PROKARYOTIC & EUKARYOTIC Part of our definition/description of what it means to be a i!

in" thin" on Earth inc udes the assertion that i!in" thin"s are made of ce s and ce products# In other words$ we consider the ce to be a prett% fundamenta structura aspect of ife# Ce s in our wor d come in two basic t%pes$ pro&ar%otic and eu&ar%otic# 'Kar%ose' comes from a (ree& word which means '&erne $' as in a &erne of "rain# In bio o"%$ we use this word root to refer to the nuc eus of a ce # 'Pro' means 'before$' and 'eu' means 'true$' or '"ood#' )o 'Pro&ar%otic' means 'before a nuc eus$' and 'eu&ar%otic' means 'possessin" a true nuc eus#' This is a bi" hint about one of the differences between these two ce t%pes# Pro&ar%otic ce s ha!e no nuc ei$ whi e eu&ar%otic ce s do ha!e true nuc ei# This is far from the on % difference between these two ce t%pes$ howe!er#

*ere+s a simp e !isua comparison between a pro&ar%otic ce and a eu&ar%otic ce ,

This particu ar eu&ar%otic ce happens to be an anima ce $ but the ce s of p ants$ fun"i and protists are a so eu&ar%otic#

-espite their apparent differences$ these two ce t%pes ha!e a ot in common# The% perform most of the same &inds of functions$ and in the same wa%s# .oth are enc osed b% p asma membranes$ fi ed with c%top asm$ and oaded with sma structures ca ed ribosomes# .oth ha!e -/A which carries the archi!ed instructions for operatin" the ce # And the simi arities "o far be%ond the !isib e00ph%sio o"ica % the% are !er% simi ar in man% wa%s# 1or e2amp e$ the -/A in the two ce t%pes is precise % the same &ind of -/A$ and the "enetic code for a pro&ar%otic ce is e2act % the same "enetic code used in eu&ar%otic ce s# )ome thin"s which seem to be differences aren+t# 1or e2amp e$ the pro&ar%otic ce has a ce wa $ and this anima ce does not# *owe!er$ man% &inds of eu&ar%otic ce s do ha!e ce wa s# -espite a of these simi arities$ the differences are a so c ear# It+s prett% ob!ious from these two itt e pictures that there are two "enera cate"ories of difference between these two ce t%pes, si3e and comp e2it%# Eu&ar%otic ce s are much ar"er and much more comp e2 than pro&ar%otic ce s# These two obser!ations are not unre ated to each other# If we ta&e a c oser oo& at the comparison of these ce s$ we see the fo owin" differences, Eu&ar%otic ce s ha!e a true nuc eus$ bound b% a doub e membrane# Pro&ar%otic ce s ha!e no nuc eus# The purpose of the nuc eus is to se4uester the -/A0re ated functions of the bi" eu&ar%otic ce into a sma er chamber$ for the purpose of increased efficienc%# This function is unnecessar% for the pro&ar%otic ce $ because its much sma er si3e means that a materia s within the ce are re ati!e % c ose to"ether# Of course$ pro&ar%otic ce s do ha!e -/A and -/A functions# .io o"ists describe the centra re"ion of the ce as its 'nuc eoid' 50oid6simi ar or imitatin"7$ because it+s prett% much where the -/A is ocated# .ut note that the nuc eoid is essentia % an ima"inar% 'structure#' There is no ph%sica boundar% enc osin" the nuc eoid# Eu&ar%otic -/A is inear8 pro&ar%otic -/A is circu ar 5it has no ends7# Eu&ar%otic -/A is comp e2ed with proteins ca ed 'histones$' and is or"ani3ed into chromosomes8 pro&ar%otic -/A is 'na&ed$' meanin" that it has no histones associated with it$ and it is not formed into chromosomes# Thou"h man% are s opp% about it$ the

term 'chromosome' does not technica % app % to an%thin" in a pro&ar%otic ce # A eu&ar%otic ce contains a number of chromosomes8 a pro&ar%otic ce contains on % one circu ar -/A mo ecu e and a !aried assortment of much sma er circ ets of -/A ca ed 'p asmids#' The sma er$ simp er pro&ar%otic ce re4uires far fewer "enes to operate than the eu&ar%otic ce # .oth ce t%pes ha!e man%$ man% ribosomes$ but the ribosomes of the eu&ar%otic ce s are ar"er and more comp e2 than those of the pro&ar%otic ce # Ribosomes are made out of a specia c ass of R/A mo ecu es 5ribosoma R/A$ or rR/A7 and a specific co ection of different proteins# A eu&ar%otic ribosome is composed of fi!e &inds of rR/A and about ei"ht% &inds of proteins# Pro&ar%otic ribosomes are composed of on % three &inds of rR/A and about fift% &inds of protein# The c%top asm of eu&ar%otic ce s is fi ed with a ar"e$ comp e2 co ection of or"ane es$ man% of them enc osed in their own membranes8 the pro&ar%otic ce contains no membrane0bound or"ane es which are independent of the p asma membrane# This is a !er% si"nificant difference$ and the source of the !ast ma9orit% of the "reater comp e2it% of the eu&ar%otic ce # There is much more space within a eu&ar%otic ce than within a pro&ar%otic ce $ and man% of these structures$ i&e the nuc eus$ increase the efficienc% of functions b% confinin" them within sma er spaces within the hu"e ce $ or with communication and mo!ement within the ce # E2amination of these differences is interestin"# As mentioned abo!e$ the% are a associated with ar"er si3e and "reater comp e2it%# This eads to an important obser!ation# Yes$ these ce s are different from each other# *owe!er$ the% are c ear % more a i&e than different$ and the% are c ear % e!o utionari % re ated to each other# .io o"ists ha!e no si"nificant doubts about the connection between them# The eu&ar%otic ce is c ear % de!e oped from the pro&ar%otic ce # One aspect of that e!o utionar% connection is particu ar % interestin"# :ithin eu&ar%otic ce s %ou find a rea % fascinatin" or"ane e ca ed a mitochondrion# And in p ant ce s$ %ou+d find an additiona fami % of or"ane es ca edp astids$ the most famous of which is the renowned ch orop ast# ;itochondria 5the p ura of mitochondrion7 and ch orop asts

a most certain % ha!e a simi ar e!o utionar% ori"in# .oth are prett% c ear % the descendants of independent pro&ar%otic ce s$ which ha!e ta&en up permanent residence within other ce s throu"h a we 0&nown and !er% common phenomenon ca ed endos%mbiosis# One structure not shown in our pro&ar%otic ce is ca ed a mesosome# /ot a pro&ar%otic ce s ha!e these# The mesosome is an e aboration of the p asma membrane00a sort of rosette of ruff ed membrane intrudin" into the ce # This dia"ram shows a trimmed down pro&ar%otic ce $ inc udin" on % the p asma membrane and a coup e of mesosomes# A mitochondrion is inc uded for comparison,

The simi arities in appearance between these structures are prett% c ear# The mitochondrion is a doub e0membrane or"ane e$ with a smooth outer membrane and an inner membrane which protrudes into the interior of the mitochondrion in fo ds ca ed cristae# This membrane is !er% simi ar in appearance to the pro&ar%otic p asma membrane with its mesosomes# .ut the simi arities are a ot more si"nificant than appearance# .oth the mesosomes and the cristae are used for the same function, the aerobic part of aerobic ce u ar respiration# Ce u ar respiration is the process b% which a ce con!erts the raw$ potentia ener"% of food into bio o"ica % usefu ener"%$ and there are two "enera t%pes$ anaerobic 5not usin" o2%"en7 and aerobic 5re4uirin" o2%"en7# In practica terms$ the bi" difference between the

two is that aerobic ce u ar respiration has a much hi"her ener"% %ie d than anaerobic respiration# Aerobic respiration is c ear % the e!o utionar% offsprin" of anaerobic respiration# In fact$ aerobic respiration rea % is anaerobic respiration with additiona chemica se4uences added on to the end of the process to a ow uti i3ation of o2%"en 5a !er% common e!o utionar% pattern00addin" new parts to o d s%stems7# )o it+s prett% reasonab e of bio o"ists to thin& that a mitochondrion e!o !ed from a once0independent aerobic pro&ar%otic ce which entered into an endos%mbiotic re ationship with a ar"er$ anaerobic ce # )o is there an% rea e!idence that the distant ancestors of mitochondria were independent ce s< =uite a ot$ actua %# And of a !er% con!incin" t%pe# ;itochondria 5and ch orop asts$ for that matter7 ha!e their own "enetic s%stems# The% ha!e their own -/A$ which is not dup icated in the nuc eus# That -/A contains a number of the "enes which are necessar% to ma&e the materia s needed for aerobic ce u ar respiration 5or photos%nthesis$ in the case of the ch orop ast7# ;itochondria and ch orop ast -/A mo ecu es are na&ed and circu ar$ i&e pro&ar%otic -/A# These or"ane es a so ha!e their own popu ation of ribosomes$ which are sma er and simp er than the ribosomes out in the "enera c%top asm# ;itochondria and ch orop asts a so di!ide on their own$ in a manner simi ar to the binar% fission of pro&ar%otic ce s# Then there+s that interestin" outer membrane$ another feature ch orop asts share with mitochondria# The manners b% which ar"e ob9ects enter ce s automatica % create an outer membrane 5actua % a part of the bi" ce +s p asma membrane7 around the incomin" ob9ect# This discussion su""ests a !er% interestin" 4uestion# Endos%mbiosis is a !er% widespread phenomenon# The more we oo&$ the more e2amp es we find throu"hout the &in"doms of ife# )o$ if a mitochondrion is the distant descendent of an independent pro&ar%otic ce $ is it then an or"anism i!in" inside a ar"er ce < Or is it 9ust a part of that ar"er ce < Is it an independent or"anism or not< .efore %ou eap to a conc usion$ thin& a bit# Certain %$ mitochondria are abso ute % dependent upon the ce s in which the% reside# >i&e an% on"0time endos%mbiont$ the% on" a"o "a!e up man% of the basic ife processes needed for independent ife# And the ce s in which the% reside are comp ete % dependent upon their mitochondria$ because the

anaerobic respiration the% cou d do without the mitochondria wou dn+t pro!ide near % enou"h ener"% for the ce +s needs# In fact$ it+s !er% probab e that the e!o ution of bi"$ comp e2 eu&ar%otic ce s wasn+t possib e unti the 'in!ention' of aerobic respiration# .ut there are man% endos%mbiotic re ationships in nature which are 9ust as interdependent# 1or e2amp e$ no termite cou d sur!i!e without the popu ation of endos%mbionts that i!es inside its "uts$ di"estin" its wood% diet for it# And the protists and bacteria that ma&e up that popu ation can+t sur!i!e outside the termite# Comp ete interdependenc%# /ow$ the termite and its passen"ers oo& a ot more i&e independent creatures to us than a ce and its mitochondria# .ut the% are actua % no more independent of each other# )o if we decide that the mitochondrion is 9ust a part of the ce $ then don+t we ha!e to a so decide that the endos%mbionts inside the termite+s "uts are 9ust parts of the termite< If not$ how do we 9ustif% insistin" that there+s a difference< .efore %ou "et too frustrated tr%in" to sort this out$ a ow me to re ie!e %our mind# There is$ in fact$ no answer to this 4uestion# ?ust the reinforcement of a !er% important esson# -espite our human need to sort our wor d into neat$ c ean cate"ories$ the rea uni!erse often doesn+t cooperate$ and this is 9ust such a case# :e want to be ab e to decide 'two separate or"anisms' or 'parts of the same or"anism' in cases i&e this$ but rea it% shows us that there are man% situations which fa somewhere between these two cate"ories# This is a esson we earned when we e2amined the 'a i!e' !s 'not a i!e' issue$ and a"ain when we tried to decide how to functiona % describe species# :e want neat cate"ories8 nature doesn+t cooperate# Cop%ri"ht @ ABBB Co e"e of -uPa"e Center for Independent >earnin" 5CDB7 EFA0AGHI fancherAJcdnet#cod#edu Updated BE )ept ABBF