Study guide exam 3 – UH 290 2012

Bioluminescence

Bioluminescence •Bioluminescence is widespread across most forms of metazoan . •Light is typically generated by the itself and only rarely due to bacterial symbionts. Example bacterial symbionts: anglerfish, bobtail squid •Light emitters (luciferins) enzymes (luciferases) •Based on the chemical mechanisms known, luminescence has evolved independently more than 40 times.

bioluminescence •It serves a variety of functions, both offensive and defensive, even within a single organism. •a bioluminescent flash can be seen from tens to hundreds of meters away –Even a single-celled dinoflagellate 0.5 mm long can send a signal to a large 5 m away—the equivalent of a 2 m tall human being able to communicate over a distance of 20 km.

Functions of bioluminescence Offensive – lure prey, stun prey, illuminate prey Defensive – startle, counterillumination, smokescreen, distractive body parts, sacrificial tag, burglar alarm Mating attraction/recognition

Bioluminescence •Luciferin/luciferase normally emits at 470 nm (blue) •Chemical reaction: Luciferin +  LIGHT + oxyluciferin. This reaction is catalyzed by the enzyme luciferase. Sometimes the reaction requires ATP. •Some medusae (cnidarians) emit green light because they have a green flourescent protein • Dragon fish red light emission •Light in the (a light-producing ) doesn't start out deep •red. Initially the light has a short wavelength (red is long-wavelength •light). This light is absorbed by a fluorescent pigment inside the •photophore, which takes the energy and re-emits it as red light sacrificial tag •In this situation, an organism may lose part of its body to a predatory encounter. •These lost tissues can continue to glow for hours afterward even within the predator’s stomach •the glowing tissue can draw attention to the predator, making it risky to consume bioluminescent prey. •This is thought to be the selective force driving the presence of so many black- or red-pigment guts in otherwise transparent –most red and orange pigments absorb blue light cookie-cutter shark •relatively small feeds by taking bites out of the bodies of much larger fish, cetaceans, and squid •unclear how it might get close enough to attack these fast-moving prey. •hypothesized that an optical “flaw” in the counterillumination pattern—a dark band below the mouth—looks like the silhouette of the prey of one of these larger species. When the fish or squid draws near to attack this apparent prey, it is instead attacked by the shark. counterillumination •the silhouette of an against the sunlight filtering down is an easy target. •hatchetfish is prey to bigger fish swimming below it. •narrow silhouette and silver sides make it harder to see, but it also emits light from its belly that is a perfect match in color and intensity to sunlight filtering down Hawaiian bobtail squid •Houses bioluminescent Vibrio bacteria in a “crypt”. Uses the light for counterillumination when they hunt at night •There is a reflector and lens as part of the light organ spookfish •two fields of view in each eye, one focusing upward with a lens and one looking sideways with a •mirror, so it can hunt while keeping an eye on what is happening below

Marine

Type of symbiotic associations - partners mutually benefit (+ +). - one partner derives some benefit while the other is unaffected (+ 0). - one partner benefits at the expense of the host (+ -)

•Degree of intimacy –* Ectosymbiont (or Episymbiont) - a symbiont that lives attached to the surface of its host (including inner surfaces and chambers, such as the cavity or ducts of endocrine glands). – * Endosymbiont - a symbiont that lives within its host, some are within the host cells and are intracellular symbionts.

•Dependence on the symbiosis – * Facultative symbiont - independent, able to exist in a free-living condition. – * Obligate symbiont - dependent, so highly adaptive that they have lost the ability to exist independently.

Evolution by association •By forming a symbiotic association, the capabilities of both partners are added together. This can be faster than “slow” evolution of new capabilities. •We have seen an example of this before with the evolution of eukaryotes

Types of symbiosis •Mutualism, commensalism, parasitism •not always easy to determine which type it is mutualistic symbioses Algal- symbiosis Examples already discussed: Algal-invertebrate symbiosis • anemones •Corals •Upside down jellyfish •Lake Palau jellyfish •Tridacna Some considerations •The algal symbionts are usually acquired from the environment, and usually one species of is always present in a given host species. How does the host recognize the symbiont? •Foreign cells in animals are usually destroyed. Yet the symbionts are not destroyed •The algae are somehow induced to release sugars and amino acids to the host. •The host probably benefits the algae Symbiont transmission •Often “horizontal” –symbionts come from environment •In some cases it is “vertical” –symbionts are in the – symbioses such as Solemya – if this is the case then the symbiont may have no free-living existence and may be on its way to evolving into an organelle

Sea slug Elysia •When it feeds on the alga Vaucheria litorea, it will turn green, by putting the algal chloroplasts into its own gut cells. Is this mutualistic?

Deep sea vent – a variety of symbioses found at deep sea vents Alvinella pompejana – deep sea worm that lives in hot and keeps a covering of bacteria on its back that may insulate it from heat Scaly foot gastropod •Has iron scales secreted by the foot. •Gills have chemoautotrophic bacteria symbionts Yeti •Has hairs that are covered with bacteria may be involved in nutrition

Commensal symbioses Clownfish/ Cleaner shrimps Decorator crab •It is usually found covered in that acts as •Pea •Small crabs often found in oysters and other bivalves tongue-eating isopod ( Cymothoa exigua ). •isopod gets into host’s mouth and grabs onto tongue with seven hooklike legs. Over time, the tongue degenerates, leaving the parasite hanging on to its stub. •This change actually has very little effect on the fish, because it can still hold prey with the parasite. The isopod doesn’t just replace the tongue physically, it also replaces it functionally. •Is this commensal or parasitic? What additional information would be necessary to evaluate this? Whale Coronula diadema • the whale’s skin is drawn into the spaces between the plates, permanently stitching the barnacle’s shell to the whale. •Is this commensal or parasitic? What additional information would be necessary to evaluate this? •

Parasitic symbioses

Remora –usually attaches to sharks, rays, turtles Sea lice •Parasitic copepods on hammerhead shark eye maggot •The eye maggot of sprat - a parasitic copepod attaches to eye Roundworms in fish muscle Parasitic castration •Common parasitic castrators include larval trematodes in snailsan individual parasitic castrator can usurp all the reproductive energy from a host

Animal behavior

Konrad Lorenz nobel prize for Lorenz, von Fritsch, and Tinbergen •During the first decades of this century research concerning animal behaviour was on its way to being stuck in a blind alley. •The vitalists believed in the instincts as mystical, wise and inexplicable forces inherent in the organism, governing the behaviour of the individual. •On the other hand reflexologists interpreted behaviour in an one-side mechanical way •behaviourists were preoccupied with learning as an explanation of all behavioural variations. •The way out of this dilemma was indicated by investigators who focused on the survival value of various behaviour patterns in their studies of species differences. Behaviour patterns become explicable when interpreted as the result of natural selection, analogous with anatomical and physiological characteristics.

The “cute response” –Lorenz theorized that certain "infantile features"—like big heads, large eyes, button noses, and round bodies—trigger a nurturing response in adults –Evolutionarily, this makes us more likely to care for our offspring, but our preference for cuteness is so strong it spills over to other species.

An Evolved Mickey Down Through the Ages Mickey’s Transition from Rat to Human Baby –His ears and head became much larger over time –His nose became less sharp and pointy –His eyes widened from two black dots to large, white, oval circles

Behavioral phylogeny – evolution of web type in spiders, “pushup” behavior in lizards, these behaviors seem to be in certain “branches” of animal phylogenetic trees showing that they evolve like other physical traits

Functions of communication •agonistic interaction: everything to do with contests and aggression between individuals. •courtship rituals: signals made by members of one sex to attract or maintain the attention of potential mate, or to cement a pair bond. ownership/territorial: signals used to claim or defend a territory, food, or a mate. •food-related signals: many animals make "food calls" that attract a mate, or offspring, or members of a social group generally to a food source. •alarm calls: signals made in the presence of a threat from a predator, allowing all members of a social group (and often members of other species) to run for cover, become immobile, or gather into a group to reduce the risk of attack. •metacommunications: signals that modify the meaning of subsequent signals. The best known example is the play face in dogs, which signals that a subsequent aggressive signal is part of a play fight rather than a serious aggressive episode.

Do animals have language? •They certainly have signals, as do humans •In the case of Jackdaws, their calls appear to be genetically determined since both German and Russian jackdaws give the same calls •Often these signals reflect moods •In humans there are also such “mimetic” signals such as yawning, smiling, wrinkling the brow In many instances animal signaling is much more sophisticated than in humans –Can humans using body language alone easily indicate to other humans that he/she is going home vs. going on a long journey –Jackdaws accomplish this via body language •One jackdaw will fly up to a tree branch and perch without causing the whole flock to also follow •But in other cases, the jackdaw will fly off to travel a distance and the others will follow. –Lorenz argues that the transmitting and receiving apparatus for animals can be more efficient than in humans transmitting and receiving apparatus for animals can be more efficient than in humans

•Example: dogs knowing when human is leaving room to do something mundane vs. to take the dog on a walk •Lorenz’s dog Tito would bite the posterior people who got on Lorenz’s nerves. •This would happen even if Tito was under the table and could not see faces and gestures. Dogs and horses that can count or do math –The dog watches the human intently and probably realizes when the right number of barks have been given –Horse paws ground, doesn’t appear to be looking at the human, but actually horses have acute peripheral vision and can detect even slightest movements –Interesting experiments –Showed dog cards with a math problem. On the back was the wrong problem. The owner of the dog saw the wrong problem and would always answer the problem incorrectly. Grey Parrot “Alex” –Irene Pepperberg research –model rival technique – –two trainers, one to give instructions, and one to model correct and incorrect responses and to act as the student's rival for the trainer's attention –parrot tries to reproduce the correct behavior. –Parrots and primates live in “societies” in the wild, this is different than a “herd” –Vocabulary of 150 words – names of 50 objects- could describe their colors, shapes and the materials they were made from. –He could ask for things—and would reject a proffered item and ask again if it was not what he wanted. – He understood, and could discuss, the concepts of “bigger”, “smaller”, “same” and “different”. –He could count up to six, including the number zero (and was grappling with the concept of “seven” when he died). Dog communication –recorded growls from 20 pet dogs in three different situations: a tug-of-war game with their owner, competing with another dog for a bone and growling at an approaching stranger. –played the recordings to 36 other dogs that had each been left to gnaw on a bone. Only those that heard the food-guarding growls tended to back off from the bone and stay away. –It seems that dog growls communicate context

Chapter 11

Between the

What is the ? •The intertidal zone is the area between the mean low and mean high tide. •By contrast, the subtidal zone is the area that is always submerged.

Problems Associated with the Intertidal Zone •Due to the exposure seen in the intertidal zone, face a variety of challenges, including: –Desiccation (water loss) – changes (can be extreme) – changes (can be extreme) –Interrupted feeding –Wave action and tides –Oxygen availability and build-up of CO2 –Limited space

Dessication •When exposed, organisms must deal with potential water loss. •Water loss is more pronounced on hot, dry days or windy days. •Organisms can deal with the potential water loss by hiding or “clamming up.” •Hiding may involve moving to a or an area with more moisture (motile organisms) •Hiding may also mean that some organisms only live in areas where moisture will remain when the tide is out (such as crevices in rocks or low spots in soft bottoms) “Clamming up” consists of closing shells or otherwise walling yourself off from the environment in an attempt to conserve moisture (such as an oyster closing its shell or a walling itself off using its operculum). •While moisture can be conserved in this way, there is a down side – no exchange of gases or feeding occurs.

Changing •The wide variety of temperatures that must be tolerated by organisms in the intertidal can be severe. •Imagine, one morning the temperature may hover around 60 degrees with an afternoon spent in temperatures exceeding 100 degrees.

Changing Salinity •Salinity can change dramatically due to temperatures or weather. •Normal salinity is around 33-35 ppt. •Estuarine salinity normally varies between 5 ppt and 30 ppt. •Organisms in the intertidal are normally (can tolerate a wide variety of ); subtidal organisms by contrast are normally (tolerate a very low range of salinities) •Salinity may begin at 20-25 ppt and climb dramatically during the day due to water loss when the intertidal is exposed. •Salinity may also drop during the day due to a sudden influx of freshwater provided by a passing thunderstorm.

•Now, imagine a salinity change of possibly 20 ppt or more combined with a temperature change of 40 degrees or more. Organisms must be extremely adaptable or perish.

Wave Action •Waves can be a problem for organisms that live in intertidal communities directly exposed to them. This action may dislodge organisms from their . •Rocky shorelines are often exposed to significant wave action.

Oxygen Availability •Oxygen can be exhausted if an organism “clams up” during low tide. •It can also be exhausted in tidal pools if many organisms have sought refuge there. •Carbon dioxide can also build up to toxic levels when organisms are not able to exchange these gases with their environments.

Limited Space •In some intertidal communities, proper space may be limited. •This is particularly true in rocky intertidal environments where the amount of surface area is limited.

Zonation in the Intertidal

•Zonation in the intertidal consists of upper intertidal (most exposed) to middle and lower intertidal (least exposed).

• will be greater in the lower intertidal because it is the least “severe” of the zones since it is exposed to a lesser degree than the upper intertidal.

•The lower intertidal is always more species rich for the same reason (although species will vary greatly by location).

Competition

•Some organisms are better competitors than others are will exclude other organisms if the is left undisturbed.

•Top in any community that have the ability to change community composition significantly are known as keystone predators.

***Intertidal experiments: be prepared to interpret data from intertidal experiments***

Chapter 12

Estuaries

Types of Drowned river valleys •Bar built - –Built by the accumulation of sediments into bars or barrier –Ex: North Carolina (seen in lower part of the photo to the right near Hatteras) • – –Deep channels cut in the coastal zone as a result of retreating glaciers –Ex: Alaska, Norway –Puget is a type of glacial Puget Sound •Over millions of years canadian glaciers formed and retreated at least 4 times •20,000 years ago, glacier 1 mile deep •After glaciers was a large lake •Sea levels eventually rose and filled the estuary •http://exhibits.pacsci.org/puget_sound/graphics/ps_glaciationsm.mov

Development of Estuaries •Estuaries are the best developed in areas where the is flat and the is wide. i.e. East US •The opposite is true in areas with steep, narrow continental shelves and coastal plains, i.e., Washington, west coast US

Physical Characteristics of Estuaries •Salinity- –Can vary from 5 – 30 ppt –Salinity varies according to distance from saltwater (tides) or freshwater (river) input –Can also vary as a result of –Depth also contributes to salinity –The diagram to the right illustrates that the salinity is not uniform (saltwater is heavier and sinks below freshwater) – this is known as a salt wedge

Dealing with Salinity Changes •Organisms in the estuaries are normally euryhaline (can tolerate a wide variety of salinities) •Some are osmoregulators that have mechanisms for keeping their internal concentration stable •Others are osmoconformers with internal concentrations that vary with their surroundings

Dealing with Salinity Changes •Flowering plants in the estuarine community must either expel excess solutes (such as Spartina and trees using salt glands) or concentrating solutes in specific tissues (such as in Salicornia, a.k.a. “pickleweed” or glasswort

Mudflat ecology •Only the upper few mm of the sediment is oxygenated, the rest is anoxic and rich in hydrogen sulfide •Very little hard substrate which is preferred by many organisms •Burrowing organisms can oxygenate their environment and also provide oxygen for other organisms that live in their burrows • can provide substrate and also oxygenate the sediment

•“Oyster reefs” are found in many estuaries •These congregations of oysters are the platform on which dozens of other species grow and thrive •The oysters provide the only hard substrate in many estuaries

•Saltmarsh •The vegetated area of an estuary •Spartina species predominate near the water in most estuaries Zostera marina Eel grass seagrasses Eelgrass distribution in puget sound – 33% reduced Eelgrass increases the number of species able to live in the estuary by acting as substrate and cover for organisms, they also oxygenate the sediment Spartina cordgrass in Puget sound are an Spartina is native to the East Coast and Europe Invaded Puget sound and elsewhere in the west coast No native species can colonize the but invasive spartina was able to take over radically altering the estuary

Worldwide Distribution of Saltmarshes and Mangrove Forests In the tropics Mangrove trees are found instead of saltmarsh grasses

This version does not include deep sea ecology which will be covered on 4/16 and 4/18