College
The World of Lichens Part of: Joint Science Education Project at Dartmouth Developed by: Ruth Heindel, Earth Sciences Department, Dartmouth College Overview Lichens are incredible symbiotic organisms that
More informationClimate Change. Unit 3
Climate Change Unit 3 Aims Is global warming a recent short term phenomenon or should it be seen as part of long term climate change? What evidence is there of long-, medium-, and short- term climate change?
More informationWeather is the day-to-day condition of Earth s atmosphere.
4.1 Climate Weather and Climate Weather is the day-to-day condition of Earth s atmosphere. Climate refers to average conditions over long periods and is defined by year-after-year patterns of temperature
More informationFernando A. O. Silveira Universidade Federal de Minas Gerais, Brazil
Fernando A. O. Silveira Universidade Federal de Minas Gerais, Brazil www.leept.webnode.com Inselbergs are geologically-ancient, nutrientimpoverished granitic and gneiss monoliths that rise sharply above
More informationChapter 4 AND 5 Practice
Name: Chapter 4 AND 5 Practice 1. Events that occur in four different ecosystems are shown in the chart below. Which ecosystem would most likely require the most time for ecological succession to restore
More informationSAMPLE PAGE. pulses. The Ice Age By: Sue Peterson
Page 61 Objective sight words (pulses, intermittent, isotopes, chronicle, methane, tectonic plates, volcanism, configurations, land-locked, erratic); concepts (geological evidence and specific terminology
More informationDescribe how ecosystems recover from a disturbance. Compare succession after a natural disturbance with succession after a human-caused disturbance.
1 2 Objectives Describe how ecosystems recover from a disturbance. Compare succession after a natural disturbance with succession after a human-caused disturbance. 3 Succesion Cartoon Guide to the Environment
More informationChapter 52: An Introduction to Ecology and the Biosphere
AP Biology Guided Reading Name Chapter 52: An Introduction to Ecology and the Biosphere Overview 1. What is ecology? 2. Study Figure 52.2. It shows the different levels of the biological hierarchy studied
More informationMarine Resources Development Foundation/MarineLab Grades: 9, 10, 11, 12 States: AP Biology Course Description Subjects: Science
Marine Resources Development Foundation/MarineLab Grades: 9, 10, 11, 12 States: AP Biology Course Description Subjects: Science Highlighted components are included in Tallahassee Museum s 2016 program
More informationLecture 24 Plant Ecology
Lecture 24 Plant Ecology Understanding the spatial pattern of plant diversity Ecology: interaction of organisms with their physical environment and with one another 1 Such interactions occur on multiple
More informationClimate change: How do we know?
Climate change: How do we know? This graph, based on the comparison of atmospheric samples contained in ice cores and more recent direct measurements, provides evidence that atmospheric CO2 has increased
More informationEcological Succession
Ecological Succession Primary succession will ALWAYS be on areas where there is NO SOIL a) bare rock from a retreating glacier, b) newly cooled lava; c) abandoned lot with an impervious surface (parking
More informationPee Dee Explorer. Science Standards
Science Standards About Pee Dee Explorer What does it mean when someone says they are from the "Pee Dee" of South Carolina? A place is bigger than its physical geography. A "sense of place" weaves together
More informationName Hour. Section 4-1 The Role of Climate (pages 87-89) What Is Climate? (page 87) 1. How is weather different from climate?
Name Hour Section 4-1 The Role of Climate (pages 87-89) What Is Climate? (page 87) 1. How is weather different from climate? 2. What factors cause climate? The Greenhouse Effect (page 87) 3. Circle the
More informationGlacial Modification of Terrain
Glacial Modification Part I Stupendous glaciers and crystal snowflakes -- every form of animate or inanimate existence leaves its impress upon the soul of man. 1 -Orison Swett Marden Glacial Modification
More informationEssential Questions. What factors are most significant in structuring a community?
Community Ecology Essential Questions What factors are most significant in structuring a community? What determines a communities species composition and the relative amount of species present? What is
More informationEssential Questions. What is erosion? What is mass wasting?
Erosion Essential Questions What is erosion? What is mass wasting? What is Erosion? Erosion The transportation of sediment from one area to another Caused mainly by running water but also caused by glaciers,
More informationCo-invasion of invasive trees and their associated belowground mutualists
Co-invasion of invasive trees and their associated belowground mutualists Martin. A. Nuñez*, Nahuel Policelli & Romina Dimarco *Grupo de Ecologia de Invasiones INIBIOMA, CONICET/U. del Comahue, Argentina
More informationSUCCESSION Community & Ecosystem Change over time
Schueller NRE 509: Lecture 23 SUCCESSION Community & Ecosystem Change over time 1. Forest study revisited 2. Patterns in community change over time: 3 cases 3. What is changing? 4. What determines the
More informationHow does the greenhouse effect maintain the biosphere s temperature range? What are Earth s three main climate zones?
Section 4 1 The Role of Climate (pages 87 89) Key Concepts How does the greenhouse effect maintain the biosphere s temperature range? What are Earth s three main climate zones? What Is Climate? (page 87)
More informationAP Environmental Science I. Unit 1-2: Biodiversity & Evolution
NOTE/STUDY GUIDE: Unit 1-2, Biodiversity & Evolution AP Environmental Science I, Mr. Doc Miller, M.Ed. North Central High School Name: ID#: NORTH CENTRAL HIGH SCHOOL NOTE & STUDY GUIDE AP Environmental
More informationAbout me (why am I giving this talk) Dr. Bruce A. Snyder
Ecology About me (why am I giving this talk) Dr. Bruce A. Snyder basnyder@ksu.edu PhD: Ecology (University of Georgia) MS: Environmental Science & Policy BS: Biology; Environmental Science (University
More informationWhat is environmental science? Rehash, Lecture 1: Rehash, Lecture 1: Environmental Issues & Problems ENV 150. Natural Capital Degradation
Environmental Issues & Problems ENV 150 Guillaume Mauger What is environmental science? Today s Schedule: 1. Review Syllabus, Schedule, Assignments 2. Rehash of last week s topics 3. Biodiversity and Rehash,
More informationEnvironmental Science
Environmental Science A Study of Interrelationships Cui Jiansheng Hebei University of Science and Technology CH06 Kinds of Ecosystems and Communities Chapter Objectives After reading this chapter, you
More informationFigure 2 If birds eat insects that feed on corn, which pyramid level in the diagram would birds occupy? 1. A 3. C 2. B 4. D
Ecology Week 1 Assignment. This week's assignment will count as a quiz grade. Please speak to Mr. Roes about any questions that you would like help on! 1. The fact that no organism exists as an entity
More informationECOLOGICAL SUCCESSION. Prof :DEEPAK SAINI HOD ZOOLOGY J.C.D.A.V. College,Dasuya
ECOLOGICAL SUCCESSION Prof :DEEPAK SAINI HOD ZOOLOGY J.C.D.A.V. College,Dasuya Primary succession: The gradual establishment, through stages, of a climax ecosystem, that has not been occupied before. Primary
More informationMycorrhizal dependence and growth habit of warm-season and cool-season tallgrass prairie plants
Mycorrhizal dependence and growth habit of warm-season and cool-season tallgrass prairie plants B. A. Daniels Hetrick, D. Gerschefske Kitt, G. Thompson Wilson Canadian Journal of Botany, 1988, 66(7): 1376-1380,
More informationSection 8. North American Biomes. What Do You See? Think About It. Investigate. Learning Outcomes
Section 8 North American Biomes What Do You See? Learning Outcomes In this section, you will Define the major biomes of North America and identify your community s biome. Understand that organisms on land
More informationD. Adaptive Radiation
D. Adaptive Radiation One species new species: A new species: B new species: C new species: D Typically occurs when populations of a single species... invade a variety of new habitats, evolve under different
More information10/6/ th Grade Ecology and the Environment. Chapter 2: Ecosystems and Biomes
7 th Grade Ecology and the Environment Chapter 2: Ecosystems and Biomes Lesson 1 (Energy Flow in Ecosystems) Each organism in an ecosystem fills an energy role. Producer an organism that can make its own
More informationEcology Review. 1. Fly larvae consume the body of a dead rabbit. In this activity, they function as
Name: ate: 1. Fly larvae consume the body of a dead rabbit. In this activity, they function as. producers. scavengers. herbivore. parasites 4. n earthworm lives and reproduces in the soil. It aerates the
More informationFundamental ecological principles
What Important Ideas Will Emerge in Your Study of Ecology? Fundamental ecological principles Application of the scientific method to answer specific ecological questions Ecology is a quantitative science
More informationEcosystems. 1. Population Interactions 2. Energy Flow 3. Material Cycle
Ecosystems 1. Population Interactions 2. Energy Flow 3. Material Cycle The deep sea was once thought to have few forms of life because of the darkness (no photosynthesis) and tremendous pressures. But
More informationMetacommunities Spatial Ecology of Communities
Spatial Ecology of Communities Four perspectives for multiple species Patch dynamics principles of metapopulation models (patchy pops, Levins) Mass effects principles of source-sink and rescue effects
More informationUnderstory loss, reduced diversity
Understory loss, reduced diversity Characteristics of an Urban Ecosystem Fauna Generalists Predators Non-native Species Monk Parakeets Congress Avenue Bridge 1.5-2 Million MexicanFree Free-tailed Bats
More informationBiogeochemical cycles
Lecture -2: Biogeochemical cycles ENV 107: Introduction to Environmental Science Dr. A.K.M. Saiful Islam Case Study: Lake Washington The city of Seattle, USA lies between two major bodies of water- saltwater
More informationSLOSS debate. reserve design principles. Caribbean Anolis. SLOSS debate- criticisms. Single large or several small Debate over reserve design
SLOSS debate reserve design principles Single large or several small Debate over reserve design SLOSS debate- criticisms Caribbean Anolis Pattern not always supported Other factors may explain diversity
More informationThe State of the cryosphere
The State of the cryosphere Course outline Introduction The cryosphere; what is it? The Earth; a unique planet Cryospheric components Classifications Lecture outlines The State of the cryosphere The State
More informationEarth s Major Terrerstrial Biomes. *Wetlands (found all over Earth)
Biomes Biome: the major types of terrestrial ecosystems determined primarily by climate 2 main factors: Depends on ; proximity to ocean; and air and ocean circulation patterns Similar traits of plants
More informationUNIT 5: ECOLOGY Chapter 15: The Biosphere
CORNELL NOTES Directions: You must create a minimum of 5 questions in this column per page (average). Use these to study your notes and prepare for tests and quizzes. Notes will be stamped after each assigned
More informationChapter 6, Part Colonizers arriving in North America found extremely landscapes. It looked different to region showing great.
Social Studies 9 Unit 1 Worksheet Chapter 6, Part 1. 1. Colonizers arriving in North America found extremely landscapes. It looked different to region showing great. 2. The Earth is years old and is composed
More informationBIOGEOCHEMICAL CYCLES
BIOGEOCHEMICAL CYCLES BASICS Biogeochemical Cycle: The complete path a chemical takes through the four major components, or reservoirs, of Earth s system (atmosphere, lithosphere, hydrosphere and biosphere)
More informationHW/CW #5 CHAPTER 3 PRACTICE
HW/CW #5 CHAPTER 3 PRACTICE 1. The portion of Earth in which all life exists is known as A) the climax stage B) the biosphere C) a population D) a biotic community 2. The study of the interactions between
More informationDynamic and Succession of Ecosystems
Dynamic and Succession of Ecosystems Kristin Heinz, Anja Nitzsche 10.05.06 Basics of Ecosystem Analysis Structure Ecosystem dynamics Basics Rhythms Fundamental model Ecosystem succession Basics Energy
More informationEcology - the study of how living things interact with each other and their environment
Ecology Ecology - the study of how living things interact with each other and their environment Biotic Factors - the living parts of a habitat Abiotic Factors - the non-living parts of a habitat examples:
More informationHow do glaciers form?
Glaciers What is a Glacier? A large mass of moving ice that exists year round is called a glacier. Glaciers are formed when snowfall exceeds snow melt year after year Snow and ice remain on the ground
More informationThe Prokaryotic World
The Prokaryotic World A. An overview of prokaryotic life There is no doubt that prokaryotes are everywhere. By everywhere, I mean living in every geographic region, in extremes of environmental conditions,
More informationQuantum Dots: A New Technique to Assess Mycorrhizal Contributions to Plant Nitrogen Across a Fire-Altered Landscape
2006-2011 Mission Kearney Foundation of Soil Science: Understanding and Managing Soil-Ecosystem Functions Across Spatial and Temporal Scales Progress Report: 2006007, 1/1/2007-12/31/2007 Quantum Dots:
More informationREADING GUIDE CHAPTERS 3-4. Name Class Date
READING GUIDE CHAPTERS 3-4 Name Class Date Levels of Organization 1. Come up with a thinking map or other visual representation that shows the relationships between the terms BIOSPHERE, ECOSYSTEM, COMMUNITY,
More informationUsing Soil Microbes to Enhance Restoration of Native FL Scrub. Ben Sikes University of Texas at Austin
Using Soil Microbes to Enhance Restoration of Native FL Scrub Ben Sikes University of Texas at Austin Talk Outline The role of soil biota in ecosystem processes and plant Current uses of soil microbes
More informationTrophic and community ecology
Trophic and community ecology Top carnivore Trophic levels Carnivore Herbivore Plant Trophic ecology Trophic related to feeding Autotrophs: synthesize their food Heterotrophs: eat other organisms Trophic
More informationWhat determines: 1) Species distributions? 2) Species diversity? Patterns and processes
Species diversity What determines: 1) Species distributions? 2) Species diversity? Patterns and processes At least 120 different (overlapping) hypotheses explaining species richness... We are going to
More informationExtent of Periglacial = Global Permafrost Permafrost: Soil and/or rock where temperatures remain below 0 degrees C for 2 or more years.
Geog 1000 - Lecture 34 Periglacial Environments and Paleoclimatology http://scholar.ulethbridge.ca/chasmer/classes/ Today s Lecture (Pgs 422-434) 1. Exam questions from last week, and today 2. Extent of
More informationCh.5 Evolution and Community Ecology How do organisms become so well suited to their environment? Evolution and Natural Selection
Ch.5 Evolution and Community Ecology How do organisms become so well suited to their environment? Evolution and Natural Selection Gene: A sequence of DNA that codes for a particular trait Gene pool: All
More informationFig.1. The largest moss peat bank on Signy Island. There is a metre rule visible in the middle distance (1975). All pictures by James Fenton
ANTARCTIC MOSS PEAT: MY WORK AS A BAS BOTANIST IN THE 1970s by James Fenton I graduated with a degree in botany from the University of Durham in 1973 and my first job was working for the British Antarctic
More informationATOC OUR CHANGING ENVIRONMENT
ATOC 1060-002 OUR CHANGING ENVIRONMENT Class 22 (Chp 15, Chp 14 Pages 288-290) Objectives of Today s Class Chp 15 Global Warming, Part 1: Recent and Future Climate: Recent climate: The Holocene Climate
More informationRainforest Ecosystems
Rainforest Ecosystems Ecosystems: A Brief Review Collection of interdependent parts Environment provides inputs Ecosystem produces outputs Hydrosphere Atmosphere Lithosphere Inputs Abiotic Inputs Energy
More informationEvolution Problem Drill 09: The Tree of Life
Evolution Problem Drill 09: The Tree of Life Question No. 1 of 10 Question 1. The age of the Earth is estimated to be about 4.0 to 4.5 billion years old. All of the following methods may be used to estimate
More informationOutline 23: The Ice Ages-Cenozoic Climatic History
Outline 23: The Ice Ages-Cenozoic Climatic History Continental Glacier in Antarctica Valley Glaciers in Alaska, note the moraines Valley Glaciers in Alaska, note the moraines Mendenhall Glacier, Juneau,
More informationInteractions Between Microorganisms and Higher Plants from Competition to Symbiosis p. 184
Introduction What Are Soils? p. 3 Introduction p. 3 Soil Genesis p. 4 Rock Weathering or Decay p. 4 Importance of Soil Texture p. 5 Input of Organic Matter into Soils and Aggregation p. 7 Migration Processes
More informationLecture 2 Carbon and Energy Transformations
1.018/7.30J Fall 2003 Fundamentals of Ecology Lecture 2 Carbon and Energy Transformations READINGS FOR NEXT LECTURE: Krebs Chapter 25: Ecosystem Metabolism I: Primary Productivity Luria. 1975. Overview
More informationModule 4: Community structure and assembly
Module 4: Community structure and assembly Class Topic Reading(s) Day 1 (Thu Intro, definitions, some history. Messing Nov 2) around with a simple dataset in R. Day 2 (Tue Nov 7) Day 3 (Thu Nov 9) Day
More informationCh20_Ecology, community & ecosystems
Community Ecology Populations of different species living in the same place NICHE The sum of all the different use of abiotic resources in the habitat by s given species what the organism does what is
More informationChapter 6 Reading Questions
Chapter 6 Reading Questions 1. Fill in 5 key events in the re-establishment of the New England forest in the Opening Story: 1. Farmers begin leaving 2. 3. 4. 5. 6. 7. Broadleaf forest reestablished 2.
More informationFind this material useful? You can help our team to keep this site up and bring you even more content consider donating via the link on our site.
Find this material useful? You can help our team to keep this site up and bring you even more content consider donating via the link on our site. Still having trouble understanding the material? Check
More informationCommunity Ecology. PowerPoint Lecture Presentations for Biology Eighth Edition Neil Campbell and Jane Reece
Chapter 54 Community Ecology PowerPoint Lecture Presentations for Biology Eighth Edition Neil Campbell and Jane Reece Lectures by Chris Romero, updated by Erin Barley with contributions from Joan Sharp
More informationWednesday, November 15, 2017
Wednesday, November 15, 2017 Northern Europe: Physical Geography Objective: Locate and describe the various traditional regions of Western Europe. Outline how the physical geography varies from region
More informationAP Biology Ecology Review and Study Guide
AP Biology Ecology Review and Study Guide 2017-2018 1. Which pattern represents the predator, which the prey? 2. Give two justifications for your selection: 3. Describe the population dynamics of A: 1.
More informationPhylogenetic diversity and conservation
Phylogenetic diversity and conservation Dan Faith The Australian Museum Applied ecology and human dimensions in biological conservation Biota Program/ FAPESP Nov. 9-10, 2009 BioGENESIS Providing an evolutionary
More informationSOIL: DEFINITION, FORMATION! & LAYERS"
SOIL: DEFINITION, FORMATION & LAYERS" What Is Soil? soil - upper-most (relatively thin) layer of Earth s crust, which supports terrestrial plants, animals, & microorganisms basic natural resource that
More informationOutline Classes of diversity measures. Species Divergence and the Measurement of Microbial Diversity. How do we describe and compare diversity?
Species Divergence and the Measurement of Microbial Diversity Cathy Lozupone University of Colorado, Boulder. Washington University, St Louis. Outline Classes of diversity measures α vs β diversity Quantitative
More informationChapter 2: Physical Geography
Chapter 2: Physical Geography Pg. 39-68 Learning Goals for Chp2: q q q q q Explain how the Earth moves in space and why seasons change. Outline the factors that influence climate and recognize different
More information