Lecture 2 and text (pg. 1,2,12-14) 1. What are some properties of life? Adaptations Growth and Development Order - ordered structure Energy processing Reproduction Regulation of body processes Response to environment Evolutionary adaptation 2. Evolution is the "overarching theme in biology". 3. Why do biologists pose questions and then try to answer them, rather than simply setting out to "study" some phenomenon? By asking questions, forming hypotheses, we have a focused topic that can be experimented on. 4. What is a hierarchical system of classification? Linnean classification is a kind of taxonomy which names individuals based upon levels of similarity. The following are also examples of classification: Taxonomy, Emergent Properties, Levels of organisms and life on earth 5. The terms Ursus and genus (Fig. 1.14) are not synonymous but they are linked. How are they similar and how are they different? Genus is the classification name, Ursus is a one of those names and is a genus 6. What are the three domains of life? Archaea, Bacteria, Eukarya 7. What is the difference between a prokaryotic and a eukaryotic cell (check your glossary)? Eukaryotes have a nucleus enclosed in a membranous sac 8. What are the four traditional kingdoms of eukaryotes? Which one is now thought to comprise numerous kingdoms? Protists are now numerous Anamalia Plantae Fungi Protists Things to think about Think of this course as the diversity and UNITY of life However, we consider viruses as non-living because they cannot replicate on their own Insects are arthropods, which is the largest group of living organisms whereas Mammals are a relatively inconsequential and diverse grouping in terms of size/numbers Two complementary approaches to our studies •Diversity of Life •Unity of Life Classification Classification shows connections/unity through similarities and diversity through differences Characters are any attribute which is considered separately from the whole organism for the purpose of comparison, identification, or interpretation Character/ expression of character can be used to find the state. For example, our hand has the character of digits. The state of fingers is yes or no (either you have fingers or you don't). The state is usually four fingers, one thumb. Characters are chosen by distinction - characters vary. Groups We define groups in different ways. Hierarchical Grouping - you can have groups within groups Can be shown through tree diagram Hierarchical Systems Aristotle - species as fixed and arranged on scala naturae - classified animals into two groups Blood - born Bloodless - insects shelled animals *Now called vertebrate and invertebrate Aristotle's Basic Questions What is the vital principle (life)? Is it the same for all creatures? What is required to maintain life (physiology)? How does "like" beget "like"? What is responsible for the diversity of creatures? What can account for the similarity among creatures? Linnaneus Used artificial classification Binomial Nomenclature - all species have a latin name of genus and epithet (adjective) or species Linnaeus - Species Plantarum Post Linnaean Systems Strove to be "natural" systems based on overall similarity Lecture 3 (453, 537-538,1246-1250) 1. How did Aristotle view biodiversity? Life-forms could be arranged on a ladder of increasing complexity: the scala natura or scale of nature. 2. What is binomial nomenclature and how does it relate to Linnaeus's views of classification? Linnaeus grouped similar species into increasingly general categories. Similar species were grouped into a similar genus, but had their own specific epithet, making a binomial name. 3. What is a taxon? The named taxonomic unit at any level of the hierarchy 4. What components make up a species' scientific name? Genus and specific epithet 5. Provide an example of a category, as the term is used in Linnaean classification. Kingdom, phylum, class, order, family, genus, and species are categories 6. What are the three main levels at which biological diversity can be described? genetic, species, ecosystem 7. Identify three main threats to biodiversity. 8. Describe an example of a benefit to humans derived from biodiversity 9. What is meant by the term ecosystem services? Modern Views of Biodiversity Current Hierarchical System Domain Kingdom Phylum Class Order Family Genus Species P P E Binomial Name Both genus and epithet and in the species name eg) Canis lupus - wolf Zea mays - corn Hierarchical Systems can be shown as tree diagrams Darwin and the Galapagos Recent volcanic origin - important bcuz species on island came there by dispersal, islands were never connected to another land mass Most species were unique Most were similar to species in South America - consistent with idea that species on Galapagos got there from long distance dispersal from nearest land mass Finches showed diversity based on different islands and habitats - spatial isolation supported Darwin's idea of the roles in natural selection and adaptation in evolution What is Biodiversity? Genetic Diversity - differences in DNA and traits Species Diversity - human diversity Ecosystem Diversity - differences in climates Loss of Biodiversity Genetic diversity Genetic variation is diminished with population extinction Species diversity Numerous species endangered or threatened Species loss has escalated dramatically in the recent past Ecosystem Diversity Highlights interactions among species and environment Some keystone species in danger of extinction (pollinators) Some habitats are disappearing quickly (wetlands, riparian systems) Why preserve biodiversity? Species and genetic diversity o biophilia o plants are sources of many economic products including medicines o Taq polymerase example Ecosystem services o The notion that we benefit from proper operation of the ecosystems we inhabit o Example of New York City Loss of Biodiversity Genetic variation is diminished with population extinction Numerous species endangered or threatened Species loss has escalated dramatically in the recent past Highlights interactions among species and the environment Some keystone species in danger of extinction (pollinators) Some habitats are disappearing quickly Why preserve biodiversity? Species and genetic diversity - Biophilia Humans like diversity and feel it is a tragedy to lose species - Plants are the sources of many economic products including medicine Taq polymerase - the enzyme that allows the polymerase chain reaction to occur for DNA replication Ecosystem Services Lecture 4 (14-18, 452-458) 1. Evolution is the process that best explains the unity of life. 2. The character that reflects the overall unity of life is DNA. 3. In Darwin's view, the source of the selective force acting on populations over many generations was Natural Selection. 4. How are the broad sense and narrow sense definitions of evolution (pg. 452) directly related to each other? 5. How did the Linnaean system of classification support the theory of evolution? 6. Distinguish between catastrophism and uniformitarianism and identify the major proponents of each theory. 7. How were Lamarck's ideas of evolution different from our current understanding of the process? 8. Darwin's observations of South American species (both extant (living) and extinct (fossil)) provided important insights. What were his conclusions? 9. Darwin's observations of Galapagos species provided important insights. What were his conclusions? 10.Who was A.R. Wallace and what were his contributions to evolutionary biology? 11. Darwin adopted the analogy of a "tree" or branching diagram as a means to present evolutionary relationships. How do these trees show both descent and modification? The Greatest Threat to Biodiversity A. Habitat Loss agriculture, cities, industrial processes, global warming B. Introduced Species global travel, species freed from natural constraints Zebra mussels, purple loosestrife C. Overexploitation Over-harvesting Conservation Biology Protect genetic, species, and ecosystem diversity Darwin and Evolution Embraced idea of gradualism Geology - change over time - Gradualism and Uniformitariansim - earth changes slowly Hutton and Lyell - Catastrophism - periodic catastrophes explained evolution of the earth Cuvier - species occurred at one time then became extinct Lamark - biologist - species could change - inheritance of acquired characteristics - use and disuse Darwin Descent with Modification Natural Selection explains modification Darwin and the Galapagos •Recent volcanic origin •Most species were unique •Most were similar to species in South America •Finches showed diversity based on different islands and habitats Propinquity of Descent Kinship/ relationship: offspring look like parents Common Ancestry explains similarity Used a classification system to come up with this idea Hierarchy exists because of this Credits Linneaus - Characters do not make the genes, BUT genes make characters - Something more is included in this classification than mere resemblance Unity and Diversity of Life Descent with Modification is based on the 'Malthusian struggle' is similar to artificial selection Lecture 5 (458-461) 1. What is artificial selection and how does it work? 2. How does natural selection differ from artificial selection? 3. What contribution did Malthus make to Darwin's ideas on natural selection? 4. What were Darwin's four observations and two inferences, relating to his theory of natural selection? 5. How does natural selection result in adaptation? 6. What biological entity responds to natural selection? (i.e. what is it exactly that evolves over time?) 7. In the studies of guppy natural selection, two important selective forces were identified. What are they and how do they work? 8. What were the results/conclusions of the guppy study? 9. How long does it take for HIV resistance to evolve in a patient taking HIV drugs? Natural Selection - based on "Malthusian struggle" - similar process to artificial selection (practiced in agriculture, breeding) Overproduction of Offspring Occurs to sustain population Unchecked, over reproduction results in an extraordinary number Artificial Selection, Natural Selection, and Adaptation •Darwin noted that humans have modified other species by selecting and breeding individuals with desired traits, a process called artificial selection •Darwin then described four observations of nature and from these drew two inferences Darwin's Observations 1. members of a population often vary greatly in their traits 2. traits are inherited from parents to offspring 3. all species are capable of producing more offspring than the environment can support 4. owing to lack of food or other resources, many of these offspring do not survive Darwin's Inferences 1. Individuals whose inherited traits give them a higher probability of surviving and reproducing in a given environment tend to leave more offspring than other individuals 2. This unequal ability of individuals to survive and reproduce will lead to the accumulation of favorable traits in the population over generations see ch 1 review p. 25 population of organisms flow chart Biology 1020 Lecture 6 (468-471, 478-481, 483-484) 1. What is the smallest unit that evolves? Populations. Microevolution is evolutionary change below the species level; change in the allele frequencies in a population over generations. 2. Who discovered genetics and when? Mendel 1865, after Darwin's 1859 Origin of Species 3. What is a "cline" in geographical variation? A graded change in a character along a geographic axis. 4. Define mutation. A change in the nucleotide sequence of an organism's DNA, ultimately creating genetic diversity. 5. How does a point mutation differ from a gene duplication? 6. How does sexual reproduction generate variation among organisms? 7. How does gene flow between populations shape patterns of genetic variation among populations? 8. What is relative fitness, and why is it important? 9. Is relative fitness a characteristic of the gene, the genotype or the individual? Individual 10. Distinguish between directional, stabilizing and disruptive selection. 11. Describe two examples of balancing selection. 12. Why, in spite of natural selection, are organisms not perfectly adapted to their environment? Descent (Unity) with Modification (Diversity) Natural Selection explains diversity, therefore diversity would be maintained between populations. However, it would reduce variation within a population. Evolution - the change in the genetic composition of a population from generation to generation Arguments against Natural Selection 1. Darwin embraced Lamarckian inheritance which was disproved 2. Lacked any explanation of inheritance Darwin vouched for "blended" inheritance 3. Difficult to explain the evolution of complex characters Mendel and Modern Synthesis Mendel's work published in 1865 (Darwin's was same time, 1859) Hugo Devries rediscovered Mendel's work in 1900 Genetics is incorporated into evolutionary theory in the 1940s (Fisher and Wright) Genetic and Non-genetic Variation Biology 1020 Lecture 7 Text Reading questions (487-498) 1. What is the biological species concept? Definition of a species as a population or group of populations whose members have the potential to interbreed in nature and produce viable, fertile offspring, but do not produce viable, fertile offspring with members of other such groups. It highlights the importance of gene flow as a unifying force 2. What is the importance of gene flow in the evolution of species? Diversity and variation 3. Reproductive isolating mechanisms can be classified into two groups. What are the two groups? 4. What are the kinds of isolating mechanisms in each group? 5. Why is the presence of Allele 1 in Population B (Fig. 24.3) evidence of a gene flow event? 6. What are two limitations of the BSC (biological species concept)? 7. What are three alternative species concepts? How do they differ from one another? 8. Distinguish between the two main speciation models (allopatric vs sympatric). Allopatric Speciation - geographical isolation Sympatric Speciation - same country 9. What is the evidence that reproductive isolation evolves as a by product of divergence, rather than via direct selection for the trait itself? 10.What is polyploidy and how does it result in sympatric speciation? 11.What are two other factors that may lead to sympatric speciation? Species - A population or group of populations whose members have the potential to interbreed in nature and produce viable, fertile offspring, but do not produce viable, fertile offspring with members of other such groups. Speciation - an evolutionary process in which one species splits into two or more species. Microevolution - evolutionary change below the species level; change in the allele frequencies in a population over generations. Macroevolution - evolutionary change above the species level, including the origin of a new group of organisms or a shift in the broad pattern of evolutionary change over a long period of time. Examples of macroevolutionary change include the appearance of major new features of organisms and the impact of mass extinctions on the diversity of life and its subsequent recovery. Population - group of individuals that interbreed -- gene flow occurs Cline - a graded change in a character along a geographic axis. Selection requires genetic variation How does genetic variation arise? - mutation o point mutations o macromutations like mouse example where chromosomes change doubling genome we have 46 chromosomes - sexual reproduction o mixes genes into new combinations to maintain diversity Gene Flow affects variation patterns within and between populations To prevent gene flow in plants near or further from a mine, a wind breaker could be set up. Copper tolerance is important in plants nearest the mine. Natural Selection From a populational standpoint When we look at a phenotype, we can describe them quantitatively to show the numbers expressing each trait. Directional selection - selection of one or the other extreme of a trait either white or black color) cows that produce most milk Disruptive selection - selects against the mean of a trait Stabilizing selection - selects for the mean - the average is the best and reduces the extremes Natural Selection Results in Adaptation The Preservation of Genetic Variation - Diploidy o One haploid genome may compensate for harmful mutations in other o Heterozygote advantage - Balancing Selection o maintains two or more phenotypic forms in a population. - Frequency dependent o decline in the reproductive success of individuals that have a phenotype that has become too common in a population The existence of anti-malarial drugs: Increase disadvantage of heterozygote Heterozygotes are malaria resistant The sickle-cell allele is most common Adaptation is imperfect 1) works on existing variation 2) historical constraints a. different groups have different traits b. design based on what was there before 3) Time lag a. Environment changes first, characters change after 4) Trade-offs a. compromises b. A character may be functioning in two different ways that may be contradictory c. Call of the frog attracts mates, but also notifies predators therefore the mating call is also bad 5) Chance a. Cataclysmic events b. Plays a role in evolution Biology 1020 - Text Questions Lecture 8 Hybridization (498-504) 1. What is a hybrid zone? 2. What are the possible outcomes of a hybrid zone once it has formed (Fig. 24.14) 3. What is reinforcement and why does it occur? 4. What factors might promote fusion of two species? 5. What factors might promote the formation of a persistent hybrid swarm? 6. What is the average time required for speciation to occur? What is the fastest time recorded? 7. Does speciation require divergence among a large number of different genes or can it occur with minimal genetic divergence? Two patterns of evolutionary change (1) Anagenesis - populations replace each other over time (2) Cladogenesis - divergence: change that involves a branching path Speciation causes divergence Species Concept How species are looked at Biological Species Concept Morphological species concept - species are fundamentally the same Phylogenetic species concept - branch of a tree Ecological species concept - species occupies unique niche Reproductive Isolation Mechanisms Prezygotic Barriers Postzygotic Isolation Example: donkeys and horses interbreed but have infertile offspring Habitat Isolation Two species never interbreed or exchange genes because they have different habitats Temporal Isolation Reproductive patterns separated by time Behavioral Isolation Based upon mate selection Example: mating dances in birds Mechanical Isolation Genetalia do not align correctly Gametic Isolation Example: sea urchins project spores/ gametes into water Postzygotic Barriers Reduced hybrid viability Reduced hybrid fertility Example: donkey and horse produce a mule Barrier to gene flow Hybrid breakdown First hybrid is viable but generations become weaker Gene Flow Allopratric Speciation - physical separation Isolating mechanism is geographic Sympatric Speciation - even though gene flow is possible, some factor prevents Habitat differentiation Polyploidy Sexual Selection Gene flow is a barrier to sympatric speciation Biology 1020 - Lecture 9 (461-466, 536-540) 1. What two major evolutionary phenomena are well documented in the fossil record? Extinction and change 2. How does the fossil record provide a means of testing evolutionary hypotheses? Age of different fossils = what evolved from what 3. Define homology? Similarity in characteristics resulting from a shared ancestry. 4. Why do homologies show a nested pattern? 5. How can ontogeny (=developmental patterns) provide evidence of evolutionary relationships? 6. How does the evolution of analogous structures by convergent evolution differ from the evolution of homologous structures? 7. How does our understanding of continental drift provide evidence supporting evolution? 8. What is a phylogeny? the evolutionary history of a species or a group of species 9. What are sister groups (=sister taxa) in a phylogeny? 10. How is classification linked to phylogenetic trees? branching diagram that represents a hypothesis about the evolutionary history of a group of organisms Sympatric Speciation Polyploidy - doubling of the genome Autopolyploidy Doubling of the chromosomes Failure of cell division after chromosome duplication gives rise to tetraploid tissue The gametes that are produced are diploid or 2n Offspring with tetraploid karyotypes may be viable with others of this kind and fertile and would have 4n genome Allopolyploidy Doubling of the chromosomes More common, although more complex than autopolyploidy Functions as a gamete with twice as many chromosomes New species is a hybrid Allopolyploidy Example - Fast speciation event Species A: 2n = 6 Normal gamete: n = 3 Species B: 2n = 4 Unreduced gamete: n = 4 Hybrid Species 2n = 10 Habitat Differentiation Often relates to host shift The environment of the parasite is a host, sometimes it can shift from one host to another host. For example: parasite moving from hawthorne tree to apple tree. The individuals on the apple tree will interbreed. The parasites on the two trees are ABLE to interbreed because gene flow is possible. However, the habitats now differ. Sexual Selection Mate selection Female chooses a specific type of male Gene flow is possible, but females choose against it For example, female could chose different appearance, causing divergence Possible Hybrid Outcomes for species: Reinforcement o hybridization is disadvantageous oMore difference between sympatric populations than allopatric ones oPre-zygotic isolation mechanism in overlapping areas to prevent hybrids oHybrid zone lessens Fusion o Potential in a hybrid zone where convergence occurs back to one species Stability o Stable Hybrid Swarm - 3 species involved o status quo relatively maintained Hybrids could backcross with their parents and dilute the parental gene pools. This implies the ability of selection into the future. Why might reproductive barriers.? If the hybrids are unfit or sterile, they only have so much reproductive potential. This is a disadvantage. Hybrids reduce the reproductive capacity of their parents. You would expect pre-zygotic isolating mechanisms to be: More common between sympatric species pairs Because hybridization can only occur within sympatric species pairs How fast can speciation occur? 6 million years average But speciation CAN happen very fast Patterns of Divergence in the Fossil Record Provides evidence of speciation and evolution (a) punctuated pattern a. quickly at the beginning, becomes stable (b) gradual pattern a. change is slow Biology 1020 - Lecture 10 text questions (540-548) 1. •What is a homoplasy? Similar (analogous) structure or molecular sequence that has evolved independently in two species. 2. •How/why do homoplasies evolve? Convergent evolution 3. •What is a clade? A group of species that includes an ancestral species and all its descendants. 4. •Distinguish among mono, para and polyphyletic groups. Monophyletic: group of taxa that consists of a common ancestor and all its descendants. A monophyletic taxon is equivalent to a clade. Paraphyletic: group of taxa that consists of a common ancestor and some, but not all, of its descendants. Polyphyletic: group of taxa derived from two or more different ancestors. 5. •What is the difference between a shared ancestral homology and a shared derived homology? 6. •What is an outgroup? What is an ingroup? 7. •What information does an outgroup provide in phylogenetic analyses? 8. •What is the principle of parsimony? What information does it provide in phylogenetic analyses? The principle states that when considering multiple explanations for an observation, one should first investigate the simplest explanation that is consistent with the facts. Phylogenies are a simple as possible 9. •What is the principle of maximum likelihood? States that when considering multiple phylogenetic hypotheses, one should take into account the hypothesis that reflects the most likely sequence of evolutionary events, given certain rules about how DNA changes over time. 10. •What is phylogenetic bracketing? On what principle is it based? Anagenesis: speciation over time without branching Cladogenesis: speciation with branching Speciation builds the tree Common ancestor - > speciation - > divergence Homology How do we recognize homology? Similarity in characteristics resulting from shared ancestry Similar structure, same ancestor If it is a complex structure and still similar, it is likely homologous (underlying bone structure of limb in human, cat, whale, and bat) Similarity = homology 1. Complex structures 2. Development 3. Genes Developmental processes In early stages of development of an organism, we see shared characteristics that are not observed in adults. This suggests developmental processes have evolved only in later stages and therefore later stages are more derived (early stages are conserved). Human embryo and chick embryo are similar with pharyngeal pouches and post-anal tail. Since humans and chickens have similar developmental processes and underlying bone structure, they are likely homologous Ontogeny recapitulates Phylogeny Development (repeat evolutionary stages as embryo) Phylogeny Embryological homology Analogy Similarity resulting from convergent evolution Analogy: wing of bats and birds Common ancestor did not have wings Wings evolved at different times and after divergence from ancestry Flying is a result of convergent evolution Homology: forelimb of bats and birds Far back ancestor had a forelimb before divergence occurred Analogy: Fish with fins and Whale has a swimming forelimb Phylogenic Trees In constructing a phylogenic tree, we ignore analogies There is one true evolutionary tree of the history of life forms on Earth. The history predates humans, so it is hard to know it is true. How do we find it? Our classification should reflect it TREE TERMINOLOGY Common Ancestor is represented by a node of the tree We construct the tree to be dichotomous A polytomy is more than two branches from a node A taxon - species, genus, or family depending on what level you are constructing your tree Sister taxa - arise from the same common ancestor and are therefore each other's closest relative Molecular Homologies can be very detailed Tree Constuction - based on two main concepts: (1) The principle of parsimony based on the assumption that evolution proceeds by a smaller rather than a larger number of events Occam's Razzor way of reasoning when have little information where you don't know, go for the simplest hypothesis/ fewer steps Can be used to construct a tree with characteristics Can demonstrate evolution reasonably and helps us choose the best tree (2) The use of shared derived homologies Ancestral trait that changed into a derived trait Homology must be traceable - talking about same character Derived Homologies are important because they represent change Change is what defines a phylogenic tree How do we know what is derived? Fossil evidence - more complete the fossil record is, the better a source Outgroup Comparisons Sister group of the Ingroup, assume outgroup has ancestral trait Use parsimony - the simplest way for a trait to have been derived Construction of Phylogenetic Trees Choose a study group - ingroup Designate the outgroup - the closest relative Choose characters (homologies) Polarize the characters (ancestral vs. derived through outgroup comparisons) Score the characters (A, B, C, ) Construct the most parsimonious tree If you are building a phylogeny of cats in the broad sense, you need an outgroup that is NOT in the ingroup as a comparison to the felines Ingroup: lion, tiger, leopard, domestic cat Outgroup: wolf How do we translate phylogeny into classification? Monophyletic - We look mostly for monophyletic groups based upon shared derived characters - common ancestor and all of its descendants Paraphyletic - shared ancestral homologies, not consistent with the tree (if you have to cut away twice to separate the group) - common ancestor and some descendants Polyphyletic - based on shared analogies, not what we want to classify on - does not include common ancestor Fundamental Questions on the Origin of Life How could complex organic molecules form abiotically? How could metabolism and replication evolve simultaneously? How could cell membranes develop? How could all of this happen so fast? Life arose about 3 to 4 billion years ago When? • Earth formed about 4.6 bya (billion years ago). • BUT: Planet continued to be bombarded by asteroids and comets for at least 500 million years. • Evidence suggests life originated at least 3.8 bya. • Either life originated fairly quickly (< 500 million years) or it originated in "Hadean" conditions. cyanobacteria - photosynthetic bacteria in a filamentus form 3.4-3.5 billion years ago evidence of Bacteria we believe that something came before these bacteria because photosynthesis was not the first source of energy and there are simpler forms How? • Abiotic synthesis of small organic molecules such as amino acids and nucleotides - provided building blocks of life. • Joining of these small molecules into macromolecules (proteins and nucleic acids) • Packing of these molecules inside a membrane. • Origin of self-replicating molecules. Abiotic synthesis on earth • Miller-Urey experiment in 1953 showed that organic molecules could be produced abiotically, under conditions plausible for the early earth. • Early atmosphere may have been composed of methane, hydrogen, water vapour, and amonia. Organic molecules can be created from these compounds. Abiotic synthesis in space • Evidence from meteorites shows organic synthesis is common in space. Meteorites contain carbon. • Murchison Meteorite showed bases found in nucleosides, and also amino acids. • Found L forms of amino acids slightly more common than D - basis for homochirality. They are mirror forms of each Living cells only use the L-form • THEORY: Organic molecules may have rained down on earth during its early history. What conditions? Conditions that favour the formation of molecules essential to life on Earth • Many different hypotheses because different organic molecules seem to require different conditions for abiotic synthesis. • Boiling temperatures aid synthesis of cytosine and uracil. • Freezing conditions aid the synthesis of adenine and guanine. • Freezing may also aid polymerization of RNA - a macromolecule Joining of these small molecules into macromolecules (proteins and nucleic acids) Polymer formation • Generally polymerization is energetically unfavorable reaction. • Some conditions may have favored polymerization: - On a drying clay surface. - Using energy from redox reactions, perhaps using high energy sulphur compounds from volcanic processes. A problem • Proteins require DNA as a template for synthesis. • DNA requires proteins as a catalyst for synthesis. • How could either be formed without the other? So which came first? Neither! RNA came first because it can do both jobs. RNA World Hypothesis • Ribosymes can act as both a template and as a catalyst, and therefore could potentially act as a catalyst for its own production - autocatalysis. Peptidyl transferase, a ribosyme which forms part of the ribosome Road to replication •Autocatalysis •Natural Selection A+B=C >C - the catalyst additional presence Packing of these molecules inside a membrane. Droplets and vesicles • Lots of evidence that a variety of lipids can spontaneously form small droplets with a bilayer at the surface. • These can be relatively stable and even split to form "daughters". • Phospholipid bilayers are highly impermeable to large organic molecules. • Simpler lipids such as fatty acids or monoglycerides are more permeable, and allow nucleotides and amino acids to enter. • Mansy et al. (2008) created simple vesicles with a single-stranded DNA template inside. Origin of self-replicating molecules. Where did the origin of life take place? Places where self-replicating molecules occurred • Drying coastal beaches or pools. • Deep sea vents. • Deep in the earth's crust. • In outer space - meteorite from Mars, where conditions for life at one time Three Domains of Life Domain Eukarya Domain Bacteria Domain Archaea - extreme environments single celled more closely related to Eukarya than other prokaryotes (bacteria) Great Moments in Evolution •Virtually all important metabolic pathways evolved in Bacteria and Archaea. •Photosynthesis - 2.7 billion years ago •Aerobic respiration - 2.1 billion years ago Evolution of Eukaryotes About 2.1 bya •Nuclear envelope •Membrane- bound organelles •cytoskeleton Closing thoughts • Experiments and fossil evidence has begun to suggest some plausible pathways for abiotic synthesis of monomers and polymers, and eventual evolution of protobionts. • The RNA Hypothesis has gained much more credibility based on Lincoln and Joyce. • By 2 bya, most important metabolic pathways had evolved, and life had diverged into 3 main branches. Sex and Multicellularity (More Great Moments in Evolution) Mitosis Meiosis Sexual life cycle with alternation of haploid and diploid stages Evolution of sex Sex provides more recombination, which spreads favorable mutations faster. • First step is evolution of mitosis. • Problem: Mitosis is a complex process: How could it evolve? Organs of extreme perfection To suppose that the eye with all its inimitable contrivances for adjusting the focus to different distances, for admitting different amounts of light, and for the correction of spherical and chromatic aberration, could have been formed by natural selection, seems, I freely confess, absurd in the highest degree. -- Charles Darwin "Reason tells me, that if numerous gradations from a simple and imperfect eye to one complex and perfect can be shown to exist, each grade being useful to its possessor... then the difficulty of believing that a perfect and complex eye could be formed by natural selection, though insuperable to our imagination, should not be considered as subversive of the theory." •Cytoskeleton evolved first in Eukaryotes, likely as ameoba-like locomotion. •Microtubule-organizing complexes (MOC) evolved initially to control flagellae. •MOC incorporated into nucleus for separation of chromatin. •MOC later function in both nucleus and controlling flagellae (not at the same time) •Evidence for this evolution based on examination of oddball single-celled Eukaryotes which don't have mitosis, or have non-standard mitosis. Gamete production • Isogamy - All gametes are identical. - May be flagellated or unflagellated • Anisogamy - Gametes differ in size or appearance - May be flagellated or unflagellated • Oogamy - Large unflagellatd egg and small flagellated sperm Gametic Meiosis Fig. 13.06a Sporic Meiosis Fig. 13.06b Zygotic Meiosis Fig. 13.06c Alternation of Generations • Characteristic of Sporic Meiosis • Two generations may be identical - Isomorphic • Two generations may look very different - Heteromorphic Benefits of diploid-dominance • Greater genetic variability within offspring (or gametes) because many more gametes are potentially produced from each zygote. Benefits of Haploid dominance • Faster growth of vegetative stage. • May reduce mixing of vertically transmitted (gamete to zygote) parasites. Benefits of Alternation of Generations • Haploid and diploid stages may be selected for different environments or functions. • But - what about isomorphic life cycles? Evolution of Multicellularity Benefits of Multicellularity • Recall our plucky little eukaryotic cell. • It's MOC can run flagellae or cilia giving the cell mobility, OR they can produce the spindles used in mitosis. But NOT both at the same time. • A colony of cells can get around this problem with cell specialization. Some cells specialize in motility, while others carry out mitosis. How? • Multicellular individuals seem to be "organs of extreme perfection." • Need to understand the transitional steps which allowed multicellularity to evolve. What about cheaters? • Multicellularity requires some cells to reduce or halt reproduction. • Cells that "cheat" will increase their own reproductive success, and so may be favored by natural selection. • Mechanisms to prevent cellular "cheating" are common, particularly in animals. E.g. germ- line sequestration, early determination of cell fate. Closing thoughts • Large innovations such as mitosis and multicellularity evolved through a series of small transitional steps, each providing a benefit. • The costs and benefits of different lifestyles and life cycles are likely specific to individual lineages, producing the high degree of variation we see in nature. • Sex likely evolved to provide greater recombination rates, perhaps as a way to combat parasitism. PROTISTS Pages 575-589 What are protists? • Eukaryotes which aren't animals, plants or fungi. Survey of the Protists Diversity Modes of Nutrition • Photoautotroph • (chemo) Heterotroph - Predators (Herbivores/Carnivores) - Parasites - Decomposers/Detritivores Modes of Locomotion • Cytoplasmic streaming or amoeboid motion • Flagellae or cilia Life Cycles • Gametic, Zygotic and Sporic Meiosis • Simple and complex (mainly parasites) • Sexual and Asexual SUPERGROUP EXCAVATA • Some members have grooves or "excavations" on their body Euglenozoans • Metabolically extremely diverse. • Some members are predatory or parasitic heterotrophs. Others are autotrophs • Some are mixotrophic - can shift between p/s and heterotrophy. - long flagellum eyespot short flagellum contractile vacuole nucleus chloroplast plasma membrane pellicle Chromalveolata Dinoflagellates Apicomplexans Aid in digestion Ciliates Phagocytosis and food vacuoles Oral groove Cell mouth Food vacuole Contractile vacuole Macronucleus Micronucleus Cilia Diatoms Brown Algae Dominant sporophyte (2N) Oomycetes Diploid (2N) Hyphae dominant Asexual Reproduction Zoosporangium Zoospore Cyst Germ tube Hyphae Sexual Reproduction Hyphae undergo Meiosis Oogonium contains egg nucleus (N) Antheridial hyphae with sperm nuclei (N) Fertilization Zygote Zygote germination Hyphae Closing Thoughts • Protists show tremendous diversity in structures and behavior. • While they may often go unnoticed in our day to day lives, they form critical parts of the biosphere (more on this next week). • The protists can be organized into five supergroups, but members of the groups are extremely diverse, and some groups are not universally recognized yet. SUPERGROUPS Pages 589-599 Excavata Euglenia - photosynthetic Chromalveolata Autotroph or photosynthetic Rhizaria Forams Pseudopodia projecting from shell are slender Slender openings in the shell from which pseudopodia extend Heterotroph Fossilize when sink to bottom of ocean We can use them as markers to date particular sedimentary layers Radiolarians Shelled amoeba Plasma membrane is outside of shell Slender pseudopodia Delicate shell made of silicon (like diatom shells) Archaeplastida Red Algae Can be single celled or multicellular Photosynthetic Most have cell walls made of cellulose Mostly chlorophyll a and accessory pigments No flagellated stage Red algae are red because Physiology: They contain accessory pigments that absorb blue and green light Phylogenetic: Their ancestors were red Functional: So they can live in deep water Green and blue light penetrate deeper through water Life Cycle Sporic meiosis Carposporophyte Also have a cystocarp which is an intermediate Tetrasporophyte Nori red algae are used to wrap sushi Land Plants Green Algae Chlorophytes Charophyceans Chlorophyll a and b Cell wall made of cellulose Large surface area: gases exchange or light collecting area Life Cycle Zygotic meiosis Some few examples of sporic meiosis Morphology can be single-celled colonial/ filamentous multicellular Unikonta Fungi Animals Plasmodial Slime Molds Cytoplasmic Streaming Similarity to fungus, are mobile through cytoplasmic streaming Live off of rotting vegetation or logs Live in damp terrestrial conditions Multinucleid Not multicellular One common cytoplasm that flows down the veins Plasmodium - feeding structure Like animals, Diploid is the adult part of the life cycle Gametic meiosis Spores are produced through meiosis Diversity of sporangium Life Cycle: Zygote (2N) Feeding Plasmodium Mature Plasmodium (preparing to fruit) Young sporangium Mature Sporangium on a stalk MEIOSIS Spores (N) Germinating spore -Flagellated cells (N) -Amoebid cells (N) FERTILIZATION Cellular Slime Molds Asexual Haploid dominant amoeba (N) Spend most of time as single-celled amoeba Spread spores Sexual Zygotic meiosis Diploid Zygote (2N) Produce haploid amoebas Cells secrete cAMP Gymnamoebas Naked amoeba Unshelled amoeba No cell wall, just a cell membrane Contractile valve Food vacuole Heterotrophic Feed by phagosytosis Move by cytoplasmic streaming which is done through cytoskeleton Fresh water Entamamoebas Parasite Live in guts of host Choanoflagellates We think the earliest animals were closely related to Choanoflagellates Found in colonies Colonies have similar structure to a spinal column Cell body, microvilli, and flagellum Ecological and Economic Role of Protists Algae are the largest component of photosynthesis in the oceans Diatoms and Dioflagellates Protists are important to marine and fresh water food chains, recycling CO2 Open ocean: Surface waters teem with microscopic protists, such as diatoms. Shallow coastal waters: Gigantic protists, such as kelp, form underwater forests. Intertidal habitats: Protists such as red algae are particularly abundant in tidal habitats. PATHOGENS Malaria is caused by Chromealveta Toxoplasmosis Dianoflagellates Non-pathogenic Archeaplastida - photosynthetic Rhizaria - shelled Protist Symbionts • Symbiosis means "living together" • Can be parasitic or mutualistic Corals Symbiotic dinoflagellate cells produce sugars from photosynthesis Supply corals with sugar Closing Thoughts • Protists show immense diversity of metabolism, habitat and life cycle, which is not surprising, given how long many of the lineages have been diverging. • Protists are essential to global food chains and carbon cycling, and have huge impacts on human health and wealth. • Protist taxonomy and phylogeny remain controversial, and a subject of active research. Lecture 16 (590-592, 600-601) 1. What characters unite the members of Archaeplastida? Red algae, green algae (chlorophytes and charophyceans), and land plants: Monophyletic group descended from an ancient protist that engulfed a cyanobacterium for photosynthesis 2. What are the characteristics of red algae? Reddish colour from accessory pigment, can live in very deep water, some are heterotrophic parasites, most are multicellular, diverse life cycles of alternation of generations, no flagellated stages, depend on water currents for fertilization. 3. Why are green algae currently a paraphyletic group? Common ancestor: Archeaplastida Land plants and red algae are not part of the group containing green algae 4. What are the two main groups of green algae? Chlorophytes - mostly fresh water, can be unicellular or exist in colonies Charophytes - complex life cycles, alternation of generations 5. Provide examples of chlorophytes you have seen in lab. Volvox 6. What characterics define the Streptophyta? Charophyceans and Embryophytes 7. What challenges needed to be met as plants evolved to inhabit a terrestrial environment? Water and nutrients PROTISTS There are many paraphyletic groups that remain well accepted in our classification. Paraphyletic groups help to highlight major adaptive shifts that have occurred in evolution. Archaeplastida The Archaeplastida is a monophyletic group including only: Red Algae Green Algae (Chlorophytes, Charophyceans) Land plants Rhodophyta - Red Algae Mostly marine Cell walls of cellulose and alginic acid (agar) No motile stages Parasites - mostly parasitic on each other Complex Life Cycles Pit Connections Porphyra Source of traditional Japanese food Seaweed grown on nets What is a Pit Connection? There are two types relating to how cells connect with each other Primary - cells connected by cytoplasm when daughter cells do not entirely split after division Secondary - formed between two cells which are not sister but still end up connected by protrusions to connect cytoplasm How parasitism takes place in red algae Parasitic Red Algae Parasite uses secondary pit connections Parasite sends nuclei into the host Revs up metabolism of the host Parasite harvests what is being grown by the host Life Cycle of Red Algae Sexual Reproduction Male and female gametophytes (2n) Spermatangia (n) and egg (n) Fertilization occurs Zygote nucleus migrates to auxiliary cell through secondary pit connection Zygote nucleus (2n) Red Algae - Deep Water algae Where is the compensation point where photosynthesis and respiration are equal? As long as plant is consuming more CO2 than respiring - it will grow. 0.005% of 1% of surface illumination Red Algae are extraordinarily efficient 268 meters deep Green Algae Green Algae are a paraphyletic group of Chlorophytes and Charophyceans Just like plants, green algae have: Chlorophyll A and B Cellulose cell wall Storage product: starch Motile via flagella Chlorophytes Chlamydomonas Small single cell with two flagella Volvox Colony shows multicellularity Division occurs inside parent colony To become motile, volvox must grow and then turn itself inside out because to begin with, all flagella are pointing inwards Sea Lettuce Large sea animal like seaweed Charophyceans Closest algal relatives to land plants Relatively small, microscopic Spirogia Micrasterias Derived characters of Charophyceans and Land Plants Mode of cellulose synthesis Peroxisome enzymes Sperm structure Phragmoplast present Spindle fibers outside of chromosomes Characteristic of cell division Fundamental to show sister taxa Archaeplastida Ancestral Algae Red Algae, Chlorophytes, Charophyceans, Embryophytes Viridiplantae Chlorophytes Charophyceans Embryophytes Streptophyta Charophyceans Embryophytes Plantae Embryophytes Why are the plants not classified as green algae? There are clear characters that separate them Embryo retention Grows in a chamber on the gameotophyte Sporic meiosis ALL plants have it - not just some True alternation of generation Terrestrial Adaptive shift Movement to land Green algae Can have alternation of generation Not all are aquatic Needs to survive: light, oxygen, CO2, water, nutrients, PLANTS The movement to land Requirements for plant life Light Water Nutrients CO2 / Oxygen Temperature Reproduction Lecture 17 (600-604, 612, 738-745) 1. What are four derived traits shared among plants? alternation of generations walled spores multicellular gametangia apical meristems 2. What is xylem and phloem? How do they differ? Vascular tissue made of schlerenchyma cells Xylem - water and mineral transport Includes tracheids (tube-shaped cells) Phloem -transport sugar, amino acids, and other organic products 3. What is lignin and how is it important in the evolution of vascular plants? Lignin - phenolic polymer that strengthens cell walls making them inflexible 4. What are the three cell types and how do they differ from one another (744)? Parenchyma - thin and flexible primary cell walls Collenchyma - unevenly thick primary cell walls for support Sclerenchyma - dead at maturity, and empty straws, with lignin inflexible 5. What is a tissue? 6. What are the three tissue systems (regions)? Dermal tissue - epidermis, cuticle Vascular tissue - transport Ground tissue - everything else Requirements for Plant Life Water Light Nutrients Carbon dioxide/ Oxygen Temperature Reproduction Nutrients C. HPKNS CaFe Mu An Mo Bo Cu Cl Carbon, hydrogen, phosphorus, potassium, nitrogen, sulfur, calcium, iron etc Survival Strategies on Land Absorption and Retention Water Light and Nutrients require absorption and some require retention as well Absorption Water and nutrients Large below ground surface area Light Large above ground suface area CO2 and O2 Large above ground Retention Water and nutrients small surface area above ground Conflict between small above ground for water retention vs. large above ground for light and CO2 and O2. Plants have cuticle to retain water The First Plants Vegatative Simple prostrate structures Flat, protected photosynthetic surfaces Rhizoids for absorption and anchoring Selection for support/water retention leads to structural complexity Kingdom Plantae Many homologies are shared with the green algae Differences relate to: -Selection for structure/ water retention -Selection for protection and dispersal Plant Anatomy Cell types versus tissue types Cell Types Based upon cell wall structure Relates to selection for support All plants are made of three main cell types: (1) Parenchyma cellulose in primary cell wall cell wall is relatively elastic - expand and stretch a little basic plant cell living cell capable of biological processes (2) Collenchyma uneven thickening of primary wall some portions thicker than others provides more support living cell capable of biological processes (3) Sclerechyma Two cell walls Primary(outside) and secondary (inward) walls Lignin - hardening component of secondary wall, impenetrable by water In most cases, these cells are very mature, and dead Plant Tissue Tissue - integrated group of cells with a common function or structure Simple Tissues Made up of one cell type: Parenchyma Collenchyma Sclerenchyma Complex Tissue Made up of more than one cell type Xylem Phloem Three Tissue Regions Dermal Tissue - outside layer Epidermis Ground Tissue - everything else Cortex Pith Vascular Tissue - transport /support Xylem Phloem VASCULAR TISSUE Vascular Tissue region containing two complex tissues which each contain two different cell types. Both Xylem (transport water hard for support) and Phloem (transport sugars) are involved in transport. Xylem: many sclerenchyma cells for hard support Tracheids - has pits Functions in both support and water transport Tracheids diverge to form vessel elements (conduction) and fibres (support). Vessel elements - perforated end walls Specialized for water transport Fibre Cell - specialized for support Phloem: parenchyma cells Transports sugars From where it is produced to where it is needed Sieve-tube elements Clicker Question Vessel elements are sclerenchyma cells that are part of the xylem and function in transport of water. Lecture 18 (600-604, 612-613, 746-751) 1. What is an organ? A specialized center of body function composed of several different types of tissues. 2. What is a stele? A vascular cylinder, with core of xylem and phloem A root has a protostele that is hard 3. How did leaves evolve? 4. What is an apical meristem and how does it relate to indeterminate growth? It is embryonic plant tissue in the tips of roots and the buds of shoots. The dividing cells of an apical meristem enable the plant to grow in length. 5. What is primary growth? Growth produced by apical meristems, lengthening stems and roots 6. What is the structure of a root apical meristem? 7. What is the structure of the root stele (Fig. 35.14a)? 8. How do lateral roots form? 9. How is the shoot apical meristem different from the root apical meristem? 10. What is the structure of the shoot stele (Fig. 35.17 a and b) 11. What are stomata and where are they normally located? Holes controlled by guard cells on the underside of a leaf for gas exchange 12. Why is the leaf mesophyll separated into two layers? Name the layers. 13. How is the vascular tissue arranged in leaves? Dermal Tissue Region Epidermis Outer layer or covering of the plant Usually consists of a single layer of cells No intercellular spaces Interface - between plant and environment Controls transfer: gas exchange Stomata: holes in epidermis can open and close Leaves: Cuticle covering - prevent water loss Roots: Water absorption is much more important Epidermis has root hairs to increase surface area Root Hair Region can be regenerated after time, but consists of most of absorption region of the plant Ground Tissue Everything else in the plant not vascular tissue or dermal tissue Basic cell type Unspecialized Organ Development Shoot - includes stem and leaves Bud Leaf: blade and petiole Root - underground portion of the plant Shoot Apical Meristem Between leaf primordia Branches off developing vascular strand Tips of plants never get old - it is still embryonic As it divides, it grows taller Produces leaves from apical meristem Leaf primordium on either side of apical meristem Axillary bud meristems occur beside apical meristem However, these produce branches, which in turn will have apical meristems Each meristem is like an individual, which reacts to environment through differential growth in order to maximize light or water absorption etc. Leaf is below a branch (or a bud) Opinion Clicker Question Roles of the stem/branches in order of importance: 1. Support 2. Transport 3. Reproduction 4. Photosynthesis 5. Storage Stele - the pattern of vascular tissue in stem or root Eustele - scattered bundles vs. Protostele - sclarenchyma tissue is central core Rhizome - an underground stem Mostly functional in vegetative production Underground storage eg) potatoe Above ground: photosythentic stems Cacti - stem is photosynthetic Leaves are sharp and protective Leaves are part of the Shoot Region Mostly have a role in photosynthesis Leaves are below the axillary bud Node - leaves attached to a stem at the node Internode - the gap between two nodes on a stem Simple Leaf Petiole is central vein of leaf Compound Leaf Pertiole looks like a smaller stem Has individual leaflets Double Compound Leaf Has grouped leaflets We know the difference between leaf and leaflet by the placement of bud Veins include both xylem and phloem Vascular tissue is arranged with xylem on the top, phloem on the bottom Think of the stem: Xylem stays in middle As these bundles move out into branches, the xylem is on the top Further stratification in the leaf Palisade mesophyl - light absorption Larger surface area, photosynthesis occur both sides of elongated cell Spongy tissue - gas is moved throughout the leaf Less surface area, big holes are to allow gas movement Guard cells are at bottom of leaf Root Apical Meristem Has a root cap Root cap protects embryonic cells from soil as roots grow down Stele - protostele in the roots Root does not need extra support Lecture 19 (604-610) 1. What two evolutionary events are highlights in the evolution of land plants? Vascular plants and bryophytes 2. Land plants can be divided into two groups. Which one is paraphyletic? Non-vascular bryophytes are paraphyletic Vascular plants have derived traits and are monophyletic 3. How many phyla of land plants are there? numerous 4. Name the three phyla of non-vascular plants or bryophytes. Phylum Hepatophyta - liverwort Phylum Anthocerophyta - hornworts Phylum Bryophyta - mosses 5. What is distinctive about the non-vascular plant life cycle? Roots begin growth at the base NOT at the tip, even though the apical meristem is at the root tip near the root cap, utilizing an active zone Branches in roots/ lateral roots are of endogenous origin Stems have exogenous origins Potato A potato is a stem, not a root If the potato is left to germinate they grow buds from the eye If you look into the eye of a potatoes you find an external origin = proves a stem Shoot meristem - apical Exogenis origin Root meristem - sub-apical Endogenesis origin THE FIRST PLANTS Vegetative Simple, prostrate structures Flat, protected photosynthetic surfaces Rhizoids for absorption and . Reproductive Antheridia and archagonium Sexual Reproduction on land requires protection Gametes Gametes must be able to fuse Fusion of gametes occurs in fluid/water However, a cell covering is needed for protection Multicellular gametangia A multicellular structure Males: Antheridium - cellular structure around with sperm inside Females: Archegonium - cellular structure with pore and single egg inside Retention of the egg Sperm released into water Zygotes and Embryos Need protection until can develop own protection Retention on gametophyte Sporophyte embryo develops on the parent plant called the gametophyte Effective dispersal on land requires a new strategy Sperm need another stage in the life cycle to promote dispersal (not open water) Gametes cannot be protective because they need to fuse So, we find on land: - increased spore production - increase dispersal in dry or terrestrial environment - how can this be accomplished? - After meiosis of the zygote from 2N to haploid N, repeated mitosis of spores occurs for many identical haploid cells - Mitosis of zygote increases diploid cells, which then each undergo meiosis, producing more spores that are genetically diverse Heteromorphic Alternation of Generation Haploid and diploid stages look different Isomorphic Alternation of Generation Haploids and diploids appear the same Non-Vascular Land Plants Bryophytes Bryophyte Vegetative Traits Light and CO2 absorption .. Bryophyte Reproductive Traits Swimming sperm Increased dispersal Need to know life cycle: Lecture 20 Questions from the text (610-616) 1. When does the fossil record indicate that vascular plants originated? Present-day vascular plants date 420 million years 2. How does the life cycle of vascular plants differ from that of non-vascular? Vascuar plants have branched sporophytes that are not dependant on the gametophyte for nutrition. 3. How and why did leaves originate? Leaves increase the surface area of the plant body and serve as the primary photosynthetic organ of vascular plants. 4. How are the lycophytes related to the rest of the vascular plants? Sister taxa Lycophytes are the oldest lineage of present-day vascular plants. Only lycophytes have microphylls, which are small, usually spine-shaped leaves supported by a single strand of vascular tissue. Almost all other vascular plants have megaphylls, which are leaves with a highly branched vascular system; a few species have reduced leaves that appear to have evolved from megaphylls. Microphylls originated from sporangia located on the side of the stem. Megaphylls, by contrast, may have evolved from a series of branches lying close together on a stem. Lycophytes are a sister group to the rest of the vascular plants. Lycophytes are seedless; so are some vascular plants. This is why we separate lycophytes from everything else (of vascular). 5. Where does meiosis occur in ferns? 6. What characters unite horsetails and ferns on the same clade? Homosporous 7. What is peat moss and where does it grow? "Sphagnum" is a wetland moss that forms extensive deposits of partially decayed organic material known as peat. Clicker questions (1)The plant gametophyte produces gametes via mitosis. (2)Alternation of generation refers to: Alternation of sporophyte and gametophyte ONLY. This is two multicellular stages. One haploid, one diploid. (3) The bryophyte life cycle has a: Dominant gametophyte, dependent sporophyte Sporophyte meiosis Meiosis - movement from diploid to haploid (N) Fertilization - two haploids make a diploid (2N) Mosses Fig 29-8-3 Meiosis occurs in sporangium in the sporophyte They germinante to generate protonemata (haploid = N) Gametophyte (haploid = N) includes both the gametophore and protonemata Male gametophyte vs. female gametophyte Antheridia male vs. archegonia female Raindrop transfers male spore INTO the female archegonium for fertilization. Zygote grows up the archegonium, sometimes pushing up haploid remnants. The female gametophye supports the sporophyte (diploid = 2N) until it grows up. The gametophyte supports mature sporophyte Method of dispersal: When cap comes off, teeth push into spore mass so when it gets drier the spores are dispersed into the dry air. *Must know comparison of bryophyte phyla: Liverworts, hornworts, and mosses All have dominant gametophyte and dependent sporophyte Peat Mosses "Sphagnum" Specialized leaves Live in very acidic environment - bogs Commercially important Living cells of peat leaves are small. Most cells are empty and dead. The dead cells take up and hold water to keep moss moist. The increased acidity also provides a wound dressing as an anticeptic. Peat fills in lakes Peat in pushed down to the bottom A peat bog is a highly acidic filled in lake People have found perfectly preserved bodies in peat bogs Excellent source of fossil information Peat is an excellent, clean-burning fossil fuel Peat is more difficult to transport, so it is really only useful small scale In Canada we harvest peat by strip mining Some people are concerned we are destroying peat bogs about to lose them Vascular Plants Monophyletic group Dominant sporophyte (2N) and dependent or independent gametophyte (N). Plant life in the Devonian Vascular plant occurs first in fossil record: believed they preserve better. Psilophyton Oldest vascular plants Kidney shape structure on side of stem = lateral sporangia Flaps of tissue on stems Enations - these flaps of tissue do not have vascular tissue Cooksonia Looks different in that it has swellings the tips which are terminal sporangia No enations The first vascular plants No leaves No roots Simple, small Two main groups: Terminal sporangia Lateral sporangia and enations How did leaves evolve? Two theories: evidence that both happened (1) Enation Theory - Microphylls Vascular tissue > sporangia > microphyll (2) Telome Theory - Megaphylls Overtopping growth > other stems become reduced and flattened > webbing develops in the megaphyll leaf Modification of the branches LIFE CYCLE THEORY What is the nature of the sporophyte? (1) Sporangium antithetic theory (2) Same as gametophyte homologous theory Isomorphic alternation of generation is an ancestral trait in plants according to the Homologous Theory but a derived trait according to the Antithetic Theory. Comparison of bryophyte phyla All have dominant gametophyte and dependent sporophyte Liverworts (Hepatophyta) Gametophyte prostrate thallus with gametangia (antheridia and archegonia) often on stalks Small sporophyte (sporangium) Hornworts (Anthocerophyta) Gametophyte Similar to liverworts, gametangia embedded in gametophyte Larger photosynthetic sporophyte Mosses (Bryophyta) Gametophyte Axial gametophores plus protonemata Complex sporophyte Lecture 21 (618-621) 1. What important innovation in the plant life cycle did reduction in gametophyte size facilitate? Tiny gametophytes can develop from spores retained within the sporangia of the parental sporophyte. This protects the female gametophytes from environmental stresses. 2. What kind of spores produce male gametophytes and what kind of spores produce female gametophytes? Seed plants are heterosporous. Megasporangia produce megaspores that give rise to female gametophytes. Microsporangia produce microspores that give rise to male gametophytes. 3. What are the layers that make up an ovule? Integument - sporophyte tissue envelops and protects megasporangium Gymnosperm megasporangia are surrounded by one integument (whereas those in angiosperms usually have two integuments) The whole structure: megasporangium, megaspore, and their integuments 4. What, in a bryophyte life cycle, is homologous to a pollen grain? A microspore develops into a pollen grain that has a male gametophyte enclosed within the pollen wall, which protects the pollen grain in pollination. Bryophytes have flagellatted sperm to travel short distance. 5. What is the relationship between an ovule and a seed? The ovule is the stage before complete fertilization. Fertilization of the ovule initiates the transformation into a seed. 6. How are spores and seeds different? How are they similar? Similar: Protective Dispersal Tiny size Different: Spores are unicellular; seeds have multicellular layer of tissue Seeds have a supply of stored food The seed can then germinate on its own Lycophytes (Phylum) .. (Phyum..) eg) lycopodium obscurum Longitudinal Section of Strobilus consisting of a central stem supporting a cluster of Leaf and sporangia are associated in location Lateral sporangia Cones are terminal Homosporous: produce spores that are all the same Typically a bisexual gametophyte that produces both sperm and eggs Heterosporous: Megasporangium/ megasporophyll > produces megaspore > female gametophyte > eggs Microspoarangium/ microsporophyll > produces microspore > male gametophyte > sperm All emerged from enations Phylum Pterophyta (Ferns and relatives) Leaves - megaphylls Sporangia - terminal - tips of "branches" Includes both ferns and horsetails Horsetails - Sphenophytes Sporangia on sporangiophores, in cones Leaves are small and whorled (they come off in a ring/ collar of leaves around stem) silica in cell walls Non-photosynthetic reproductive stem Vegetative, photosynthetic stem with branches Sporangia on cones is made up of modified branches No leaves in the cone - all little stalks What theory explains this branch modification? Telome theory: modification of branches Ferns Sporangia on megaphylls Sporangia clustered in sori (sing.= sorus), not in cones Modification of the stem Considered so similar even though they look very different Rhizome - underground stem Frond - the leaf in the fern Rachis - the stalk (like a petiole) LIFE CYCLE OF FERNS Must know how it is similar and different to bryophyte Fern gametophyte is independent Does not require sporophyte for nutriets Homosporous Telome Theory: Where would you expect the ancestral position of sporangia to be? The tips of veins Veins are homologous to branches Sporangia are terminal As ferns have evolved, this may not always be true The underside of the leaves of ferns Sori are located under leaf Sorus: cluster of sporangia Inside a sporangium are many spores Dominant independent sporophyte Mosses and other Nonvascular Plants Constraints on the Life Cycle Necessity of water: gametophyte needs water for sperm dispersal How can this problem be solved? Two issues (1) Where would the gametophyte live? What clues to strategy do bryophytes provide? The sporophyte lives on the gametophyte Moist inside sporangium Germinates inside sporangium (2) How will the sperm be transferred? Text questions (621-625, 751-754) 1. What important characteristics link Archaeopteris to the evolution of seed plants? The heterosporous Archaeopteris had characteristics of seed plants such as a woody stem. It is called a progymnosperm - an extinct seedless vascular plant that may be ancestral to seed plants. 2. When (geological period) did seed plants begin to dominate in the fossil record? 360 million years ago, which is about 200 million years before angiosperms 3. Where are ovules located in a pine tree? How does fertilization occur in a pine tree? Is it unisexual or hermaphroditic? -Ovules are in cones: most conifers have both ovulate and pollen cones -Pines are heterosporous. -Pollination occurs when a pollen grain reaches the ovule. The pollen grain then germinates, forming a pollen tube. -Fertilization does not occur until the eggs are mature. By this time, two sperm cells have developed in the pollen tube, which extends to the female gametophyte. Fertilization occurs when the sperm and egg nuclei unite. 4. How does secondary growth affect the development of the mature plant body? Secondary growth is the growth in thickness produced by lateral meristems and adds girth to stems and roots in woody plants. The secondary plant body consists of the tissues produced by the vascular cambium and cork cambium, which thicken the stems and roots of woody plants. 5. What are the two main secondary meristems? (1) Lateral meristems (2) vascular cambium 6. Where is the vascular cambium located and what does it produce? It is in between primary xylem and primary phloem in the center of a stem or root. The vascular cambium is a cylinder of meristematic cells, often only one cell thick, which thickens roots and stems. It increases in circumference and also adds layers of secondary xylem to its interior and secondary phloem to its exterior. It increases vascular flow and support for the shoot system. (1) Elongated initials of the vascular cambium produce cells such as the tracheids, vessel elements, and fibers of xylem, as well as the sieve-tube elements, companion cells, parenchyma, and fibres of the phloem. (2) Shortened initials produce vascular rays - radial files of cells to connect 7. What is the difference between heartwood and sapwood? Heartwood layers are older and closer to the center, no longer transporting water and minerals (xylem sap). Sapwood is the newest outer layers of secondary xylem. 8. What is periderm and how is it related to cork? Each cork cambium and the tissues it produces comprise a layer of periderm. Dispersal How will sperm be transferred? Retain gametophyte in the sporangium, to provide a moist environment Intermediary Wildlife Insects Raindrops Splash sperm to female gametophyte Retain moisture so sperm can swim Gametophyte Antheridium and archegonium are on gametophytes You could move the whole gametophyte if it is small Could use wind to disperse male gametophyte to stationary female Egg is larger than sperm, so it stays stationary Division of labour A heterosporous plant is more successful! Both sexes are produced in a heterosporous plant Fertilization Male gametophyte lands on female sporangium and then can grow through the sporangium and deposit sperm Conifers: cones and pollen grains Packaging your own water Pollen grain - male gametophyte Not spores because are more than one cell How will the sperm be captured? An ovule Ovule - an integumented megasporangium Sporangium contains megaspore that germinates into female gametophyte Integument is a layer of tissue derived from branches Female structure of layers around the female gametophyte that include the sporangium, and lobes of integument Ovule: Integument > sporangium wall > female gametophyte Fig 30-3-2 Textbook is wrong in this figure Spore wall is not living and does not expand That is the megasporangium wall Sporangium wall become nucellus Seed - fertilized ovule Development of seeds is a major evolution in plants The seed shows homologies with other plants: gametophyte and sporophytes Monophyletic group Gymnosperms "naked seeds" mostly cones Angiosperms "enclose seeds" mostly flowers Seed Plants Heterospory Eusteles Presence of bifacial cambium o Xylem to the inside, phloem to the outside o structure that produces wood Where did seed plants come from? Progymnosperm ancestor Heterosporous Megaphyllous W oody Vascular Cambium Division into xylem and phloem Secondary tissue is produced by vascular cambium Wood = secondary xylem How would you characterize bark? When you peel off bark on a tree trunk you are left with secondary xylem The boundary between bark and wood must be the cambium Everything outside wood Mostly parenchyma cells Bark includes: secondary phloem (closest to cambium), primary phloem (further out) cortex cork cambium cork (tissue harvested from surface of tree) Rings in tree trunks Cambium divides in spring and summer Grows quickly and grows large cells in spring, small cells later fall It is dormant in the winter Growth rings are the difference in cell size between end of last year and spring. Asymmetrical growth can occur for the tree to adapt and grow in a direction Periderm - refers to cork and cork cambium Lecture 23 Text questions (625-630, 801-804) 1. Name the parts of the four rings of floral structures that make up the flower. Carpels Stamens Petal Sepal 2. What are the floral organs derived from? 3. What is a fruit? A mature ovary 4. How many cells (or nuclei) make up the male and female gametophyte (respectively) of the angiosperm? 5. What is endosperm and what is its function? 6. What is double fertilization? 7. What is an embryo sac? 8. Distinguish between the terms carpel, ovary and pistil as used in describing an angiosperm flower. Gymnosperms Four phyla Phylum Cyc Phylum Ginkgophyta Only contains one species Ginkgo biloba Two big lobes on leaf Pollen-producing tree in China Separate male and female trees Phylum Gnetophyta 3 very distinct genera Gnetum Source of ephedrine - strong stimulant - good for tea Welwitschia Phylum Coniferophyta Confirs: Fir, Larch, Pine, Sequoia, Juniper Megaphyll Leaf Sporangia is on underside of leaf Female Cone Bract Scale (branch) - ovules located here on top of scale (leaf) Bud (becomes branch) is located above the leaf Angiosperms = Phylum Anthophyta angio = container for sperm = seed Bisexual reproductive structures Eg) barley, water lily, oak trees, birch trees, poplars, most are wind-pollinated FLOWERS Calyx - sepals Corolla - petals Androecium - stamens (leaves with microsporangia on them) Anther filaments Gynoecium - female - made up of carpels Ovaries, stigma, style Four parts of the plant Carpels It is a leaf unit that makes up the ovary Stamens Petal Sepal Ovules are in an ovary Seeds are in a fruit A fruit is a mature ovary Lecture 24 (630-632, 804-805, 809-811) 1.Which of the two angiosperm groups (monocots or dicots) is paraphyletic? Monocot - member of a clade consisting of flowering plants that have one embryonic seed leaf, or cotyledon. Dicot - a term traditionally used to refer to flowering plants that have two embryonic seed leaves, or cotyledons. Recent molecular evidence indicates that dicots do not form a clade; species once classified as dicots are now grouped into eudicots, magnoliids, and several lineages of basal angiosperms. 2. What is the major constraint relating to pollen dispersal that is less of an issue with seed dispersal? 3. What are the characteristics of wind pollinated flowers? 4. How do bee pollinated flowers differ from bird pollinated flowers? 5. What develops into the pericarp (fruitwall)? The ovary wall becomes the thickened wall of the fruit, or pericarp 6. Distinguish among simple, aggregate, multiple and accessory fruits. simple fruit - a fruit derived from a single carpel or several fused carpels. aggregate - derived from a single flower that has more than one carpel. multiple fruit - a fruit derived from an inflorescence, a group of flowers tightly clustered together 7. What are two main types of abiotic and two types of biotic seed dispersal? Abiotic: wind water Biotic: animals insects Derived characters defining angiosperm Flowers Closed Carpels Fruits Vessel members in the xylem Reduced gametophytes Double fertilization Xylem cells in angiosperms Tracheids diverge to form vessel elements (conduction) and fibres (support) Reduced Gametophyte Male Gametophyte Microsporangium is the anther pollen sac Produces microsporocyte (2N) meiosis POLLEN GRAIN 4 microspores (N) Each of 4 microspores Generative cell (N) Female Gametophyte Megasporangium (2N) Megasporangium wall Megasporocyte (2N) Integuments (2N) Micropyle meiosis Surviving megaspore (N) EMBRYO SAC Ovule 3 antipodal cells (N) 2 polar nuclei (N) 1 egg (n) 2 synergids (N) TOTAL: 8 cells highly reduced gametophyte Clicker The closest relative to an angiosperm is a gymnosperm A reduced angiosperm female gametophyte is beneficial because less maternal commitment prior to fertilization and less weight for the sporophyte to carry and detrimental because another food source for the embryo must be found. Life Cycle of Angiosperms Mature flower Meiosis in the anther Pollen grain lands on stigma, pollen tube grows down stile Release sperm into ovule Fertilization occurs Double fertilization Fertilization of both the egg and the polar nuclei in the ovule 1. Endosperms nucleus (3N) 2 polar nuclei plus sperm 2. Zygote (2N) egg plus sperm Coevolution Angiosperms use animals to aid their dispersal Two different dispersal stages Seed dispersal syndromes Abiotic: wind, water Biotic: animals Active: sometimes involves attraction/reward brightly colored fruit food sugar around seed in ovary wall seed has oils that are not attractive to animal Passive biotic dispersal such as burrs latch onto animal Pericarp (fruit wall) is often involved in promoting dispersal It acts as an intermediary, and offers protection eg. coconut - strong fibrous pericarp would float in salt water ocean Wind dispersal Dandelion "parachute" white hairs break off of flower stem Tumble weed Animal dispersal Seed caches Excretion of whole seeds Simple Fruits Peas are an example of a simple fruits Strawberries Yellow things are fruits/ ovary walls Red part is stem Accessory Fruit: red part attractive portion are not ovary Aggregate Fruit: single flower, aggregation of fruits from single flower Multiple Fruits Pineapple Many flowers, one carpel Pollination syndromes -Biotic/Abiotic dispersal of pollen grains -Biotic involves attraction and reward Attraction: color of flower Reward: fruit, nectar -Precise destination of the pollen to a stigma of a receptive flower Clicker Opinion Question True: Flowers should evolve to attract as many different pollinators as possible so that reproduction is maximized. False: precise destination, need specialized subset of pollinators otherwise pollen is wasted going to places where there are no receptive Pollination syndromes and strategies Wind Insect Bird Introduction to Fungi pages 636-643 Fungi are absorbtive heterotrophs Fungal Characteristics • Absorbtive heterotrophs. • Cell walls of chitin (protein found in insect and crustacean exoskeletons) • Life cycle with zygotic meiosis (except a few chytrids) • Decomposers, parasites (and predators) • Multicellular - mycelium made of hyphae • Or Single-celled (yeast) Fungal Morphology Fungus can be very large, extending underground and can be colonial Fungi have hypha to absorb nutrients Fungi can be predatory (on nematodes) (a) septate hypha - discrete cells divided by a septum (b) coenocytic hypha - all nuclei float around in common cytoplasm (also aseptate) Movement of Cytoplasm allows for fungi to grow quickly Fungal Life Cycles Asexual Reproduction Germination Mycelium - haploid dominant mycelium organism Spore-producing structures Spores Sexual Reproduction Plasmogamy (fusion of cytoplasm) Heterokaryotic stage (unfused nuclei from different parents) Karyogamy (fusion of nuclei) Zygote Meiosis Spores Germination Heterokaryotic Stage Gametangia containing multiple nuclei Two hypha grow together Compatible + and Two parents have nuclei float around in cytoplasm Dikaryotic Stage In septate, we get a single nuclei from each parent in same cytoplasm Fungal sex, sexes and parasex • Some fungi can sexually reproduce with themselves. • In those that don't, different "sexes" consist of individuals with different alleles at one or two loci. May be many "sexes" in population. • Often these genes are responsible for forming the dikaryotic phase. • Asexual fungi can also produce recombination through a strange process called parasex. Fungal Ecology Decomposition Fungi can break down cellulose, lignin, karatin, etc. that are hard compounds Fungal mycelia convert cellulose and lignin (from wood) into sugars and other small organic compounds Fungi can also attack living trees Adaptations in Fungi Adaptations for dispersal of spores -Shoot spores out by building up hydrostatic pressure -Spores land in grass Can be eaten by herbivores, and deposited in a new habitat Economics of Fungi Plant parasites Important fungal disease agents Leaves toxins on our food and harm our plant-life Grains: mildews, black rust on wheat, corn light Trees: white rot, dutch elm disease Fruit: mold Rye: ergots causes hallucinations (Salem witch trials) Fungi as Food Mushrooms Molds (to flavour cheeses) Truffles Brewing and baking Yeast Wine and Beer Pharmaceuticals Penicillin comes from a mold called penicillium Fungi for Biological Control Protect plants by killing off insect parasites Fungi have an ability to secrete enzymes that break down chitin in the body of the insects Evolution of Fungi Fossil Record Suggests animals and fungi separated before we see them in the fossil record Very little evidence Linked to the evolution of land plants in movement to terrestrial environments Symbiotic relationships Chytrids • 1000 spp. Found in freshwater and soil. • Single celled and multicellular. • Decomposers, parasites, and mutualists. • Likely the first fungal lineage to diverge. - Only group with flagellated stages. Chytrids can be parasitic Can contribute to the decline of amphibians by causing skin diseases Digestion Chytrids can help break down grass in the stomach of cows Closing thoughts • Fungi are absorbtive heterotrophs with unique lifestyles, metabolism and life cycles. • Fungi have important roles in the global nutrient cycles, and major economic importance, particularly as plant pathogens. • Fungi evolved from flagellated protists. • The primitive Chytrids, a paraphyletic group, are the only fungi which retain flagellae. • Chytrids are an inconspicuous group, but contain some important pathogens and mutualists. Fungi pages 643-646 Zygomycota (Glomeromycota) Ascomycota Only zygotic meiosis No flagellated stages Adapted for terrestrial life Zygomycete Member of the fungal phylum Zygomycota, characterized by the formation of a sturdy structure called a zygosporangium during sexual reproduction. Phylum Zygomycota • 1,000 species. • Aseptate fungi. • Almost all decomposers (molds) • Small and short-lived heterokaryotic stage. • Asexual reproduction more common than sexual. Life Cycle Mating type + and mating type Gametangia with haploid nucleus PLASMOGAMY Young zygosporangium (heterokaryotic) Sexual reproduction KARYOGAMY Diploid nuclei MEIOSIS Sporangium Haploid spores Dispersal and germination Mating type + and mating type Asexual Reproduction Spore dispersal and germination Mycellium Sporangia Ecology of Zygomycetes • Mainly decomposers reason why not to put bread in refridgerator: grow in cool damp conditions Parasitic zygomycetes Colonize insects, spreading throughout body and killing insect Predatory Zygomycetes Makes rings to catch nematodes Spore dispersal mechanisms of zygomycetes Live in cow dung and shoot spores several meters into grass Asexual spores attach selves to fly, reproduce, then explode out of fly body Spirodactylon spores in rat pellets disperse by rats carrying them, the spores are spiral "burrs" that latch onto rat fur, rats lick, ingest, excrete, smell pellets Glomeromycota • Only 160 species • Aseptate. • Sexual reproduction unknown. • Extremely important as mycorrhizae (we will discuss this next day). • Reproduce asexually though sporangia at ends of hyphae. Ascocarps Ascomycota • 65,000 species. Most diverse group of fungi • Septate hyphae • Some unicellular forms (yeasts) • Terrestrial, marine and freshwater. • Sexual reproduction through ascospores produced in asci. • Longer lived and larger dikaryotic stage. Life cycle Sexual Reproduction Two hypha come together Mating type (+) and mating type (-) Plasmogamy Dikaryotic hyphae Ascus (dikaryotic) - small clear sac, plural asci, each containing 2 nuclei Karyogamy Diploid nucleus zygote Meiosis Four haploid daughter nuclei Mitosis Eight ascospores In an ascocarp, many asci, may be both haploid and diploid Dispersal Germination Mycelia Asexual reproduction Haploid spores Dispersal Germination Hypha Mycelium Conidiophore Haploid spore on end of hypha (conidia) Mycelium is the dominant part of the life cycle Relatedness to parent in Ascomycota: Conidia is exactly 100% related to parent (asexual reproduction) Ascospore average 50% Types of ascocarp Related to method of spore dispersal: • Apothecium - Open. Cup or saucer shaped wind dispersal • Perithecium - Small opening (ostiole). Flask or pear shaped. can act like a canon to fire out spores wind dispersal • Cleistothecium - no opening. Spherical. animal dispersal ascocarp provides protection for spores Asexual reproduction via conidia • Produced entirely by mitosis. • Often produced "naked" on the end of a specialized hypha - a conidiophore. • Sexual and asexual reproduction often occur at different times and on different substrates, leading to confusion and delay. • Typically produced in huge numbers. Survey of the Ascomycota About 700 molds Grain infection: ergots Infect insects Powdery mildew Athlete's foot is caused by fungi Penecillium Can grow with little water present Yeasts • Unicellular fungi. • Not a monophyletic group. Description of morphology, not a taxonomic group • Reproduce mainly by budding. Budding = asexual reproduction • Some can reproduce sexually by asci. Closing thoughts • Ascomycota are extremely diverse but monophyletic group. • Defined mainly by sexual reproduction, but evidence suggests many asexual species are Ascomycetes. • Contains many of the most important plant pathogens. Fungi and Fungal Friends Basidiomycota, Lichens, and Micorrhizae pages 646-652, 795-797 Ascomycota and Basidiomycota The two most similar phyla of fungi Basidiomycota • 30,000 species. • Septate hyphae • Mainly terrestrial • Sexual reproduction through basidiospores produced on a basidium • Even longer lived and larger dikaryotic stage. Structure of septa Ascomycetes and basidiomycota have a wararin body to block pores between cells Function seems to be to ensure that nuclei do not move between cells Life Cycle Sexual Reproduction Haploid mycelia Mating type (+) and (-) Plasmogamy Dikaryotic mycelium Gills lined with besidia (bottom ribbing on a mushroom) Basidiocarp - dikaryotic (n+n) Basidia (n+n) Karyogamy Diploid Nuclei - zygote is short-lived Meiosis Basidium Basidium with four basidiospores Basidium containing four haploid nuclei Basidiospores (n) Dispersal and germination Haploid mycelia (1) Class Hymenomycetes • Spores exposed at maturity. • Many have mechanisms to shoot them, but usually only a very short distance. Commonly called "gill-less" fungi Shelf fungi (important decomposers of wood) appear as half circles on logs (2) Gill Fungi • Mainly decomposers or ectomycorrhizae, especially with conifers. • Basidia produced on gills or lamellae. Toadstool mushroom Anatomy of a mushroom Cap - umbrella like structure on top that keeps spores dry Gills - underside of cap Annulus - veil that covers stipe, falls off at maturity Stipe - grows up out of ground, like a stem (3) Class Gasteromycetes Distinguished by their spore dispersal Puff-balls (4) Rusts and Smuts • 6000 rusts and 1000 smuts. • All parasitic on plants, and include many common crop diseases. • No basidiocarps. • Different septal pores from other Basidiomycetes. The odd ones of fungi, most are parasitic Complex life cycle between wheat and barberry dikaryotic spores infect wheat plant basidiospores grow into barberry Some structures similar to land plants Fairy rings Produced by a fungus living underground Middle of fairy ring no longer has enough nutrients Basidiocarps are produced around the entire perimenter Mycophagy Edible fungi are mostly found in basidiomycota Lichens • Lichens are symbiotic associations between a fungus and a green algae (or cyanobacteria). • Most common with Ascomycota (98%) which form lichens, but Basidiomycota, Zygomycota and Glomeromycota are all known to form lichens. • About 20,000 lichens known. Three layers in a cross section of a lichen (Fungal hyphae is most of body) (1) Fungal layer Asci produced by fungus (2) Algal layer Soredia - Asexual reproduction through soredia Fungus may be parasitic to algae (3) Fungal layer Substrate One organism or two? Three Morphologies of Lichen 1. Crustose (encrusting lichens that live on rocks) Extract minerals from rock Absorb moisture from air 2. Foliose (leaflike) Often grow on trees 3. Fruticose (shrublike) Lichen Ecology Long lives Can live in extreme cold or hot environments Arctic - cold environment, lichens grow on rocks Lichen as Bioindicators Sensitive to acid rain and air pollutants When lichens disappear, we need to be concerned about air quality Mycorrhizae • Mutualistic association between plant roots and fungi. Plants grow better with Mycorrhizae Expansion of root system of plants Improve ability to absorb minerals, nutrients, and water Some evidence of protection from pathogens • Probably date back to the earliest land plants and fungi. • Two main types: - Ectomycorrhizae EMF form sheaths around roots and penetrate between root cells, providing a new route for sugar transport Basidiomycota and a few Ascomycota - Arbuscular Mycorrhizae AMF Contact to plasma membrane in root cells AMF penetrate cell walls, but not plasma membrane Glomeromycota (1) Ectomycorrhizae • 2000 plants and 5000 fungal partners. • Many commercially and ecologically important tree species included. Development and morphology Seeds grow on plant root, mycorrhizae colonize root Form sheath around root Greater surface area Plant no longer needs to put energy into producing root hairs (2) Arbuscular Mycorrhizae • 300,000 plants but only 160 fungi, all in the Glomeromycota. • Penetrate cell walls of root cortex, and form tree-like arbuscules which are completely surrounded by the plasma membrane of the cells. • Roots are also surrounded by fungal hyphae growing in the soil and extending the volume and surface area for absorption. Development and Morphology Reproduction • Almost entirely asexual reproduction. • Produce very large, thick-walled spores filled with storage lipids. These can survive for long periods until conditions are suitable for growth. (Grow under similar conditions to partner plant seeds). Spores often spread with seeds AM in agriculture Use mycorrhizae fungi to help plants grow Other fungal mutualisms Endophytes reduce presence of pathogens Ants take leafs to fungi inside nest Closing thoughts • Basidiomycota are less morphologically and metabolically diverse than the Ascomycota, but still very common decomposers and ectomycorrhizae. • Lichens are a common and diverse association between fungi and algae, which behave like single organisms despite their dual nature. • Ecto and arbuscular mycorrhizae are almost universal in plants, and go back to the earliest invasion of land by both kingdoms. Introduction to Animals Characteristics of animals Multicellular, ingestive heterotrophs No cell walls Many have muscles and nerves Genome contains Hox genes Main Lineages in Animals See Ch.32 Lineage figure Protostomes Grades - basic similarities (eg) worm-like bodies No longer used Clades - branches on evolutionary tree Discovered from biological evidence Body Symmetry Assymetry - no symmetry, sponges Radial Symmetry - jellyfish Bilateral Symmertry - lizard, human, have an anterior and posterior end Tissue Layers Sponges • Some animals have no true tissues (sponges) Eumetozoa • Diploblastic animals have two tissue layers - Ectoderm - Endoderm • Triploblastic animals have three tissue layers - Ectoderm - Mesoderm - Endoderm Fate of Tissue Layers Ectoderm - epidermis, nervous system Mesoderm - notochord, skeletal and muscular system, reproductive system Endoderm - inner organs Body cavities Example: Flatworm Body covering from ectoderm Tissue-filled region from mesoderm Wall of digestive cavity Example: Pseudocelum Body covering Muscle layer No inner layer of muscle, rather a fluid cavity Digestive tract from endoderm Example: Hydrostatic skeleton of a nematode Fluid-filled pseudocoelom (under pressure - creates tension in the body wall) Example: Coelomate Tissue layer lines the coelom and suspends internal organs (from mesoderm) Function: Peristalsis Peristalsis allows food to move through gut independent of body movement Refer to: Animal Phylogeny from Campbell 5th ed. is no longer correct Animal development Sexual Reproduction Gametic Meiosis with oogamy (egg and sperm) Asexual Reproduction Mitosis Animal development Zygote Cleavage Eight-cell stage Cleavage Blastula (hollow ball of cells) Gastrulation Gastrula Blastopore (opening) Archenteron (developing gut) Ectoderm Blastocoel Endoderm Protostomes vs. Deuterostomes Early stages of cleavage can help distinguish between Protosomes/ Deuterostomes in the eight-cell stage. We can also look at the fate of the blastopore and coelom formation. Protostomes spiral and determinate eight-cell stage of cleavage Deuterostomes Radial and indeterminate eight-cell stage of cleavage solid masses of mesoderm split and form coelom folds of archenteron form coelom mouth develops from blastopore anus develops from blastopore Form and Function in Animals • Gas exchange • Digestion • Osmoregulation - balance of ions, excretion of metabolic waste • Move to land Is bigger better? • Not all animals are big • But generally, multicellular animals are bigger than the single celled protists from which they evolved. • What problems has this created, and how has natural selection produced answers? Cube - Square Law As size increases, surface area does not increase at the same rate as volume Surface to volume ratio decreases as animal gets bigger Gas Exchange - larger animals needs to increase surface area for gas = development of high surface area lungs Animals then evolved circulatory systems to carry oxygen to muscles and other areas of the body Digestion (esp. Absorption) - intestine is folded and has villi to increase surface area for nutrient and water absorption Cube-square law affects more than just diffusion • Heat exchange with the environment. • Muscle strength • Friction and terminal velocity. - "You can drop a mouse down a thousand-yard mine shaft; and, on arriving at the bottom, it gets a slight shock and walks away, provided that the ground is fairly soft. A rat is killed, a man is broken, a horse splashes." JBS Haldane Closing thoughts • Animals are multicellular ingestive heterotrophs. • Animals are a diverse and species rich group. Most of the species are in a few phyla. • Cleavage, a blastula and gastrulation, and fundamental and almost universal elements of animal development. • Much of animals' organ structure can be explained as ways to increase surface area. Sponges Water is drawn into the interior cavity, called the spongocoel. Feeding by phagocytosis Silicea Protein - spongin Sponging is elastic Sponges Water is drawn into the interior cavity, called the spongocoel. Feeding by phagocytosis Silicea Protein - spongin Spongin is elastic Histoincompatibility - body's rejection of foreign tissues Reason for organ transplant rejection Same individual can be separated, then fused back together You cannot make two different sponges grow together Sponges Gas Exchange Choanoflagellates are sessile protists; some are colonial. Sponges are multicellular, sessile animals. Water current out of sponge Sponge feeding cell - choanocyte Phyla Calcarea and Silicea • Marine & a few freshwater • 5500 species • No muscles, organs or nerves • Filter feeders. • 5mm to >1min length. Body wall of sponge EXAMPLES Calcarea - spicules of CaCO3 Silicea -- spicules of silica Silicea - spongin Functions of choanocytes • Generate the currents that draw seawater into the interior of the sponge. • Capture small food particles. • Capturing incoming sperm during mating. Functions of amoebocytes • Digest food collected by choanocytes. • Store food. • Give rise to eggs and sperm (most sponges) • Eliminate wastes • Become specialized to: - secrete sponge skeleton - become epidermal cells Sexual Reproduction • Usually hermaphroditic. • Eggs and sperm produced by amoebocytes (usually). • Eggs (usually) retained, while sperm is released. • Incoming sperm is captured by choanocytes and transferred to mesohyl for fertilization. • Embryo (usually) develops internally to free- swimming stage. Development Develop into flagellated free- swimming larva. Larvae quickly settles onto substrate and develops into adult sponge. Asexual reproduction Simple but sophisticated and efficient • Sponges show histoincompatibility, an animal characteristic. • A + A : Fuse and grow • B+B: Fuse and grow • A + substrate : Grow • A+B: Death Gas Exchange • All animals are aerobic heterotrophs. • Require oxygen in and carbon dioxide out. • Gas exchange depends on surface area. • Because of cube-square law, bigger animals need more elaborate gas exchange organs. • Metabolically active animals will need more gas exchange as well. • Larger animals also need to transport oxygen throughout the body - circulatory system. Basal animals use diffusion Closing thoughts • Sponges are simple congretations of cells, but function as efficient filter feeders. • Gas exchange mechanisms maximize surface area, minimize distance between water/air and cells, and often connect to circulatory systems. Cnidaria and Ctenophora Feeding Animals formerly known as Radiata Phylum Cnidaria • 10,000 species, marine and a few freshwater. • All predatory with stinging tentacles. • Have nerves and muscle. • <5mm to >2m in diameter. • Four classes. Morphology Cnidocytes Thread shoots out and into the prey animal from the tentacle Movement Bell moves down and contracts to push body through water Development Polyp - asexually reproducing Mitosis produces many medusa (2N) Medusa produces egg and sperm by meiosis Female egg and male sperm (N) Fertilization Zygote (2N) Mitosis grows zygote into a polyp Planula larva Senses and Nerves • Nerve net. No centralization. Sensory information is sent throughout entire body, no brain to receive it • Balance organs. Statoliths used for knowing orientation of body • Light receptors. • Touch receptors. FOUR CLASSES (1) Class Scyphozoa • ~200 species, all marine • No true polyp stage. • Include some of the largest sea jellies. • Four gastric pouches. Circular canals (outer edge) Mouth Radial canals (Water flows from mouth at center and radiate out to edge) (2) Class Hydrozoa • 3,000 species, mostly marine. • Polyp and medusa equally prominent. • Jet propulsion aided by a velum. • Some form colonies with specialized polyps. Velum Opening in the bell is controlled by the circular skin called the velum Bottom opening in bell has water that is forced out for movement Specialized colonies Portugese Man O War has three types of polyp attached to a float made from a medusa. Cubozoa Few species, all marine. Small, but with deadly toxins. Complex nervous system and eyes. Anthozoa • Sea anemones and corals. • 6,000 species, all marine. • Medusa reduced or absent. • Some symbiotic with algae. Phylum Ctenophora (Comb Jellies) 100 species, all marine. Swim using ciliated combs called ctenes. Catch prey with sticky tentacles. No larval stage. Hermaphrodites with external fertilization Feeding Extracellular digestion Closing Thoughts • Cnidarians have a life cycle which alternates between sexual medusae and asexual polyps. • Cnidarians and Ctenophora are both diploblastic and radial, with a simple gastrovascular cavity. Cnidarians have cnidocytes and Ctenophora have ctenes. • Digestive systems can be understood as adaptations to the diet and lifestyle of the organism, and a way to increase SA for absorption. LOPHOTROCHOZOA • Diverse group of Bilaterians • Some have a Lophophore or a trochophore larva. Some have neither, grouped mainly by molecular data. • 18 phyla - we will look at 4 - Platyhelminthes - Rotifera - Mollusca - Annelida Platyhelminthes and Rotifera Platyhelminthes are triploblastic Locophores function in suspension feeding in adults Trocophore larva feed and swim via cilia Phylum Platyhelminthes • 20,000 species in 3 classes. • Free-living or parasitic. • Simple excretory system. • Acoelomates with a marine flatworm flattened bodies. Osmoregulation Excretion - getting rid of metabolic wastes Managing ionic balance inside body Maintain homeostasis Class Turbellaria • 3000 species, freshwater and marine. • 2mm to 60cm • Mostly freeliving, a few parasitic. • Hermaphroditic with internal fertilization. • Larval stage rare. Morphology Mouth Pharynx - used in feeding, can be extended out of body Gastrovascular cavity Eyespots - light sensitive Ganglia - gathering of nerves: processing of nerve information Ventral nerve cords Fairly well-developed muscle system Adaptations for Internal Parasitism • Reduction of feeding, sensory and locomotory organs. • Expansion of reproductive organs. • Internal parasites don't need to hunt for food or avoid being eaten. • Biggest problem is finding a new host. Class Cestoidea (Tapeworms) • About 1000 species All internal parasites. • No head or gut. Absorb food from intestine of host through body wall • Segmented body consisting of proglottids. Proglottids - repeating segments of egg and sperm-producing factories Scolex (head) - attach to gut wall with hooks and suckers The largest tapeworms can be up to 40m long! Tapeworm life cycle Adult tapeworm in definitive host. Eggs or gravid proglottids released in host feces. Eggs consumed by Intermediate host Egg develops into Oncosphere larva Larva encysts in organs of intermediate host. Cyst reaches definitive host & grows into adult. Adult tapeworm in definitive host Tapeworm myths and misinformation • Tapeworms won't come up into the host's mouth if you hold a piece of food close to mouth. • Tapeworms can't be used to lose weight (by any sane person). Class Trematoda (Flukes) • About 16,000 species, all parasites. • All have complex life cycle (intermediate host). • Responsible for serious human and livestock diseases. Fluke life cycle Motile larva Human host Ciliated larva from feces get into water Snail host Motile larva can penetrate skin of human to end up in kidneys Morphology Schistosomiasis and other diseases • Caused by several species in genus Schistosoma. • Intestinal and urinary forms of disease. • 200 million people have chronic symptoms, up to 800,000 deaths a year from complications. • Most common in Africa, SW Asia and South America. Monogeneans • Ectoparasites on skin or gills of fish. • Closely related to Trematoda. • Simple lifecycles (no intermediate hosts). Phylum Rotifera • About 2,000 species, mostly freshwater and a few marine. • < 50m to 2mm in length (among the smallest animals). • Mostly predatory with a few parasites on invertebrates. Morphology Crown of cilia Jaws Stomach Anus Pseudocoelomates have an enclosed body cavity partially lined with mesoderm. Mastax organ - rigid structures to grind up prey Well developed sensory organs Light receptors Concentration of nerve cells at anterior end (ganglion) Sexual reproduction is either rare or unknown Asexual for at least 40 million years In drought, can form a cyst that can remain dormant for 50 years Closing thoughts • Platyhelminthes is a phylum of acoelomate worms which include some specialized and dangerous parasites. • Rotifers are mainly freshwater predators feeding with a corona of cilia. PHYLUM ANNELIDA Which worms are Annelids? • Segmented worms. • 16,500 species found in marine, freshwater and terrestrial envts. • Three classes. • Closed circulatory system and complete digestive system. Segmentation Worms have repeating units or segments with the same organs Open and Closed Circulatory Systems Closed - blood pumped through Heart and stays in circulatory system Open - blood vessels open into cavities called sinuses (eg) grasshopper Digestive System of Worms Mouth Muscular pharynx draws food in Esophagus Crop (storage) Gizzard (mechanical mashing of food) Intestine (runs to back of earthworm) Typhiosole - provides more surface area in the intestine (1) Class Polychaeta • 11,000 species, nearly all marine. • Body features: parapodia, setae, and head appendages. • Include active predators, and filter and deposit feeders. Parapodia and setae (chetae) Parapodia are for locomotion and digging tunnels Parapodia are highly vascular and can function in gas exchange Parapodia move together like oars Setae are the small hairs branching off of the parapodia Head appendages Two well developed eyes Jaws Tentacles - filter feeding, gas exchange, or sensory Burrowing Filter feeders tend to burrow partially and stay in the same place to collect food with tentacles Reproduction They send their reproductive organs off to mate while main body stays to eat They produce an epitode which swims off to the breeding grounds Gametes are released into the water in a swarm Fertilization takes place in the water Active or "Errant" Polychaetes Active predators Sedentary Polychaetes Burrowers Can form their own tube Tentacles around head - used for collecting food in seawater Christmas tree and feather-duster worms have a coral-like lifestyle Feather-duster worms are filter feeders Worms that live in hot areas such as underwater volcanoes can have symbiotic relationship with bacteria (2) Class Oligochaeta • 3,500 species, mainly terrestrial (earthworms). • No parapodia, little encephalization, but do have setae. • Distinctive clitellum Morphology Outer waxy cuticle - retain moisture; may be porous for gas exchange Epidermis Circular muscle (can make worm longer and thinner) Longitudinal muscle (can make shorter and thicker) Setae are embedded in the muscle so they can be extended for anchorage or retracted to glide Clitellum Three layers form the cocoon Mucus sheath Secrete proteins that form actual wall Secrete food for egg Eggs laid into this cocoon Cocoon is pushed off of worm's body and left in soil Movement Contracting circular muscles gives a wave of movement Front part stretches forward; back is anchored to not move backwards Front and back anchor; middle stretches and moves forward Reproduction Worms simultaneously exchange sperm Worm A sends some sperm into B; then worm B returns sperm This mechanism has allowed hemaphrodidism to continue Diversity and Ecology Some earthworms can be very long Earthworms are good for soil and plants Aeration - open airways in soil Less surface litter More topsoil More organic carbon, nitrogen, and polysaccharides In hardwood forests, worms can be harmful if heavily infested (3) Class Hirudinea -- leeches • 500 species, mainly freshwater and terrestrial. • Parasitic and predatory. • Have clitellum, but lack setae and septa (no internal divisions). Morphology -Jaws, Pharynx, Crop, Crop cecum, Intestinum (short intestine), Posterior suction -Body walls would prevent suction of maximum blood Predation -on small minnows Mouthparts -Salivary glands secrete anti-coagulates to keep blood from clotting Leeches in medicine -bleeding Parental care -among the best parents in the invertebrate world -keep young in a pouch Leech removal • Option 1: Let it feed and drop off by itself. • Option 2: Find the anterior (skinny) end. Push sideways with a nail next to skin. Remove posterior sucker the same way. Closing thoughts • Annelids are one of many vermiform or "wormlike" phyla. • Annelids are segmented. • Polychaetes dominate the marine envt., Oligochaetes the terrestrial, and Hirudinea freshwater. Phylum Mollusca Phylum Mollusa • 90,000 species, marine, freshwater and terrestrial. • Highly varied body plans in each class. • Open circulatory system (mostly). • 7 classes, we'll look a 4 main ones. Most intelligent invertebrates: squids and octopus Morphology Shell - secreted by the mantle Mantle Mantle cavity Visceral mass - (organs) Nephridium Heart Coelom Intestine Stomach Gonads Radula - scraping organ of herbivorous used to scrape algae off of rocks Mouth Anus Gill Foot - large and muscular Nerve Esophagus cords Mollusc Circulatory system Open circulatory system Blood circulates in blood vessels Blood opens into cavities called sinuses (in the foot and gills) Radula New rasping parts always growing Similar to teeth Mollusc Development • Most have a trochophore stage, but this is usually short- lived. Veliger Larva Large velum used for gas exchange Cilia for locomotion and feeding Gut is protected by the shell Class Polyplacophora (Chitons) • 800 species, all marine. • Shell of 8 dorsal plates - 8 plates running up the back • Large foot used for locomotion and attachment. Do not move very much, so mostly foot is for suction • Use radula to feed on algae. Chiton morphology Underside: mouth and ventral foot Shell top: dorsal plates Class Gastropoda (slugs and snails) • 70,000 species, marine, freshwater and terrestrial. • Shelled (snails) and unshelled (slugs on land and nudibranchs in ocean) • All display torsion. Most are herbivores using radula Gastropod body plan Mantle cavity Stomach and Intestine in shell - twists around Anus - sticks out above head from shell Mouth on underside Move by muscular contraction of foot Gastropod Torsion In early larval stages have straight body During development, torsion or twisting occurs so that anus and mouth end up on the same side of the shell The benefit of this is in adult form for entire animal to retreat into shell Gas exchange Marine: Gills inside mantle cavity, some have terrestrial lungs used underwater sea slugs have large external gills Terrestrial: lung gas exchange to blood vessels, or gills by maintaining water in shell Diversity Carnivorous gastropods Sea snails - think of shell diversity you see wash up on beach Nudibranchs - large external gills, extremely colorful, carry toxins Land snails and slugs Class Bivalvia (clams) • 7000 species, freshwater and marine. • Hinged shell with two valves. Gives ability to clamp shut, or to open shell • No head or sensory organs. • No radula. • Filter or deposit feeders. Muscular foot is used for digging into sediments Morphology Shell Hinge area of shell Mantle Mantle cavity Coelom Gut Gonad Heart Adductor muscle Digestive gland Mouth Palp Foot - used for burrowing Anus Excurrent siphon - draws water in Gills - filters water and provides gas exchange Most clams burrow into soft subtrates and suspension feed. Gills are thin structures for gas exchange. They also trap food particles as water passes through them. Cilia move the particles to the mouth. Scallops live on the surface of the substrate and suspension feed. Symbiotic Bivalves symbiotic bacteria living in gills for protection provide ATP and NADPH to the clams bacteria are chemoautotrophs Carnivorous Clams use siphon to suck up small arthropods Class Cephalopoda • About 600 species, all marine. • Mainly active carnivores. • Big brains and lots of sensory organs. • Highly modified gills, radula and foot. (include squid and octopi) Cephalization Eye of the octopus are large and complex example of analogous/ covergent evolution with vertebrate eye The brain is in a modified foot Motion Jet propulsion Squid has a siphon used for water movement Octopus can use tentacles for propulsion Shell Diversity Nautilus - uses gas in the shell for buoyancy Cuttle bone - skeleton of cuttle fish Squid pen Octopus has lost its shell Modifications Radula modified to beak Skin can change colour and texture (muscles can contract or relax to change skin from bumpy to smooth) for camoflage Bioluminescence Ammonites Group of extinct cephalopods Gem-stones are many from colourful fossils Diversity in Cephalopods Blue ring octopus - highly poisonous Squid sizes - long tentacles for catching prey Closing thoughts • Molluscs are a diverse, species-rich phylum occupying marine, fw and terrestrial habitats. • Most have a basic body plan featuring a mantle, mantle cavity, visceral mass, shell and radula. • The body parts are modified for different functions in the different classes. Phylum Arthropoda Phylum Arthropoda • The most successful and diverse phylum of animals. • Segmented body. • Exoskeleton (or cuticle) made of chitin. • Paired, jointed appendages. Exoskeleton/cuticle Use external skeleton to attach muscles Legs move separately from body Cuticle - waxy layer preventing water loss Open circulatory system Tublular heart Large blood sinuses (1) Subphylum Cheliciformes • A few marine species, and many terrestrial, mostly in Class Arachnida. • Head appendages called chelicerae. • Lack antennae. • Usually have simple eyes. Class Arachnida • 70,000 species mostly terrestrial. • Half are spiders most of the rest are mites and ticks. •Also includes scorpions Morphology Head and thorax have become fused Abdomen behind thorax Chelicerae of spiders are fangs used for feeding Reduced celum Blood sinus Continuous duct from mouth to anus Fluid feeders Blood and sap from plants and animals Canivores, parasites (ticks), Gas exchange via book lungs or tracheae Gas exchange in thin air chambers Filters Lung slit Blood space Spiders - prey on insects Spider bites are fairly rare Spiders are not generally dangerous Mites - spider mites are destructive on greenhouse crops Dust mites in your household eat dead skin that falls off your body When you are allergic to dust, you are actually allergic to the mites in it Ticks - blood feeder parasite on animals Carry many diseases Slow steady pressure is the prescribed method for tick removal Scorpions Larger scorpions are less venomous because they can use their pincers The most venomous scorpions could kill a small child, not an adult human (2) Subphylum Myriapoda •Antennae and several pairs of mouthparts. •Uniramous (unbranched appendages). •Gas exchange by trachea. •Excretion by Malpighian tubules. •Cuticle not waxy. •Simple eyes or no eyes. •Many have repugnatorial glands (excrete sticky and smelly chemicals). Class Chilopoda Maxilliped Centipede Single pair of legs on each body section Predatory on other insects Class Diplopoda Millipede Two pairs of legs in each body segment (3) Subphylum Crustacea • 50,000 species. Large and diverse group • Mostly marine and freshwater. • Biramous appendages. • Two pairs of antennae. • Lots of diverse groups. Lobster Pincers, swimming appendages under each abdominal segment Gas exchange by gills Osmoregulation by glands Decapods - large group Includes crabs and shell fish Mostly marine Some freshwater Isopods - terrestrial Most are fairly small Copepods and Krill Marine Small and numerous Important part of zooplankton group; major food source Barnacles - crustaceans that have sedentary life Calcium carbonate shell (4) Subphylum Hexapoda • Body of head, thorax and abdomen. • Six uniramous legs on thorax. • One pair of antennae. • Compound eyes. • Mainly terrestrial. Not true insects: descended from wingless ancestors Class Insecta Morphology Compound eye Antennae Heart Cerebral ganglion artery Crop Abdomen Thorax and Head fused Wings attached to thorax Anus Vagina and ovaries in female Malpighian tubules Tracheal tubes - gas exchange; characteristic rings; breathing by pressure system (like humans); draw air in; tubes require rings for support Nerve cords Mouthparts Reproduction and Development • Sexual reproduction common, with internal fertilization and separate sexes. • Incomplete metamorphosis: Juveniles look similar to adults but go through several moults as they grow in size. • Complete metamorphosis: One or more larval feeding stages that do not resemble adults. Wings and Flight Evolutionary hypotheses: Perhaps derived in aquatic environment from excess gills could be used for locomotion Wings could have been for thermal regulation Coevolution with Angiosperms Bees and flowers Insect Social Systems Ants, bees, wasps Separation into non-reproductive and reproductive individuals (Think queen ant and the queen bee: for reproduction) Insect Pests Mosquito bites and Bee stings Only a problem when insects are carrying diseases Insects eat about the same amount of crops each year as humans do Closing Thoughts •Arthropods, esp. insects, are the dominant group of terrestrial animals, by almost any measure. Vertebrates are a very distant second. Osmoregulation; Move to Land; and Phylum Nematoda Osmoregulation/Excretion Two main functions: 1. Maintaining osmotic balance of body. 2. Excreting metabolic waste products - mainly nitrogen compounds. Two main trends to remember: • Larger and more metabolically active animals will require more sophisticated excretion system. • Problems in maintaining osmotic balance are different in salt water fresh water and air. Wastes: Urine is most concentrated in salt water fish and mammals Salt water fish excrete ammonia - most toxic Mammals excrete urea Invertebrates move to land •Fossil tracks from about 490 mya. •Arthropods with 8+ pairs of legs and a tail. Arthropod adaptations for land • Rigid support structures. • Waxy cuticle. • Internalized gas exchange surfaces. • Regulatible pores. • Internal fertilization • Use of urea or uric acid rather than ammonia for excretion. Phylum Nematoda • At least 25,000 species, terrestrial (soil), marine,freshwater and parasitic. • Most quite small with a similar body plan. • Tough cuticle of collagen. Morphology Development Movement Not very efficient No circular muscles "waggle" back and forth poor swimmers Parasitic Nematodes • Can be parasitic on invertebrates, especially arthropods (the good). • Can be parasitic on plants (the bad). • Can be parasitic on vertebrates, including humans (the truly revolting). Closing thoughts • Osmoregulation/excretion involves four basic processes: filtration, reabsorption, secretion and excretion. Excretory systems must be more elaborate in larger and more metabolically active animals. • The arthropods were the first animals to invade the land, and have many adaptations for terrestrial life. • Nematodes include some of the vilest parasites you are ever likely to encounter. Let's all go wash our hands. Phylum Echinodermata • 7000 species, all marine. • Most display secondary, pentaradial symmetry. • Have a unique water vascular system. • Internal CaCO3 skeleton. • Six classes. Pentaradial symmetry Star with 5 arms Reproduction and Development No internal fertilization Bilateral symmetric larvae Broadcast fertilization in water Internal skeleton Calcium carbonate elements Stick out of body wall in a spine Ossicles are internal If packed close together, hard shell If loosely packed, more flexibility Pedicellariae For defense and protection Two pronged Cleaning upper surface to remove debris from gas exchange surfaces Water vascular system Madreporite - sucks water in from outside body Stone Canal - draws water into middle Ring Canal - circle Radial Canal - one down each arm Ampullae - bladder on inside of body Tube feet - outside of body, extended by ampullae contracting and pushing water into tube feet for locomotion 1. Class Asteroidea (sea stars) • 2000 species, • Predators • Slow moving, using tube feet. • Found in intertidal to moderate depths. • Five (or multiple of five) arms. Sea Star Digestion Cardiac stomach ejected from body through mouth Pyloric Stomach Morphology Central disk Digestive glands Anus Stomach Gonads Spine Gills Madreporite Radial nerve Ring canal Ampulla Podium Radial canal Tube feet Gas exchange Dermal Branchia Extensions that go outside of body wall to increase surface area Regeneration Regrowth; one arm can regrow the entire body Asexual reproduction Defense Sea star diversity Colours, sizes, and shapes Sun-stars have multiple arms 2. Class Ophiuroidea (brittle stars) • 2100 species, mainly deep water. • Predators, scavengers, deposit and filter feeders. • Digestive system in disc, with no anus. • No ampullae More flexible, slender limbs Tube feet are more used for gas exchange, limbs for locomotion Look like a "spider" in the way they move with their limbs Basket Stars - sedentary filter feeders 3. Class Echinoidea (sea urchins and sand dollars) • 1000 species, mostly shallow water. • Feed on algae (urchins) or detritus (sand dollars) • Spines and pedicellariae may be poisoned (urchins) Spines can be used in locomotion Feeding Feed on algae and detritus 5-part symmetric digestive morphology: mouth, stomach Ecology Sea Urchins can be devastating to kelp forests Sea otters eat sea urchins and keep urchin populations down Sea urchin eggs are considered a delicacy 4. Class Crinoidea (Sea lilies and feather stars) • 600 species, mostly deep water or coral reef. • Most ancient class - fossils go back almost 600 million years. • Filter feeders with oral surface on top. Feather Parts Crown: Arm Calyx Pelma: Stem Radix 5. Class Holothuroidea (sea cucumbers) • 1000 species. • Soft-bodied, may be bilaterally symmetrical. See an anterior end, tube feet all on one side • Oral tube feet modified as feeding tentacles. • Filter or deposit feeders. • Internal gas exchange structures called respiratory trees. For defense, a sea cucumber would spill his guts for distraction If attacked, digestive tract is spilled out so sea cucumber can get away Cost may not be large during inactive feeding periods 6. Class Concentricycloidea (sea daisies) • Only three species known, living on submerged wood. • Biology not well understood. Food may be directly absorbed Closing thoughts • The Echinoderms are an ancient phylum in the Deuterostomes, with a unique water vascular system used in locomotion, feeding, gas exchange (and probably waste excretion). Phylum Chordata • > 50,000 species, most in Craniata (Vertebrates). • Deuterostomes. • Humble origins, but now the largest animals on earth. • Three subphyla Phylum Chordata includes the Vertebrates, along with some smaller and simpler members of the phylum. Chordate characteristics 1. Dorsal, hollow nerve cord 2. Notocord 3. Pharyngeal slits or clefts 4. Tail Spinal column Dorsal, hollow nerve cord Notochord Body parts Muscle segments Mouth Pharyngeal slits or clefts Filter feeding or gas exchange Anus Tail: Muscular, post-anal tail Subphylum Cephalochordata (lancelets) Filter feeders Bury into bottom of ocean Considered to be basal group; closest to ancestral Subphylum Urochordata (Tunicates) Simple, sedentary life as adults Filter food through Pharyngeal slits Larvae mobile; active stage; has all four characteristics • Cranium (braincase) • Encephalization and increased mobility. • Neural crest • Heart with 2+ chambers. • Hemoglobin • Kidneys • Doubling of Hox genes. Subphylum Craniata Neural Crest Formation Skull Formation and neural tube important Notochord Migrating neural cells form eyes and nerves in head Myxini (Hagfish) • Lack vertebrae although are craniates. Primitive eyes. • Notocord retained as adults. • No bone or jaws. • 30 species. Bottom- dwelling scavengers. • Produce slime. VERTEBRATES Petromyzontida (lampreys) • Vertebrae are small cartilage elements. • Notochord retained as adult. • No jaws or bone. • 35 species, marine and freshwater. • Most are parasitic. Mineralized tissues • Bone is made of calcium + phosphate mineral (apatite) - Dermal bone Humans: jaw and skull - Endochondral bone Mammals: large internal bones that anchor muscles Development: Cartilage acts as a template and is laid down first, then bone, then cartilage dissolves • Cartilage - Sharks have only cartilage and have jaws of cartilage • "Tooth minerals" (enamel, dentine, cementum) Extinct jawless fishes Have complex skeletal structures Evolution of jaws • Allowed new ways of feeding, defence and manipulation of environment. • Gnathostomes also have two pairs of fins. • Also associated with changes to vertebrae, muscles, sensory system. Stronger vertebrae for attachment of muscles Adaptations seem to be associated with becoming more active Increasing number of Hox genes could have allowed for changes to occur Evolution: Jaws evolved from the gill arches Are jaws "organs of extreme perfection"? Jaws likely evolved in stages, with different functions at each stage. 1. Improved gill ventilation by active movement of water through mouth. 2. Suction of water used to draw in prey. 3. Teeth evolve to keep prey from escaping. 4. Toothed jaws used for biting and tearing prey. Extinct jawed fish - Placoderms Large and heavily armoured (size of a bus - 10m long) Extinct jawed fish - Acanthodii Look like modern sharks and rays More closely related to the bony fish Strong spines Chondrichthyes • 750 species, mostly marine. Diverged from the bony fish: • Sharks • Skates and rays • Ratfish (chimeras) • Secondary loss of bone. May have lightened themselves • Skeleton of calcified cartilage. • Vertebrae have largely replaced notochord. Sharks Powerful swimmers Predatory Internal fertilization Many give birth to live young Heavier than water, and will sink Have a much larger upper part of the tail fin to give lift Have light oils in organs for bouyancy Shark sensory systems Olfactory systems are sensitive "Nostril" does not connect to gills, only sense of smell to detect blood Ampullae sense electrical fields in water; temperature gradients Strong low-light vision Detect pressure changes in water from vibrations Olfactory > Vision > Electrical Jaws and teeth Teeth are not very well attached, snap off easily Regenerated teeth often and quickly Skates and Rays Feed on molluscs and crustaceans Jaws for crushing hard shells Ratfish Deeper water Single gill opening LUNGS or LUNG DERIVATIVES Ray-finned fishes • 27,000 species, marine and freshwater. • Most in a derived group called the Teleosts. • Found in almost every aquatic environment. Swim bladders Associated with gut Retains buoyancy Can extract gases from the gut or through the stomach Operculum forces water past the gills; so fish can sit and still breath Fin specializations LOBED FINS Lobed-finned Fish Lung fish Fleshy fins Closing Thoughts •Craniates evolved greater body size and complexity than competitors, perhaps because of multiplication of the Hox genes. •Evolution of traits such as increased encephalization, jaws and mineralized skeletons has often been followed by adaptive radiations.