AN INTRODUCTION TO CORAL REEFS

Overview Coral reefs are among the most diverse and productive communities on Earth. They are found in the warm, clear, shallow waters of tropical oceans worldwide. Reefs have functions ranging from providing food and shelter to fish and invertebrates to protecting the shore from erosion. Through symbiosis with unicellular algae (zooxanthellae , reef!building corals are the source of primary production in reef communities (Richmond "##$ . %iologically active compounds produced by reef dwelling organisms possess antimicrobial and antiviral activity (&an 'lstyne and (aul "#)) . These compounds may be important sources for natural product based drugs and medicines. *n addition, revenue from tourists attracted to the beauty of coral reefs can be a significant source of income for human communities in these areas. +nfortunately, as our understanding of corals and coral reefs increases, it becomes apparent that the effects of the human population on these communities may be increasing as well. Reef!building (or hermatypic corals are of the order ,cleractinia in the class 'nthozoa of the phylum Cnidaria. 'pproximately -,... species of 'nthozoans exist, all of them marine ((echeni/ "##" . Coral reefs of the western (acific are much more diverse than those of the 'tlantic and Caribbean. There are up to 012 more genera and )12 more species of corals in (acific waters (3il/inson "#)0 . Reefs are formed by calcium carbonate produced by tiny coral polyps. 3hile corals are the chief architects of reef structure, they are not the only builders. Coralline algae cement various corals together with compounds of calcium, and other organisms such as tube worms and molluscs donate their hard s/eletons (Cousteau "#)1 . Together these organisms construct many different types of reefs. Reefs are important land builders in tropical areas, forming islands and altering continental shorelines (4oreau et al. "#0# . ' coral colony may consist of thousands of polyps. (olyps are typically carnivorous, feeding on small particles floating in the water. 5owever, endosymbiotic algae /nown as zooxanthellae provide important sources of nutrition to hermatypic corals (Rowan and (owers "##" . Corals reproduce both sexually and asexually. 'n entire colony many meters in diameter can start out as a single polyp. %ecause many coral reef organisms can tolerate only a narrow range of environmental conditions, reefs are sensitive to damage from environmental changes (Richmond "##$ . Corals are susceptible to diseases and bleaching. 'lso, dramatic natural events such as hurricanes can damage coral reefs. *n addition, many problems to reefs are anthropogenic (human caused . %ecause of the important ecological and economic roles coral reef communities fulfill, an understanding of the stresses and dangers to the reefs is necessary. 6ortunately, many of the human induced hazards to coral reefs can be remedied (Richmond "##$ . Anatomy of Corals Coral reefs consist of many diverse species of corals. These corals in turn are made up of tiny organisms called polyps. The structure of the polyps and the s/eleton of the coral is a rather simple combination. ' polyp is made up or two cell layers7 the epidermis and the gastrodermis. The non!tissue layer between the gastrodermis and the epidermis is called the mesoglea. The polyp contains mesentery filaments, which contain nematocysts used in food capture, a pharynx, endothecal dissepiments (horizontal layers of s/eletal material and the columella (the central axis of the corallite found below the mouth . The corallite is the part of the s/eleton deposited by one polyp. The s/eletal wall around each polyp is called the theca. 8ther structures include the calice (the upper opening of the corallite , the coenosarc (the coral tissue that stretches over the surface of the coral between the polyps , the coenosteum (the s/eletal material around the corallites , and the corallum, which is the s/eleton of the coral. The coral anatomy also includes calcareous plate!li/e structure /nown as septa. The septa radiate from the wall to the center of the corallite. There are two types of septa7 insert septa which lie below the corallite wall and exsert septa which protrude above the corallite wall. Corals are of two types7 perforate and imperforate. (erforate corals have porous s/eletons with connections between the polyps through the s/eleton. *mperforate corals have solid s/eletons. 9any corals have different growth forms. They can be plocoid as in Tubastrea coccinea (orange cup coral and Favia fragum (golf ball coral . They can also be meandroid in which corallites form a series within the same walls, as in the species Dendrogyra cylindrus (pillar coral . 8ther growth forms include cocoid, spherical shaped and phalecoid, as in Eusmilia fastigiata. Categories and Stru ture of Coral Reefs There are three basic /inds of coral reefs in the Caribbean7 fringing reefs, barrier reefs and atolls. 6ringing reefs are coral reefs that grow in shallow waters and border the coast closely or are separated from it by a narrow stretch of water. 6ringing reefs consist of several zones that are characterized by "

their depth, the structure of the reef, and its plant and animal communities. These regions include the reef crest (the part of the reef the waves brea/ over , the fore reef (the region of medium energy , and the spur and groove or buttress zone (the region of coral growth which includes rows of corals with sandy canyons or passages between each row . %arrier reefs are reefs that are separated from land by a lagoon. These reefs grow parallel to the coast and are large and continuous. %arrier reefs also include regions of coral formation that include the zones found in fringing reefs along with patch reefs (small reefs , bac/ reefs (the shoreward side of the reef , as well as ban/ reefs (reefs that occur on deep bottom irregularities . Coral reefs also include reef flats (the are of the reef not exposed , the reef crest, which runs parallel to the coast and is protected from waves, and a coral terrace (a slope of sand with isolated coral pea/s . These features are followed by another coral terrace and a vertical drop into deeper waters. The third type of coral reefs are atolls. 'tolls are annular reefs that develop at or near the surface of the sea when islands that are surrounded by reefs subside. 'tolls separate a central lagoon and are circular or sub!circular. There are two types of atolls7 deep sea atolls that rise from deep sea and those found on the continental shelf. !oo"ant#ellae :ooxanthellae are unicellular yellow!brown (dinoflagellate algae which live symbiotically in the gastrodermis of reef!building corals (4oreau et al., "#0# . *t is the nutrients supplied by the zooxanthellae that ma/e it possible for the corals to grow and reproduce ;uic/ly enough to create reefs. :ooxanthellae provide the corals with food in the form of photosynthetic products. *n turn, the coral provides protection and access to light for the zooxanthellae. %ecause of the need for light, corals containing zooxanthellae only live in ocean waters less than ".. meters deep (4oreau et al., "#0# . They also only live in waters above <. degrees Celsius and are intolerant of low salinity and high turbidity (4oreau et al., "#0# . *t was once believed that all zooxanthellae were the same species, Symbiodinium microadriaticum (Rowan and (owers, "##" . 5owever, recently, zooxanthellae of various corals have been found to belong to at least ". different algal taxa. *nterestingly, zooxanthellae found in closely related coral species are not necessarily closely related themselves, and zooxanthellae found in distantly related coral species may, in fact, be closely related (Rowan and (owers, "##" . This suggests that coral and zooxanthellae evolution did not occur in permanently associated lineages. Rather, symbiotic recombination probably shaped the evolutionary process, allowing both symbionts to evolve separately. Coral Feeding Corals can obtain food in a variety of ways. Reef!building corals rely on the photosynthetic products of zooxanthellae for the ma=ority of their nutrients. 5owever, corals also capture zooplan/ton for food. Corals are suspension feeders. They utilize two main methods of prey capture7 nematocyst adhesion and mucus entrapment (,ebens and >ohnson, "##" . ?ematocysts on the tentacles and mesentarial filaments can be used to sting prey and move it into the mouth. ,ome corals will trap prey in stic/y mucus on their tentacles and move the prey into the mouth using the mucus and cilia. (rey size is correlated to polyp size (,ebens and >ohnson, "##" . 9ost corals feed at night. This may be because night is when the zooplan/ton travel into the water column and become available for capture. @eeping the tentacles retracted during the day may also help corals avoid predation, protect themselves from +& light, and avoid shading their zooxanthellae. (rey can be brought into contact with the tentacles by direct interception (the flow of water ma/es the particles impact the tentacles , by inertial impaction (momentum of dense particles ma/es them deviate from the water flow and impact the tentacles , and by gravitational deposition (gravity causes heavy particles to fall into the tentacles (,ebens and >ohnson, "##" . Regardless of the method of capture, prey is moved through the mouth, down the pharynx (gullet , to the gastrovascular cavity for digestion. Coral Re$rodu tion Corals exhibit sexual and asexual reproduction. The coral colony expands in size by budding. %udding may be intratentacular, in which the new bud forms from the oral discs of the old polyp, as in Diploria, or extratentacular in which the new polyp forms from the base of the old polyp, as in Montastraea cavernosa. ' common type of asexual reproduction in corals is by fragmentation. %ro/en pieces of corals that land on a suitable substrate may begin growing and produce a new colony. This type of reproduction is common in branching corals li/e Acropora cervicornis in which a positive correlation was found between fragment size and survival. <

9any coral species mass spawn. 3ithin a <A hour period, all the corals from one species and often within a genus release their eggs and sperm at the same time. This occurs in related species of Montastraea, and in other genera such as Montipora, Platygra, Favia, and Favites (3allace, "##A . *n some Montastraea and Acropora species, the eggs and sperm are released in a sac/. They float to the surface where they separate and fertilization ta/es place. *ntraspecies is common but mass spawning raises the possibility of hybridization by congeneric species (3allace, "##A . The zygote develops into larvae called planula which attaches itself to a suitable substrate and grows into a new colony. ,ome species of coral brood their larvae. The sperm fertilizes the egg before both are released from the coral. The larvae float to the top, settle, and become another colony. ,pecies of Acropora release brooded larvae. Coral Diseases 6our coral conditions have been identified as diseases7 white band disease (3%B , blac/ band disease (%%B , bacterial infection, and shut down reaction (Richmond "##$ . They are also susceptible to tumors and parasitic worms. These maladies are all stress related, and anthropogenic stresses can increase a coralCs susceptibility to these diseases. Biseases such as %%B and 3%B actually /ill coral tissue while advancing in a band around the coral and leaving the white coral s/eleton behind. Edmunds ("##" stated that %%B, caused by cyanophyte (hormidium corallyticum, may have a role in maintaining coral diversity because it is most prevalent in coral species that form large colonies and provide a structural framewor/ for the reef. 3hen %%B /ills part of these colonies, the s/eleton is available to be colonized by other coral species recruits. 5owever, after <1 months, there were not coral recruits among corals infected by %%B (Edmunds "##" . 3%B, which is believed to be caused by a bacteria pathogen yet un/nown, has much of the same effect on corals, leaving behind a white, lifeless coral s/eleton. 4ladfelter ("#)< does not see 3%B as being beneficial to reefs. 5e feels that 3%B destroys the reef structure because the dead coral s/eleton brought about by the algae is colonized by algae, invertebrates, gastropods, and boring cloinid sponges that wor/ to wea/en the coral s/eletons ma/ing them more susceptible to brea/age during storms. This situation is also probable for %%B. The exact method by which the diseases are transmitted are un/nown. Even though healthy corals may get %%B through contact with an infected coral, diseased corals are not aggregated naturally on the reef and can be separated by great distances. Thus, it is also suggested that %%B can be spread by currents through trichomes that come off infected colonies and land on other colonies (Edmunds "##" . Corals under stress are more li/ely to be infected, and %%B has a higher rate of infection in warmer water. Thus seasonal temperatures affect the spread of %%B, and also any anthropogenic warming of water temperatures may increase %%B. %%B was also found to be more abundant near anthropogenic disturbances. 5owever, 3%B has not been found to be related to anthropogenic disturbances (4ladfelter "#)< . Corals may also be affected by diseases indirectly. Biseases that infect other reef inhabitants may affect ecology of corals. *n "##$, a newly encountered pathogen was found to affect coralline algae in the (acific 8cean (Dittler and Dittler "##1 . This pathogen also leaves the coralline algae s/eleton white as it progresses in a orange band, destroying the algae. Coralline algae help the coral reef community by cementing together sand, coral fragments, and other debris into a suitable hard substrate for the establishment of coral colonies and by absorbing wave energy in the outer reef rim that would otherwise erode the shoreline (Dittler and Dittler "##1 . 'nother disease which caused a Caribbean!wide mortality in the reef dwelling sea urchin Biadema antillarum was also shown to indirectly affect coral reefs (Dessions "#)) . The sea urchin functions as both a grazer on algae that can otherwise smother coral and as a bioeroder of corals as it feeds on them (Dessions "#)) . Coral %lea #ing Coral bleaching is the whitening of coral colonies due to the loss of symbiotic zooxanthellae from the tissues of polyps. This loss exposes the white calcium carbonate s/eletons of the coral colony. Corals naturally lose less than .."2 of their zooxanthellae during processes of regulation and replacement (%rown and 8gden, "##$ . 5owever, adverse changes in a coralCs environment can cause an increase in the number of zooxanthellae lost. There are a number of stresses or environmental changes that may cause bleaching including disease, excess shade, increased levels of ultraviolet radiation, sedimentation, pollution, salinity changes, and increased temperatures.

Corals tolerate a narrow temperature range between <1 degrees Celsius and <# degrees Celsius depending on location. Corals bleach in response to prolonged temperature change and not due to rapidly fluctuating temperatures. Dab experiments show that corals bleach when water reaches a constant $< degrees Celsius (%rown and 8gden, "##$ . Experiments have shown that an increase in +& light causes coral bleaching. +& light experienced by corals can increase in calm waters. The amount of mycosporine!li/e amino acids in a coralCs tissues helps to determine how much +& it can withstand without bleaching (4leason, "##$ . The exact mechanism by which corals bleach or the trigger that induces bleaching is un/nown. 5owever, there are a number of hypotheses that attempt to answer this ;uestion. Daboratory experiments have shown that the zooxanthellae are released into the gut of the polyp and then are expelled from the polyp through the mouth (%rown and 8gden, "##$ . 5owever, this has not been observed in nature. 'nother hypothesis is that stressed corals give algae fewer nutrients and thus the algae leave the polyp. 'lgae may produce oxide toxicity under stress, and these toxins may affect the polyps. T#reats to Coral Reefs Corals and coral reefs are extremely sensitive. ,light changes in the reef environment may have detrimental effects on the health of entire coral colonies. These changes may be due to a variety of factors, but they generally fall within two categories7 natural disturbances and anthropogenic disturbances. 'lthough natural disturbances may cause severe changes in coral communities, anthropogenic disturbances have been lin/ed to the vast ma=ority of decreases in coral cover and general colony health when coral reefs and humans occur together. 8ne of the greatest threats to coral reefs is human expansion and development. 's development continues to alter the landscape, the amount of freshwater runoff increases. This terriginous runoff may carry large amounts of sediment from land!clearing areas, high levels of nutrients from agricultural areas or septic systems, as well as many pollutants such as petroleum products or insecticides. 3hether it is direct sedimentation onto the reef or an increase in the turbidity of the water due to eutrophication, decreases in the amounts of light reaching corals may cause bleaching (%rown and 8gden "##$ . *n addition, increases in the amounts of nutrients enhance the growth of other reef organisms such as sponges which may outcompete the corals for space on crowded reefs. *n addition to runoff, outflows from water treatment plants and large power plants are the cause of much damage to coral reefs. ,ewage treatment facilities greatly increase the nutrient levels surrounding their outflow pipes while large power plants alter water temperatures by discharging extremely hot water into the coastal waters. 's with all these factors, the basis for the continued degradation of coral reefs is the increasing size of the human population. 's this population increases, so does the harvest of resources from the sea. Bue to overfishing, reef fish populations have been greatly decreased in some areas of the world. The removal of large numbers of reef fish has caused the coral reef ecosystems to become unbalanced and allowed more competitive organisms, such as algae, which were once controlled by large fish populations, to become dominant on reefs in many regions. Bue to decreased yields, fishermen have been forced to change their methods in order to catch enough fish to sustain their needs. *n some areas this means fish traps with small mesh diameters which catch even the small =uvenile fish. *n other areas of the world, the use of explosives or poisons has become ;uite common (Richmond "##$ . ?ot only do these practices /ill all fish in the affected areas, but they also severely damage the corals in these areas. Corals are also very popular as decorations. 8ften, when people vacation in tropical locations surrounded by beautiful reefs they want to ta/e coral souvenirs home. *n order to do this, they either collect pieces of coral themselves or buy pieces from a EcuriosE shop. These shops receive their corals from commercial collectors who select well developed colonies which will ma/e them the most money. This is very damaging because a large amount of the most healthy corals are selected. Coral reefs also receive much damage from both commercial and private vessels. The lea/age of fuels into the water and the occurrences of spills by large tan/ers are extremely damaging to local corals. %oat anchors are also very damaging to reefs by brea/ing and destroying entire colonies. The grounding of large sea!going vessels also results in large sections of coral reefs being destroyed. *t has also been found that the anti!fouling bottom paints used by many boats contribute to the formation of toxic concentrations of Tributyl tin and several other chemical compounds which may be harmful to corals or other coral species (Besrosiers, personal communication .

,ince most corals mass spawn and produce floating gametes, pollutants and toxins on the surface can effect coral reproduction and development for a large area. Therefore, much care must be ta/en to reduce or prevent the spillage and lea/age of contaminants into the water as well as to improve cleanup procedures of such accidents. 'lthough much of the coral reefs degradation is directly blamed on human impact, there are several natural disturbances which cause significant damage to coral reefs. The most recognized of these events are hurricanes, or typhoons, which bring large and powerful waves to the tropics. These storm waves cause large corals to brea/ apart and scatter fragments about the reefs. 'fter the storm, these slow growing corals might easily be overgrown by ;uic/er growing algae. *n addition, these storms generally bring heavy rain which increases runoff and sedimentation. 'nother common threat to coral populations of the (acific 8cean is the crown!of!thorns, Acanthaster planci. A. planci is a large starfish which feeds on corals by extruding its stomach out onto the coral to digest the living tissue layer (%ir/eland "#)# . These predators have had serious effects on the coral populations in many regions of the (acific, and Acanthaster brea/outs have been lin/ed to regions of increased development and eutrophication (%ir/eland "#)# . 9any other predators such as fish and gastropods also are /nown to cause damage to coral colonies, but these generally do not compare to the drastic effects which A. planci has on coral populations. There are a great number of threats to coral reefs, and most of the threats can be attributed either directly or indirectly to humans. 3or/ must be done ;uic/ly to protect our threatened resources. The list of solutions to the many coral reef problems is extensive. These range from better methods of development in order to decrease runoff to the installation of permanent moorings at heavily used anchorage sites. 3hatever the solutions, there always needs to be ade;uate enforcement to ensure proper techni;ues are being followed. +nfortunately, enforcement has not been great enough in the past and will probably not be in the future. Therefore, the education and cooperation of people throughout the world is necessary if coral reefs are to survive. Animals Asso iated wit# Coral Reefs Coral reefs provide habitats for a large variety of organisms. These organisms rely on corals as a source of food and shelter. ,ome organisms that use corals through mutualism, commensalism and parasitism are within the taxonomic groups (orifera, (olychaeta, 4astropoda, Crustacea, Echinodermata and (isces. ,ponges ((orifera are found inhabiting cavities in the reef. They remove small chips of calcium carbonate from corals. These sponges, such as Cliona, cause bioerosion in corals. ,ponges inhabit corals for the purpose of protection from predators. (olychaetes such as 5ermodice carunculata and 4astropods in the family Trochidae depend on corals for food. They feed on corals such as (orites and 'garicia. Becapod crustaceans such as shrimps and crabs depend on corals for shelter. Fantid crabs form cavities in the coral Acropora palmata. 6ish also depend on corals for protection against predators. 8ne such is the parrot fish (,caridae . Echinoderms such as 'canthaster planci are coral predators. This crown!of!thorns starfish relies on corals for food. There are many other species of fungi, sponges, sea worms, crustaceans and molluscs that bore into coral s/eletons. 8ther organisms that inhabit the coral reefs include sea urchins, =ellyfish, oysters, clams, turtles, and sea anemones.
Literature Cited
%ir/eland, C. "#)#. The 6austian traits of the Crown!of!Thorns ,tarfish. 'merican ,cientist, 007"1A!"-$. %rown, %.E., 8dgen, >.C. "##$. Coral %leaching. ,cientific 'merican, <-#7-A!0.. Cousteau, >. G. "#)1. The 8cean 3orld. 5arry ?. 'brams, *nc., ?ew Gor/, ?G, pp. "0A!"01. Edmunds, (.>. "##". Extent and Effect of %lac/ %and Bisease on a Caribbean Reef. Coral Reefs, ".7"-"!"-1. 4ladfelter, 3.%. "#)<. 3hite %and Bisease in Acropora palmata7 *mplications for the ,tructure and 4orwth of ,hallow Reefs. %ulletin of 9arine ,cience, $<(< 7-$#! -A$. 4leason, B.6. "##$. Bifferential Effects of +ltraviolet Radiation on 4reen and %rown 9orphs of the Caribbean Coral Porites astreoides. >ournal of Dimnology and 8ceanography, $)(0 7"A1<!"A-$. 4oreau, T. 6., ?. *. 4oreau, and T. >. 4oreau. "#0#. Corals and Coral Reefs. ,cientific 'merican, <A"7"<A!"$-. Dessions, 5.'. "#)). 9ass 9ortality of Diadema antillarum in the Caribbean7 3hat 5ave 3e learnedH. 'nnual Review of Ecology and ,ystematics, "#7$0"!$#$. Dittler, 9.9., B.,. Dittler. "##1. *mpact of CD8B (athogen on (acific Coral Reefs. ,cience, <-07"$1-!"$1#. (echeni/, >. '. "##". %iology of the *nvertebrates. 3m. C. %rown (ublishers, Bubu;ue, *', pp. #"!#<. Richmond, R. 5. "##$. Coral Reefs7 (resent (roblems and 6uture Concerns Resulting from 'nthropogenic Bisturbance. 'merican :oologist, $$71<A!1$-. Rowan, R. and B. '. (owers. "##". ' 9olecular 4enetic Classification of :ooxanthellae and the Evolution of 'nimal!'lgal ,ymbioses. ,cience, &ol. <1"7"$A)!"$1". ,ebens, @.(., >ohnson, '.,. "##". Effects of 3ater 9ovement on (rey Capture and Bistribution of Reef Corals. 5ydrobiologia, <<-7#"!".". &an 'lstyne, @. D. and &. >. (aul. "#)). The Role of ,econdary 9etabolites in 9arine Ecological *nteractions. (roceedings of the -th *nternational Coral Reef ,ymposium, 'ustralia, "#)), &ol. ". 3allace, C.C., 3illis, %.D. "##A. ,ystematics of the Coral 4enus Acropora7 *mplications of ?ew %iological 6indings for ,pecies Concepts. 'nnual Review of Ecology and ,ystematics, <17<$0!<-<. 3il/inson, C. R. "#)0. *nterocean Bifferences in ,ize and ?utrition of Coral Reef ,ponge (opulations. ,cience, &ol. <$-7"-1A!"-10.

Tmb."syl

Tropical 9arine %iology (%$$< ,pring <.." >ohn %ishop

S&LLA%US Course7 Tropical 9arine %iology (%$$< S #edule' 936 ""7$.!"<7<., 6 "7$1!A7<1, ,pring brea/ field trip Instru tor7 >ohn 3. %ishop, ,!"." 4ottwald ,cience Center, ().A <)#!)<$., =bishopIrichmond.edu, www.science.urich.eduJKbishop Offi e #ours7 936 ""7..!""7$. '.9., TR A7..!17.. (.9. and by appointment O()e tives7 The course is designed to ac;uaint the student with the ecology of tropical marine organisms with an emphasis on coral reefs. Readings7 The primary text is ?yba//en, >.3. <..". 9arine %iology7 'n Ecological 'pproach. 1th ed., 'ddison 3esley Dongman, *nc., ?.G. 'dditional reading assignments from @aplan, E. "#)<. 6ield 4uide to Coral Reefs of the Caribbean and 6lorida. 5oughton!9ifflin Co., %oston and =ournal articles. *edagogy7 The course relies on your active participation. *t includes a variety of pedagogical approaches ! class discussions, lectures, field observations, presentation of observations, oral and written reports, and tests. S #edule7 3ee/ >an. "1 << <# 6eb. 1 "< "# <9ar. 1 "< "# <'pril < # "<$ 9ay 1 Topic 9arine Environment (?7 "!$0 " Coral Reefs (?7 $0.!A.., @7 -!"<. 6ishes (?7 #A!".), ""1!""#, A..!A".L @7 <.-!<A" Coral Reef Bynamics (?7 A".!A"- , >ournal articles Test "L ?ational ';uarium, %altimore, 9B. ,eagrasses (?7 <".!<"), @7"$!<1 , >ournal articles 9angroves (?7 A"-!A$$ , >ournal articles Test <L (ractical %elize 6ield Trip 6ield Trip ,ummary (3 5uman *mpacts (?7 A1)!A#$ ,ubmit 6ield Trip Report (6 >ournal articles (resent 8ral 'utecology Report (resent 8ral 'utecology Report 3ritten 'utecology Report (6 6inal Exam (#!"< ?oon

". Reading assignments from ?yba//en (? and @aplan (@ are in parentheses. Reading assignments from =ournal articles will be announced. ,tudents will be given ;uestions on readings about a wee/ before class discussions of the readings. *oli ies7 ,tudents are expected to attend and participate in all class activities, and to ta/e tests and submit assignments as scheduled. 4rades for assignments submitted after the due dates will be decreased by 1 pts per day to a minimum grade of 11 if the assignment is submitted by the last day of class and . if the assignment is not submitted by the last day of class. The honor code pledge should be included on written assignments.