gaussia is a genus of copepods .

it is a " characteristic genus of the mesopelagial " , occurring at depths of 0 – 3,000 metres ( 0 – 9,843 ft ) .

the genus gaussia contains the following species : gaussia asymmetrica t. k. s. björnberg & campaner , 1988 gaussia gadusae sarkar , 2004 gaussia intermedia defaye , 1998 gaussia melanotica wolfenden , 1905 gaussia princeps ( t. scott , 1894 ) ( type species ) gaussia sewelli saraswathy , 1973

gaussia ( copepod )

bioluminescence in the mesopelagic copepod , gaussia princeps ( t . scott ) - sciencedirect
identification of two catalytic domains in a luciferase secreted by the copepod gaussia princeps .
katja schulz selected
gaussia princeps ( copepod )
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gaussia princeps ( t . scott , 1894 )
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as are most other copepod species , the deep - sea copepod is bioluminescent . . . .
identification of two catalytic domains in a luciferase secreted by the copepod gaussia princeps . - pubmed - ncbi
most copepod species are 1 - 3 mm long , but the deep - sea copepod . . .
gaussia luciferase secreted by the copepod gaussia princeps catalyzes the oxidation of coelenterazine to produce blue light . the primary structure of gaussia luciferase deduced from the cdna sequence shows two repeat sequences of 71 amino acid residues , suggesting the luciferase consists of two structural domains . two domains in gaussia luciferase were expressed independently in escherichia coli cells , purified and characterized . we found that both domains have luminescence activity with coelenterazine , and the catalytic properties including luminescence spectrum , optimal ph , substrate specificity and luminescence stimulation by halogen ions ( cl - , br - and i - ) are identical to intact gaussia luciferase . thus , gaussia luciferase has two catalytic domains for the luminescence reaction .
paul merviel added text to
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cloning and expression of cdna for a luciferase from the marine copepod metridia longa . a novel secreted bioluminescent reporter enzyme
one fascinating shrimp - like copepod that god created is the gaussia princeps . gaussia stores its supply of chemicals in glands located on its tail . when a predator approaches this vulnerable little fellow , it shoots out a stream of glowing blue goo . the predator’s attention is immediately drawn away from the copepod to the movement of the blue stream . then suddenly the goo gives off a bright flash of white light . the flash so startles the predator that it is unable to process what is happening . the copepod is able to slip away and avoid becoming a meal .
jennifer hammock set
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copepoda milne - edwards , 1840
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in the ocean you may find useful molecules , for instance gaussia luciferase ( gluc ) . this is a pretty novel biomiluminescent reporter encoded by a gene isolated from the marine copepod gaussia princeps . this luciferase does not require atp and catalyzes the oxidation of the substrate coelenterazine in a reaction that emits light ( peak at 470 - 480 nm ) like renilla and
rene noam added the english common name
deep sea coceopod
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gaussia princeps ( t . scott , 1894 )
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a . barnes & j . f . case ( 1972 ) .
bioluminescence in the mesopelagic copepod , gaussia princeps ( t . scott )
. journal of experimental marine biology and ecology 8 ( 1 ) : 53–71 . doi : 10 . 1016 / 0022 - 0981 ( 72 ) 90056 - 1 .
copepods are the dominant taxa in zooplankton communities of the ocean worldwide . although bioluminescence of certain copepods has been known for more than a 100 years , there is very limited information about the structure and evolutionary history of copepod luciferase genes . here , we report the cdna sequences of 11 copepod luciferases isolated from the superfamily augaptiloidea in the order calanoida . highly conserved amino acid residues in two similar repeat sequences were confirmed by the multiple alignment of all known copepod luciferases . copepod luciferases were classified into two groups of metridinidae and heterorhabdidae / lucicutiidae families based on phylogenetic analyses , with confirmation of the interrelationships within the calanoida using 18s ribosomal dna sequences . the large diversity in the specific activity of planktonic homogenates and copepod luciferases that we were able to express in mammalian cultured cells illustrates the importance of bioluminescence as a protective function against predators . we also discuss the relationship between the evolution of copepod bioluminescence and the aspects of their ecological characteristics , such as swimming activity and vertical habitat .
gaussia’s incredible ability reminds us of world war 2 navy ships that released depth charges that would be dropped into water close to submerged enemy submarines . a time delay allowed the ship to move away while the charges sank down to get close to the submarine before exploding . similarly , gaussia emits its bioluminescent fluid as it swims swiftly in the opposite direction . just as the bluish fluid is about to disappear , a bright flash ensues that “blinds” the predator and distracts him from pursuing the little copepod .
bioluminescent activity of zooplankton homogenates . luminous activity of homogenized copepods was measured by mixing coelenterazine solution and the supernatant of the lysate . total protein concentration was determined to compare specific luciferase activities among copepod species .
like insects on land , crustaceans fill the oceans of the world . they come in many different shapes and sizes . one very common form is the copepod . many copepods are bioluminescent . they typically release their bioluminescent chemicals into the water to produce a glowing cloud of light . this luminous smoke screen serves to distract or blind an attacker . the copepod can then swim quickly away from danger into the darkness .
wurdinger and colleagues reports to be able to assay in just 5 μl of blood ( a very little drop ) the presence of secreted gaussia luciferase , demonstrating that gluc assay is 1000 fold more sensitive than seap assay .
in this study , we isolated and characterized novel luciferase genes from five species of calanoid copepods and conducted the quantitative analyses of copepod bioluminescence using zooplankton homogenates and recombinant luciferases . by quantitative measurements , we could reveal the presence or absence of luminescence of unverified species with confirmation of verified ones . furthermore , we compared the luminous intensity of homogenates among calanoid superfamilies and those of luciferases among genera in augaptiloidea to infer the defensive function and molecular evolution of copepod bioluminescence .
issued from : r . n . wolfenden in die marinen copepoden der deutschen südpolar - expedition 1901 - 1903 , 1911 . [ pl . xxxiii , fig . 4 ] . as gaussia scotti . female : 4 habitus ( dorsal ) .
issued from : r . n . wolfenden in die marinen copepoden der deutschen südpolar - expedition 1901 - 1903 , 1911 . [ pl . xxxiii , fig . 3 ] . as gaussia scotti . male : 3 , habitus ( dorsal ) . :
we have isolated nine novel copepod luciferases using the homology - based pcr cloning strategy . a blast search using the amino acid sequences of copepod luciferases as the query revealed no highly similar proteins in the database , except for other calanoid luciferases ( gluc , mluc , and mpluc ) . this may be due to the limited number of deposited sequences of copepod proteins . so far , all of copepod luciferase genes , including gluc and mluc reported elsewhere , have been isolated from species in the augaptiloidea superfamily in calanoida . we have tried to isolate luciferase or luciferase - like genes from other superfamilies of calanoida , such as pseudocyclopoidea or centropagoidea using degenerate pcr primers . the result was no amplification or no luciferase - like sequences in the amplified dna product ( data not shown ) . however , we assume that luciferases or luciferase - like genes may exist , not only in augaptiloidea but also in other superfamilies , because the luminous assay of zooplankton lysate from these superfamilies all had a measurable , though very weak , level of activity compared with that of negative control samples ( fig . 2 ) . further screening of the calanoid luciferase or luciferase - like genes by different cloning strategies may reveal the structure and evolution of a direct ancestral copepod luciferase gene .
issued from : r . n . wolfenden in die marinen copepoden der deutschen südpolar - expedition 1901 - 1903 , 1911 . [ pl . xxxiii , figs . 6 - 7 ] . as gaussia scotti . male : 6 , p5 ; 7 , right p5 .
issued from : h . y . soh , s . ohtsuka , h . imabayashi & s . sawamoto in species diversity , 1998 , 3 . [ p . 182 , fig . 7 ] . zoogeographical distribution of gaussia princeps , g . asymmetrica and g . sewelli .
consensus amino acid residues and domain structure were revealed by an alignment of 13 copepod luciferases obtained in this study ( fig . 4a ) . highly conserved amino acid residues , c - x ( 3 ) - c - l - x ( 2 ) - l - x ( 4 ) - c - x ( 8 ) - p - x - r - c , which are present in both domains , would be one of the criteria for the copepod luciferase , although the reason for this cysteine - rich area in the conserved sequence remains obscure . because the similarity in the structure of the two domains was found in all of the copepod luciferases isolated in this study ( fig . 4b ) , we assume this similarity in the structure of the two domains is very characteristic of the luciferases isolated from augaptiloidea species .
luminous activity of zooplankton homogenates varies greatly among the copepods ( fig . 2 ) . for instance , luminous activity of an m . pacifica homogenate was ∼500 - fold higher than that of a h . tanneri homogenate , even though both species belong to the same superfamily . one possible explanation is very different levels of luciferase expression among bioluminescent copepod species . the other is a substantial divergence of specific activity of copepod luciferases . to compare the specific activity of copepod luciferases isolated in this study , 11 luciferases were expressed in hek293 cells as recombinant proteins tagged with hexahistidine at the c - terminus , concentrated from culture medium by a nickel - chelating column , and then tested for their protein expression and activity . specific activity was estimated by dividing the luminous activity of the culture media by signal intensity of polyhistidine - tagged copepod luciferases detected by an anti - polyhistidine antibody ( fig . 6a ) . results showed that there is a large difference in specific activity among copepod luciferases ( fig . 6b ) but only a slight difference in their expression levels ( fig . 6a ) . in several experiments , we consistently observed that mpluc1 showed the highest level of specific activity among tested luciferases , which was approximately 10 6 - fold higher than that of htluc2 . the specific activity of luciferases isolated from copepods in the family metridinidae ( mpluc1 , mpluc2 , moluc1 , moluc2 , paluc1 , and paluc2 ) was notably higher than that of luciferases from the heterorhabdidae family ( htluc1 , htluc2 , hmluc1 , and hmluc2 ) . loluc , isolated from l . ovaliformis , which belongs to the lucicutiidae family , showed intermediate levels of activity . these results are consistent with the luminous activity obtained by homogenate assays of calanoid copepods ( fig . 2 ) . thus , species differences in copepod bioluminescence was reflected in the substantial divergence of specific activity of the expressed luciferases .
bioluminescence in all experiments , except for analyses of the spectra and kinetics , was measured using a minilumat lb 9506 luminometer ( berthold , japan ) . the basic procedure for the luciferase assay was as follows : 5 μl of protein solution containing each of the luciferases was transferred into the luminometer tube and then placed in the luminometer . a 10 - s measurement was initiated immediately after injecting 10 μl of 1 ng / μl coelenterazine in 20 mm tris–hcl ( ph 8 . 0 ) with 50 mm mgcl 2 into the protein solution . luminescent spectra of recombinant copepod luciferases were measured using an ab - 1850c spectrofluorometer ( atto , japan ) ( slit width , 1 mm ; spectral resolution , 0 . 5 nm ) for 1 min after mixing 20 μl of copepod luciferases expressed by cultured cells and 200 ng of coelenterazine in a 0 . 2 - ml pcr tube . to determine thermostability of copepod luciferases , 20 μl of copepod luciferases expressed by cultured cells was incubated in a block heater at 60 or 80 °c for 30 min and then cooled on ice for 5 min . bioluminescence was measured by injecting 10 μl of coelenterazine solution ( 1 ng / μl ) into 5 μl of heat - treated luciferases at room temperature using a minilumat lb 9506 luminometer .
issued from : r . n . wolfenden in die marinen copepoden der deutschen südpolar - expedition 1901 - 1903 , 1911 . [ pl . xxxiii , fig . 5 , 8 ] ] . as gaussia scotti . female : 5 , last thoracic segment and urosome ( lateral , left side ) ; 8 , forehead ( lateral ) .
issued from : r . stephen , 2007 [ data sheets of nio , kochi , india ( on line ) ] . after m . saraswathy , 1973 . as gaussia scotti . female ( from w indian ) : a , last thoracic segment and urosome ( dorsal ) . male : last thoracic segment and urosome ( lateral ) .
( a ) immunoblot of polyhistidine - tagged copepod luciferases expressed by hek293 cells . proteins were concentrated from culture media on a ni - chelating column . the same volume of eluents was loaded on sodium dodecyl sulphate–polyacrylamide gel electrophoresis , blotted , and detected by anti - polyhistidine tag antibody . ( b ) specific activity of secreted copepod luciferases expressed by hek293 cells . the initial light intensity was counted for 10 s and then normalized to the protein expression level obtained by immunoblotting shown in figure 5a . luciferase activities are expressed as percentages of normalized luminescence activity ( light intensity / protein expression level ) , using the value of mpluc1 defined as 100 % . the data represent the mean for triplicate measurements .
in phylogenetic analyses of aligned amino acid sequences of copepod luciferases , 48 substitution models for proteins used in ml estimates were tested by mega5 to find the most appropriate model . the best model showing the lowest bayesian information criterion score was whelan and goldman plus gamma ( gamma distribution ) , therefore , an ml bootstrap consensus tree was generated with the model (
issued from : r . n . wolfenden in die marinen copepoden der deutschen südpolar - expedition 1901 - 1903 , 1911 . [ pl . xxxiii , figs . 9 - 12 ] . as gaussia scotti . female : 9 , p2 ; 10 , p3 ( distal segment of exopod ) ; 11 , p4 ( distal segment of exopod ) ; 12 , p5 .
one strong piece of evidence for the presence of bioluminescence in a particular organism is the molecular identification and functional analysis of luciferase , which is the enzyme directly involved in producing luminescence in an organism . in 2002 , the luciferase gene was cloned from calanoid gaussia princeps ( family metridinidae ) , which is a large ( ∼10 mm ) gray - black–pigmented copepod usually found in the mesopelagic zone ( at the depths from 200 to 1 , 000 m ) ( verhaegen and christopoulos 2002 ) . a novel feature of gaussia luciferase ( gluc ) is its relatively small size ( ∼20 kda ) and secreted luminescence , not only in planktonic expression but also when expressed in mammalian and insect cells , with robust stability and great luminescent activity . gluc catalyzes the oxidation of a luciferin called coelenterazine to produce the light . luciferase genes have also been found in metridia longa ( mluc ) ( markova et al . 2004 ) and metridia pacifica ( mpluc1 and mpluc2 ) ( takenaka et al . 2008 ) , both of which belong to the metridinidae family just as g . princeps does . these gluc - homologues also react with the substrate coelenterazine to produce light . the bioluminescence in these species was confirmed by identification and functional analysis of their luciferases .
issued from : e . suarez - morales in zootaxa , 2007 , 1621 . [ p . 41 , table 2 ] . male : comparison of pore number of right a1 of gaussia princeps from the caribbean ( gpc ) , the indian ocean ( gpl ) from soh & al . , 1998 , and g . sewelli ( gsi ) from saraswathy & bradford , 1980 from the indian ocean . a1 segments grouped in 3 sectors : 1st segment , segments 2 - 5 , and 6 - 12 . nota : the number and general distribution of the pores ( gland openings ) have been found to express important differences between congeners in gaussia . the caribbean specimens showed a pore pattern that differs in number with that described from specimens from the indian ocean and japan ( soh & al . , , 1998 ) on different appendages
santos , e . , yeh , r . , lee , j . , nikhamin , y . , punzalan , b . , punzalan , b . , perle , k . , larson , s . , sadelain , m . , and brentjens , r . ( 2009 ) . sensitive in vivo imaging of t cells using a membrane - bound gaussia princeps luciferase nature medicine , 15 ( 3 ) , 338 - 344 doi : 10 . 1038 / nm . 1930 .
wurdinger , t . , badr , c . and tannous , b . ( 2008 ) . gaussia luciferase blood level as an index of cell growth and proliferation . nature methods , wurdinger t , badr c , pike l , badr c , de klein r , weissleder r , breakefield xo , tannous ba . ( 2008 ) a secreted luciferase for ex vivo monitoring of in vivo processes . nature methods 5 : 171 - 173 doi : 10 . 1038 / nmeth . 1177
( a ) multiple amino acid sequence alignment of 13 copepod luciferases . positions of two putatively similar domains are underlined . outlined letters represent at least 90 % identical ( dark shade ) or chemically similar ( light shade ) amino acid residues . white letters in the black boxes are highly conserved residues in both domains . arrowhead indicates proposed cleavage sites of n - terminal secretion signals of copepod luciferases . ( b ) multiple amino acid sequence alignment of tandem repeat domains of luciferases . residues are darkly shaded and outlined if they are conserved at a level of more than 90 % among all aligned sequences . similar residues are lightly shaded and outlined . the distance between the two domains in each species is indicated as number of amino acids in parentheses at the end of the sequence . those amino acids that are conserved in both domains ( consensus ) , domain 1 only ( consensus _ domain1 ) or domain 2 only ( consensus _ domain2 ) , are indicated in the last three rows .
adult male and female gaussia princeps ( t . scott ) produce bright , extracellular luminescent displays usually coupled with the initiation of rapid swimming . glands of the head , mandibular palps , urosome , furca , and underlying at least 14 body pores contribute to the luminous response which , in free swimming copepods , is characterized by a sudden discharge to the exterior of luminous blue material that remains at peak intensity for 1–3 sec before undergoing an 80 sec decay as 8 or less discrete , low intensity points of light .
( a ) ml phylogeny of copepod luciferases based on an amino acid sequence alignment of domains 1 and 2 with a linker sequence . the tree is midpoint rooted , and numbers on the nodes indicate % bootstrap values from 1 , 000 replicates . luciferases isolated and analyzed in this study are shown in shaded boxes . scale bar indicates evolutionary distance and is in the units of the number of amino acid substitutions per site . ( b ) gene structures of moluc1 and moluc2 . introns and exons are shown as open boxes and lines , respectively .
, a calanoid copepod , measuring about 6 mm from head to telson , making it a giant among hawaiian copepods . copepods are the most abundant and diverse group in the planktonic community . they are crustaceans , related to crabs and lobsters . they are members of the phylum arthropoda , as are other crustaceans , insects , and spiders . there are planktonic , benthic , and parasitic copepods , appearing in freshwater and salt water . their distribution around the world and success in so many niches earns them the title of
the insects of the sea
.
pleuromma princeps t . scott , 1894 b ( p . 42 , descr . m , figs . m ) ; metridia scotti giesbrecht & schmeil , 1898 ( p . 107 ) ; no sewell , 1913 ( p . 354 ) ; gaussia melanotica wolfenden , 1905 a ( pl . 2 , figs . 1 - 5 ) ; gaussia scotti wolfenden , 1905 a ( p . 5 ) ; 1911 ( p . 290 , figs . f , m ) ; lysholm & al . , 1945 ( p . 32 ) ; saraswathy , 1973 a ( p . 191 , figs . f , m ) ; stephen , 2007 [ data sheets of nio , kochi , india ( on line ) ] . female metridia atra esterly , 1906 a ( p . 70 , juv . 5 : f ) ; c . b . wilson , 1950 ( p . 263 , rem . ) ; guérédrat , 1969 a ( p . 362 ) ; no g . princeps : sewell , 1932 ( p . 270 , figs . f , m ) ; 1947 ( p . 173 ) ; ? tanaka & omori , 1967 ( p . 251 )
issued from : e . suarez - morales in zootaxa , 2007 , 1621 . [ p . 42 , table 3 ] . male : comparison of pore number of p1 - p4 of gaussia princeps from the caribbean ( gpc ) , the indian ocean ( gpl ) from soh & al . , 1998 , and g . assymmetrica ( gasa ) from the south - western atlantic ocean ( björnberg & campaner , 1990 ) , and g . intermedia ( ginp ) from the north pacific ( defaye , 1998 ) cx = coxa ; bp = basipodite ; enp = endopodite ; exp = exopodite .
the reactivity of homogenates with coelenterazine indicates the expression of luciferase in these luminous species tested ( fig . 2 ) , as previously reported molecular identification of luciferase genes from single gaussia and two metridia species has indicated ( verhaegen and christopoulos 2002 ; markova et al . 2004 ; takenaka et al . 2008 ) . we amplified full - length luciferase genes from five copepod cdnas with degenerate primers designed based on the nucleotide sequences of the g . princeps , m . longa , and m . pacifica luciferases . at least two forms of luciferase genes , as identified in m . pacifica ( mpluc1 and mpluc2 ) ( takenaka et al . 2008 ) , were also identified from m . okhotensis , p . abdominalis , h . tanneri , and h . major cdnas . the gene names and biochemical features of these proteins are summarized in table 2 . they are relatively small proteins , ranging from 19 . 0 to 23 . 5 kda , and they are efficiently secreted into the culture medium when they are expressed by cultured mammalian cells . the n - terminal 17–22 amino acid sequences of all novel luciferases seem to have the features of a consensus sequence that signals secretion based on the prediction by psort ii ( urltoken ) . moluc1 , moluc2 , paluc1 , and paluc2 retained more than half of their initial activity even after 30 - min incubation at 60 °c but were largely inactivated at 80 °c . thermostability of htluc1 , htluc2 , hmluc1 , and hmluc2 could not be determined correctly because of their low initial activity . the identity of the amino acid sequence of these luciferases was notably high at the internal and c - terminal regions rather than n - terminals ( fig . 4a ) . inouye and sahara ( 2008 ) reported the identification of two catalytic domains in gaussia luciferase , and we confirmed the presence of two short repeat sequences in the primary structures of these novel copepod luciferases . alignment of two repeat sequences consisting of 62–64 amino acid residues ( referred as domains 1 and 2 , respectively ) from 12 luciferases revealed consensus sequence of c - x ( 3 ) - c - l - x ( 2 ) - l - x ( 4 ) - c - x ( 8 ) - p - x - r - c ( x , amino acid residue ) in both domains ( fig . 4b ) . substitution of all cysteine residues with alanine in domains 1 and 2 of mpluc1 resulted in complete loss of activity when it was expressed in escherichia coli ( data not shown ) , suggesting an essential role of cysteine residues in luciferase activity .
issued from : h . y . soh , s . ohtsuka , h . imabayashi & s . sawamoto in species diversity , 1998 , 3 . [ p . 178 , fig . 5 ] . female ( from suruga bay ) : a , p2 ( anterior ) ; b , p3 ( anterior ) ; c , p4 ( anterior ) ; d , p5 ( posterior ) . female ( from the indian ocean ) : e , distal seta on 3rd exopodal segment of right p5 . gaussia sewelli saraswathy , 1973 , female ( from the indian ocean ) : g , distal seta on 3rd exopodal segment of right p5 . arrowheads indicate gland openings .
dvm is a copepod behavior to stay in darker deeper layers during daytime and migrate upward to food - rich surface layers at night for feeding . a most important function of copepod dvm is considered to be avoidance from predation by visual - feeding fishes ( gliwicz 1986 ; neill 1992 ) . species characterized with strong bioluminescence ( metridinidae ) are all medium - sized suspension feeders known to perform strong nocturnal ascent dvm ( haury 1988 ; hays 1995 ) . the dominance of metridinidae in oceans world - wide can be explained by the fast swimming speed and strong dvm intensity ( mauchline 1998 ) . because of its numerical dominance , metridinidae is selectively captured by various mesopelagic lantern fishes ( merrett and roe 1974 ; hopkins and sutton 1998 ) . the bioluminescent behavior of copepods is considered to have a function of avoidance from vision - dependent predators ( herring 1988 ; widder 1992 ) . to distract or blind a predator of mesopelagic lantern fish predator , metridinidae with reinforced luciferases would have a selective advantage for their stronger bioluminescent system ( mpluc , moluc , and paluc ) . we suggest that mpluc from m . pacifica ( fig . 1 ) shows the highest bioluminescence ( fig . 6b ) because the habitat depth of m . pacifica is the epipelagic layer ( 0–200 m ) ( table 1 ) where a large number of predators exist .
issued from : m . saraswathy in handbook to the intern . zoopl . coll . , cochin - 18 , kerala state , india , 1973 . [ p . 191 , fig . 1a , 1b , 2 , 3 ] . as gaussia scotti . female ( from world - wide ) : 2 , p5 . male : 1a , right a1 ( segments 10 to 16 ) ; 1b , glandular structure on segment 12 ) ; 3 , p5 . nota : proximally directed spine on 3rd segment of p5 blunt , short and stumpy . terminal segment of right leg with prominent undulating inner border , and 4 setae ; distance between 1st and 2nd and 2nd and 3rd setae almost equal .
this study was supported by an advanced industrial science and technology research grant ( y . t . , y . s . , n . t . , and m . t . ) , japan foundation for applied enzymology ( y . s . ) , and the nig cooperative research program ( t . g . , 2010 - a66 ) . the authors are greatly indebted to the following individuals : prof . susumu ohtsuka , hiroshima university , for critical discussion for the copepod evolution ; dr steven haddock and ms lynne christianson , monterey bay aquarium research institute , for kindly providing genomic dna of g . princeps ; dr tadashi imanishi and dr akiko ogura noda for introducing mega5 software to calculate ml phylogeny ; and prof . leslie sargent jones , appalachian state university , for valuable discussions .
the origin of the bioluminescence of calanoid copepods must be carefully verified with an assessment of bioluminescence , including nonluminescent primitive species , such as the pseudocyclopoidea , and luminescent oncaea species . nonetheless , we postulate that bioluminescence in calanoid copepods could have originated from an ancient species between the families pseudocyclopoidea and augaptiloidea ( fig . 7 ) . the ancient augaptiloidea may have ventured into the surface water , which was probably caused by unexpected food or oxygen demands ( bradford - grieve 2002 , 2004 ) , and then encountered a number of difficulties for adapting to the pelagic habitat . until myelinated nerve axons evolved , the earlier evolution of bioluminescence might have been a highly efficient strategy for them to avoid surrounding predators . further analysis of copepod bioluminescence in the world ' s oceans could be a clue for understanding the ecology and evolution of marine organisms .
issued from : h . y . soh , s . ohtsuka , h . imabayashi & s . sawamoto in species diversity , 1998 , 3 . [ p . 180 , fig . 6 ] . male : ( from the indian ocean ) : a , a1 ( segments i - viii ) ; b , idem ( segments ix - xix ) ; c , idem ( segments xx - xxviii ) ; d , e ( holotype ) , anterior part of a1 segment xiv ; f , p5 ( posterior ) ; g , medial process on 2nd exopodal segment of left p5 ( anterior ) . gaussia sewelli saraswathy , 1973 ( from the indian ocean ) male : h , p5 ( posterior ; distal two exopodal segments of right p5 torted ) . arrowheads indicate inner process proximally directed on 2nd exopodal segment . p : process . s : spinule .
the phylogeny of 13 calanoid luciferases suggests a different evolution of the metridinidae and heterorhabdidae families ' luciferases ( fig . 5a ) . in the clade containing metridinidae luciferases ( clade 1 ) , two proteins in a single species are likely to have evolved independently . the similarity of primary structures among mpluc1 , moluc1 , and paluc1 ( type - i ) luciferases were greater than those of their counterparts ( mpluc2 , moluc2 , and paluc2 ) ( type - ii ) luciferases ( fig . 4a ) . the mrna coding for moluc1 or moluc2 would be transcribed from two different loci or alleles since studies of genomic sequences revealed the presence of different lengths and positions of the introns in moluc genes ( fig . 5b ) , as seen in mpluc genes ( takenaka et al . 2008 ) . gene duplication events of luciferase probably happened in the ancestral species of the metridia and pleuromamma families , thereafter , paralogous luciferases might have diverged independently . conversely , it would be expected that specialization of heterorhabdidae species preceded the duplication and diversification of luciferase genes since pairs of luciferases ( htluc1 and htluc2 ; hmluc1 and hmluc2 ) are monophyletic ( clades 6 and 7 ) within species . nucleotide identity between coding sequences of hmluc1 and hmluc2 is 91 . 8 % , whereas it is 62 . 6 % between hmluc1 and htluc1 , which were isolated from species of the same heterorhabdidae family . extensive genome survey of two lineages of metridinidae and heterorhabdidae families may shed light on the multigene structure of luciferase genes and how copepod luciferases have evolved . from the present study , we could not interpret any evolutionary novelty derived from gene duplication of luciferases in calanoid copepods . because phenotypic observation on live copepod bioluminescence is difficult without special video equipment and relevant technical skills , we are trying to evaluate any possible evolutionary novelty of the gene duplication by revealing different biochemical characteristics between type - i and type - ii luciferases identified in this study .
issued from : j . j . childress in mar . biol . , 1977 , 39 . [ p . 23 , fig . 3 ] . gaussia princeps adult males ( in experimental proceedings ) . interpolation of data to predict oxygen - consumption in the field . solid line shows oxygen - consumption rate based on temperature and pressure considerations only ; dotted line takes into account limitations of oxygen uptake by low oxygen and indicates maximum oxygen - consumption possible in the region of this limitation ; hatched area represents that part of the metabolic needs which must be accounted for by anaerobic metabolism . the two scales on the right show the temperature and oxygen characteristics of a typical area ( san clemente basin , southern california ) where the animals are found . crosses indicate primary , non - interpolated data , while dots are interpolated points . nota : gaussia princeps is found only below 400 m in the daytime and is present down to the greatest depth sampled ( 900 m ) . at night an appreciate fraction of the population migrates to depths shallower than 400 m . even at night , however , many remain below 400 m . the individuals clearly do not need to migrate above this depth on a daily basis . this species ' consumption was measured at 3 . 5 , 7 and 10°c and 1 , 14 , 28 , 61 , 121 and 181 atm of hydrostatic pressure ( 1 atm corresponds about 10 m of depth ) . hydrostatic pressure is shown to have significant effects on the oxygen - consumption rate at pressures as low as 28 atm . at all temperature and pressure combinations , the species displays a very low metabolic rate compared to shallow - living copepods . the critical oxygen partial pressure for this species is shown to be about 10 to 1 , mm hg o2 at 10 , 7 and 5 . 5 °c . this shows a higher oxygen - consumption rate at the nighttime depths and a much lower , partially anaerobic metabolism at the daytime depths .
the full coding sequences of the copepod luciferases were amplified by pyrobest dna polymerase ( takara bio ) from the template plasmids that harbor the full - length cdnas and subcloned into pcdna3 . 1 / v5 - his - topo ( invitrogen ) to express luciferase proteins tagged with 6× histidines at their c - terminus in mammalian cultured cells . all expression vectors were purified using an endofree plasmid maxi kit ( qiagen , japan ) . hek293 cells ( health science research resources bank , japan ) were maintained in megacell mem ( sigma - aldrich , japan ) supplemented with 5 % fetal bovine serum at 37 °c in a humidified atmosphere of 5 % co 2 . for transfections , 5 × 10 5 cells per well were seeded in six - well culture plates , and fugene hd ( roche applied science ) was used according to the manufacturer ' s instructions . the culture medium was incubated for 72 h after transfection and stored at −30 °c until immunoblotting and luciferase assay .
concerning the bioluminescent copepods other than calanoida , oncaea belonging to poecilostomatoida is known to have bioluminescence ( herring et al . 1993 ) . since many of the bioluminescent copepods are predatory and live moderately deep , it is unique that the tiny poecilostomatoid copepod oncaea is one example that is relatively shallow water habitat and has pseudoplanktonic behavior ( böttger - schnack and schnack 2005 ) . the most notable feature of oncaea bioluminescence is their nonsecretory and repetitive fast flashes . the reason for this different characteristic of oncaea bioluminescence compared with those of other bioluminescent copepods might lie in their smaller size and reduced motility . throwing off bioluminescence as a decoy against potential predators is not an effective protection for oncaea , which cannot escape rapidly from surrounding luminescent materials ejected from themselves , unlike calanoid copepods ( herring et al . 1993 ) . furthermore , the bioluminescence emission spectra of oncaea ( ∼470 nm ) are slightly different from those of other calanoid copepods ( 480–500 nm ) , possibly due to the structural difference of their luciferase and / or luciferin .
copepods are the most numerous taxa of zooplankton fauna in the ocean worldwide . their bioluminescent abilities and characteristics have been investigated for over a 100 years ( giesbrecht 1895 ; clarke et al . 1962 ; haddock et al . 2010 ) . luminescence of calanoid copepods is produced by glandular cells in the luminous glands , which are largely located on the caudal rami and legs ( fig . 1 ) , although their number and location vary considerably among families . most of the luminous species reported to date belong to the superfamily augaptiloidea , which contains the families arietellidae , augaptilidae , heterorhabdidae , lucicutiidae , metridinidae , and nullosetigeridae ( herring 1988 ) . within these families , the luminescent behavior of metridinidae and most heterorhabdidae , lucicutiidae , and augaptilidae are well known ( herring 1988 ) . based on a previous cladistic analysis of morphological features , the augaptiloidea is believed to have diverged very early in the evolution of calanoid copepods ( bradford - grieve et al . 2010 ) . except for the augaptiloidea superfamily , a single species in megacalanoidea ( megacalanus princeps ) was reported as luminous ( herring 1988 ) with very limited evidence , and it remains unverified . it is also unclear whether species in other superfamilies , such as centropagoidea , clausocalanoidea , and eucalanoidea , are luminous . this indicates that among the ten calanoid superfamilies , bioluminescent copepods are found almost exclusively in the superfamily , augaptiloidea . concerning bioluminescence behavior other than calanoid copepods , that of tiny pseudoplanktonic poecilostomatoid oncaea spp . is only notable ( herring et al . 1993 ) . one reason for the limited information on copepod bioluminescence is that correct copepod identification is very difficult . in addition to the species identification challenges , zooplankton sampling , especially in deep water , requires special equipment , including large - sized plankton nets , to collect sufficient numbers of specimens . furthermore , macro - and microscopic detection of weak and flash - type bioluminescence from tiny copepods is also difficult , especially if the specimen was damaged during the collection or manipulation by an investigator . therefore , a device , such as a luminometer , to detect weak bioluminescence of zooplankton is needed .
issued from : a . t . barnes & j . f . case in j . exp . mar . biol . ecol . , 1972 , 8 . [ p . 59 , fig . 3 ] . distribution of fluorescent glands , luminescent glands , and luminous pores of adult male and female gaussia princeps . a : female ; b : male ; c : lateral view of 5th thoracic segment and abdomen of female ; d : lateral xiew of female . f : fluorecent gland ; l . g . : luminescent gland ; l . p . : luminous pore . ( modified after wolfenden , 1911 , and sewell , 1932 ) . nota : adult male and female produce bright , extracellular luminescent displays usually coupled with the initiation of rapid swimming . glands of the head , mandibular palps , urosome , caudal rami , and underlying at least 14 body pores contribute to the luminous response which , in free swimming copepods , is characterized by a sudden discharge to the exterior of luminous blue material that remains at peak intensity for 1 - 3 sec before undergoing and 80 sec decay as 8 or less discrete , low intensity points of light .
copepod luciferases , isolated in this study and previous reports , are all derived from the augaptiloidea superfamily in calanoida . there are few to no reports of bioluminescence in other calanoid superfamilies . the reason for the difference in the luminous ability between the augaptiloidea and other superfamilies could be explained by their ecology ( e . g . , motility , habitat , and body size ) . although precise evolutionary positions of the pseudocyclopoidea and epacteriscoidea are not determined yet , these superfamilies are believed to be the most primitive calanoida ( fig . 7 ) ( bradford - grieve et al . 2010 ) . comparing with other eight superfamilies illustrated in figure 7 , they are highly specialized because of their adaptation to the epibenthic habitats . they are found only at shallow to shelf depth ( ohtsuka et al . 1999 ) or in submarine caves ( ohtsuka et al . 2002 ) , in contrast to the augaptiloidea , which has a greater diversity of distribution within the pelagic zones ( yamaguchi et al . 2004 ) . these observations imply little change , if any , of habitat in the evolutionary history of the pseudocyclopoidea and epacteriscoidea ( bradford - grieve 2004 ) . living on - bottom– or in - bottom–dwelling habitat would be a very effective means to find nutrients and hide from predators . there is no report , so far , describing the bioluminescence of these primitive copepods , probably because of the very limited research on these minor species in calanoida so far . however , considering their stable benthic habitat and very small body size ( below 1 mm ) ( ohtsuka et al . 1999 ) , evolving bioluminescence as a defense against the predators seems to be unnecessary for them .
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davis , 1949 ( p . 50 , figs . f ) ; c . b . wilson , 1950 ( p . 235 , figs . f , m ) ; brodsky , 1950 ( 1967 ) ( part . , p . 312 , figs . np : f , m , non figs . f , m : ind . oc , rem . ) ; vervoort , 1965 ( p . 103 , rem . ) ; owre & foyo , 1967 ( p . 74 , figs . f , m ) ; barnes & case , 1972 ( p . 53 , bioluminescence ) ; gardner & szabo , 1982 ( p . 348 , figs . f , m ) ; björnberg & campaner , 1988 ( p . 351 , 354 , fig . f , rem . ) ; hulsemann , 1988 ( p . 188 , rem . ) ; baessa - de - aguiar , 1989 ( 1992 ) ( p . 117 , figs . f , m ) ; bowlby & case , 1991 ( p . 440 , bioluminescence ) ; soh & al . , 1998 ( p . 171 , redescr . f , m , figs . f , m , juv . , fig . 7 : carte , rem . ) ; bradford - grieve & al . , 1999 ( p . 884 , 948 , figs . f , m ) ; bradford - grieve , 1999 b ( p . 111 , figs . f , m , rem . , figs . 177 , 192 ) ; vives & shmeleva , 2007 ( p . 3 , figs . f , m , rem . ) ; vives & shmeleva , 2007 ( p . 357 , figs . f , m , rem . ) ; suarez - morales , 2007 ( p . 34 , figs . m , tabl . 2 , 3 , 4 , rem . ) ; mercier & al . , 2013 ( p . 760 , neural ultrastructure )
issued from : j . m . bradford - grieve in the marine fauna of new zealand : pelagic calanoid copepoda . national institute of water and atmospheric research ( niwa ) . niwa biodiversity memoir , 111 , 1999 . [ p . 111 , fig . 74 ] . female ( from 39°49 . 9 ' s , 178°26 ' e ) : a , habitus ( dorsal ) ; b , p5 . male : c , habitus ( right lateral side ) ; d , p5 ( l = left leg ; r = right leg ) . female characteristics : - spinous prolongations of posterolateral corners of last thoracic segment divergent . - proximal part of genital segment asymmetrically inflated , with a prominent curved process usually on right side . - blunt processes on caudal rami are prominent . - penultimate segment of p5 as broad as long . male characteristics : - processes on caudal rami as in female ; - proximally directed spine on segment 3 of left p5 blunt , short and stumpy ; terminal segment of right leg with a prominent undulating border , and 4 setae ; the distance between setae 1 and 2 and 2 and 3 almost equal . remarks :
in the southwest pacific males right p5 terminal segment with the distance between spines 1 and 2 ( numbering from proximal part of segment ) is almost twice that of the distance between spines 2 and 3 ; spines 3 and 4 much shorter than spines 1 and 2 .
issued from : h . y . soh , s . ohtsuka , h . imabayashi & s . sawamoto in species diversity , 1998 , 3 . [ p . 173 , fig . 1 ] . female ( from suruga bay ) : a , habitus ( dorsal ) ; b , idem ( lateral left side ) ; c , forehead ( lateral ) ; d , genital compound somite ( dorsal ) ; e , idem ( lateral ) ; f , idem ( ventral . c : copulatory pore ; cd : copulatory duct ; g , gonopore .
issued from : h . y . soh , s . ohtsuka , h . imabayashi & s . sawamoto in species diversity , 1998 , 3 . [ p . 143 , fig . 2 ] . female : scanning electon microscope micrographs of genital compound somite ; aventral ( copulatiory pores damaged ) ; b , right gonopore ; c , left gonopore . scale bars = 0 . 3 mm ( a ) ; 0 . 05 mm ( b , c ) .
issued from : h . y . soh , s . ohtsuka , h . imabayashi & s . sawamoto in species diversity , 1998 , 3 . [ p . 175 , fig . 3 ] . female : a , left a1 ( segments i - xiv ) ; b , idem ( segments xv - xx ) ; c , idem ( segments xxi - xxviii ) ; d , idem ( segments xiv - xv ) ; e , idem ( segments xxvii - xxviii ) ; f , a2 ; g , 2nd endopodal segment of a2 ; h , labrum and paragnaths ; i , md . arrowheads indicate gland openings .
issued from : h . y . soh , s . ohtsuka , h . imabayashi & s . sawamoto in species diversity , 1998 , 3 . [ p . 177 , fig . 4 ] . female : a , mx1 ; b , mx2 ; c , mxp ; d , p1 ( anterior ) . arrowheads indicate gland openings .
issued from : t . k . s . björnberg & a . f . campaner in hydrobiomogia , 1988 , 167 / 168 . [ p . 352 , figs . d , e , f ] . female ( from chile ) : d , forehead ( dorsal ) ; e , idem ( lateral ) ; f , metasomal corners and genital segment ( dorsal ) .
issued from : t . k . s . björnberg & a . f . campaner in hydrobiomogia , 1988 , 167 / 168 . [ p . 353 , figs . b - f ] . female : b , genital segment with attached couplers ( lateral right side ) ; d , idem ( ventral ; e , right coupler with spermatophore after removal of filling material ) ; c , genital segment ( lateral right side ; another specimen ) .
issued from : e . suarez - morales in zootaxa , 2007 , 1621 . [ p . 35 , fig . 1 ] . male ( from w caribbean ) : a - b , habitus ( lateral and dorsal , respectively ) ; c , forehead lateral ; ( showing frontal process and depression ) ; d , 5th pedigerous and 1st urosomites ( dorsal ) ; e , anal somite and caudal rami ( dorsal ) ; f , same ( lateral ) . scale bars : 3 mm ( a , b ) ; 0 . 7 mm ( c - f )
issued from : e . suarez - morales in zootaxa , 2007 , 1621 . [ p . 36 , fig . 2 ] . male : a , left a1 ( segments 1 - 10 ) ; b , same ( segments 11 - 16 ) ; c , same ( segments 17 - 23 ) ; d , right a1 ( segments 1 - 10 ) ; e , same ( segments 11 - 20 ) ; f , same ( detail of modified seta and aesthetascs ) ; g , detail of segment 17 and distal acute process ( position of integumental pores indicated by solid circles . scale bars : 0 . 7 mm ( a - e ) ; 0 . 05 mm ( f ) ; 0 . 35 mm ( g ) .
issued from : e . suarez - morales in zootaxa , 2007 , 1621 . [ p . 37 , fig . 3 ] . male : a , a2 ; b , md ( mandibular palp ) ; c , md ( mandibular blade showing detail of teeth and blade ornamentation ) ; d , mx2 ; e , mxp ; e , p1 ; g , rostal area ( ventral ) . position of integumental pores indicated by solid circles . scale bars : 0 . 7 mm ( a , b - g ) ; 0 . 2 mm ( c ) .
issued from : e . suarez - morales in zootaxa , 2007 , 1621 . [ p . 38 , fig . 4 ] . male : a , p2 ; b , detail of hook - like and spiniform processes on 1st endopodal segment of p2 ; c , p3 ; d , p4 ; e , p5 ( anterior ) ; f , basipod and exopod of left p5 ( lateral view ) ; g , detail of processes of 2nd exopodal segment of left p5 ; h , detail of distal exopodal segment of right p5 ( showing ornamentation ) . position of integumental pores indicated by solid sircles . scale bars : 0 . 7 mm ( a , c - e , h ) ; 0 . 3 mm ( b , f , g ) .
issued from : e . suarez - morales in zootaxa , 2007 , 1621 . [ p . 42 , table 3 ] . male : comparison of pore number of p5 of from the caribbean ( gpc ) , the indian ocean ( gpl ) from soh & al . , 1998 , and g . sewelli ( gsi ) from saraswathy & bradford , 1980 from the indian ocean . cx = coxa ; bp = basipodite ; enp = endopodite ; exp = exopodite ; r = right ; l = left .
issued from : c . b . wilson in bull . u . s . natn mus . , 100 , 14 ( 4 ) . [ pl . 25 , figs . 377 - 378 ] . as metridia atra . female ( from 13°37 ' 40 ' ' n , 120°39 ' e ) : 377 , endopod of p2 ; 378 , p5 .
issued from : c . b . wilson in bull . u . s . natn mus . , 100 , 14 ( 4 ) . [ pl . 11 , figs . 117 - 119 ] . female ( from pacific ) : 117 - 118 , habitus ( dorsal and lateral ) . male : 119 , habitus ( dorsal ) . nota : the color is very dark , almost black , with the ventral surface of the genital segment a reddish brown .
issued from : m . saraswathy in handbook to the intern . zoopl . coll . , cochin - 18 , kerala state , india , 1973 . [ p . 194 ] . comparison between g . scotti ( = g . princeps ) and g . sewelli .
issued from : h . b . owre & m . foyo in fauna caribaea , 1 , crustacea , 1 : copepoda . copepods of the florida current . 1967 . [ p . 73 , fig . 479 ] . female : 479 , p2 .
issued from : h . b . owre & m . foyo in fauna caribaea , 1 , crustacea , 1 : copepoda . copepods of the florida current . 1967 . [ p . 17 , fig . 47 ] . female : 47 , p5 .
issued from : h . b . owre & m . foyo in fauna caribaea , 1 , crustacea , 1 : copepoda . copepods of the florida current . 1967 . [ p . 20 , fig . 74 ] . male : 74 , p5 .
sewell , 1948 ( part . , p . 330 , 503 , 516 , 521 , 530 , 548 , 558 , 561 ) ; grice , 1963 a ( p . 496 ) ; fleminger , 1967 a ( tabl . 1 ) ; grice & hulsemann , 1967 ( p . 17 ) ; 1968 ( tab . 2 ) ; morris , 1970 ( p . 2301 ) ; lee & al . , 1971 ( p . 1151 , 1152 ) ; gueredrat & friess , 1971 ( p . 187 , fig . 1 ) ; roe , 1972 ( p . 277 , tabl . 1 , tabl . 2 ) ; björnberg , 1973 ( p . 338 , 387 ) ; lee & barnes , 1975 ( p . 265 , lipids ) ; childress , 1975 ( p . 787 , respiratory rate ) ; 1977 ( p . 19 , pressure , temperature v . s . oxygen consumption ) ; vives , 1982 ( p . 293 ) ; petipa & borichenko , 1985 ( tab . 2 ) ; madhupratap & haridas , 1986 ( p . 105 , tab . 1 ) ; lozano soldevilla & al . , 1988 ( p . 59 ) ; heinrich , 1990 ( p . 18 ) ; bowlby & case , 1991 ( p . 329 ) ; 1991 a ( p . 440 ) ; suarez - morales & gasca , 1998 a ( p . 110 ) ; razouls & al . , 2000 ( p . 343 , appendix ) ; haury & al . , 2000 ( p . 69 , table 1 ) ; lenz & al . , 2000 ( p . 338 ) ; fields & al . , 2002 ( p . 173 ) ; fernandes , 2008 ( p . 465 , tabl . 2 ) ; galbraith , 2009 ( pers . comm . ) ; inouye & sahara , 2008 ( p . 96 , luciferase ) ; takenaka & al . , 2012 ( p . 1669 , fig . 3 , table 1 , bioluminescence ) ; teuber & al . , 2013 ( p . 1 , table 2 , fig . 5 , respiration rates ) ; in calcofi regional list ( mdo , nov . 2013 ; m . ohman , comm . pers . )
issued from : m . saraswathy in handbook to the intern . zoopl . coll . , cochin - 18 , kerala state , india , 1973 . [ p . 193 ] . localities from g . scotti ( = g . princeps ) ( black circle ) and g . sewelli ( white circle ) obtained .
sub - antarct . ( se pacif . ) ( in björnberg & campaner , 1988 ) , de trindade is . , off st . paul is . , off gabon , g . of guinea , cape verde is . , off morocco - mauritania , canary is . , caribbean sea , belize , florida , w sargasso sea , s . cape hatteras , indian , bay of bengal , philippines , japan , s . hokkaido , off n new caledonia , new zealand ( e north island ) , pacif . ( equatorial ) , ne pacif . ( queen charlotte is . ) , hawaii , off n hawaii , california ( off monterey bay , san pedro , santa catalina , san clemente , velero and tanner basins ) , ne easter is . , sw galapagos , pacif . ( se tropical ) , off juan fernandez is . , peru , chile ( n & s , sub - antarct . )
( 10 ) f : 10 , 5 - 10 ; ( 16 ) f : 10 , 75 - 10 , 5 ; m : 10 , 4 - 9 , 4 ; ( 59 ) f : 12 - 9 ; m : 12 - 9 ; ( 94 ) f : 10 . 5 - 11 . 4 [ indian ] ; 10 . 0 - 10 . 7 [ atlant . ] ; 10 . 0 - 11 . 6 [ pacif . ] ; m : 9 . 6 - 10 . 6 [ indian ] ; 9 . 1 - 10 . 4 [ atlant . ] ; 10 . 1 - 11 . 1 [ pacif . ] ; ( 199 ) m : 10 , 08 - 9 , 92 ; ( 770 ) f : 10 , 47 [ indien ] ; 10 , 9 [ japon : suruga bay ] ; m : 9 , 54 [ indien ] ; ( 909 ) f : 10 , 45 - 11 , 06 ; m : 9 , 39 - 9 , 85 [ n - z ] ; ( 986 ) m : 10 , 1 ; { f : 9 , 00 - 12 , 00 ; m : 9 , 00 - 12 , 00 } the mean female size is 10 . 589 mm ( n = 19 ; sd = 0 . 6712 ) , and the mean male size is 10 . 058 mm ( n = 22 ; sd = 0 . 7233 ) . the size ratio ( male : female ) is about 0 . 94 ( n = 9 ; sd = 0 . 0347 ) .
razouls c . , de bovée f . , kouwenberg j . et desreumaux n . , 2005 - 2018 . - diversity and geographic distribution of marine planktonic copepods . sorbonne université , cnrs . available at urltoken [ accessed july 09 , 2018 ]


gay trip was a racehorse noted for winning the 1970 grand national .

gay trip was a small bay gelding owned by tony chambers and trained by fred rimell .

formerly a flat racing horse , gay trip was switched to national hunt racing as a five-year-old and won the mackeson gold cup in 1969 .

in the 1970 grand national he was ridden by pat taaffe after his regular jockey terry biddlecombe was ruled out of the race by injury .

gay trip carried top weight of 11 stones five pounds despite previously never having won a race longer than 2 ½ miles and started at odds of 15/1 in a field of twenty-eight runners .

he took the lead at the second last fence and drew clear to win by twenty lengths .

since 1970 only red rum has won the grand national carrying top weight .

it was fred rimell 's third national winner having previously won with e.s.b. and nicolaus silver .

in 1972 he finished second in the grand national , ridden by biddlecombe .

gay trip

naples is a fascinating city and it makes for an interesting trip during your visit to italy . interested in learning more ? here are five experiences to enhance your visit .
we ' ve brought together our favorite and most diverse spots across mexico to create a 10 - day gay holiday trip from the caribbean coast , through central mexico , with a celebratory finish on the pacific coast . a holiday trip wouldn ' t be complete without ringing in 2019 with a bang there ' s no better place than puerto vallarta , one of the favorite nye spots for guys from all over the world .
our 11 - day trip through israel & jordan ticks off an amazing number of bucket list items , from floating weightlessly in the dead sea , to riding camelback through the desert , to gazing up at the famous buildings of petra carved into the cliffs . these 11 days in israel & jordan bring us into the middle of a region of fascinating contradictions . as gay travelers , this trip gives us the ability to take in the middle east while enjoying the comfort of gay welcoming community .
join italy gay travels on a cycle trip like no other ! the gay cycling tour takes you from the north to the south of this region , allowing you to discover many of the magical places in puglia . puglia is fast becoming the gay destination of south italy and that is why we are sure that the gay cycling tour is perfect for you ! the area is rich in culture , food , awe inspiring landscapes , white sandy beaches and some gay nightlife for good measure !
our 15 - day adventure starts and finishes in the metropolis of bangkok , the capital city of thailand and home to asia ' s best food , nightlife , and shopping . it includes everything that is experienced in our 12 - day trip , with a few extra days of exploring new destinations too ! the majority of this incredible trip will be spent on the islands and beaches of southern thailand .
owned by mr . a . chambers , gay trip was trained by fred rimmell . rimmell had been an extremely successful jockey and had won the national hunt champion jockey title four times before two falls , resulting in a broken neck , cut short his riding career .
our 8 - day peru trip showcases the jaw - dropping andean mountain range in all it ' s glory ! this trip starts in the bustling capital city of lima and finishes 8 days later in the ancient inca city of cusco . from historic mountaintop fortresses , to lush river valleys surrounded by impressive natural wonders , these 8 days in peru show off the side of peru that everyone dreams of experiencing for themselves .
a gay australia cultural tour down under , including sydney’s gay and lesbian mardi gras ! what is the mystery surrounding the great land called “oz ? ” and no , it’s not the ruby slippers… explore gay “down under” , including visiting the sydney opera house to dining with an aboriginal family . from sophisticated afternoon tea and 4 - wheeling in the outback , to sunset dinner aboard a private yacht and a fabulous gay mardi gras , this is the trip of a lifetime !
pat taaffe was the jockey chosen to ride gay trip for the race , he had already enjoyed great success with the legendary arkle winning three gold cups and a fourth on another horse , fort leney and was no stranger to the national having won in 1955 on quare times .
our 12 - day greece trip promise to deliver the best of athens and the greek islands with stops in dramatic santorini , cosmopolitan mykonos , and peaceful milos . our laid - back trip leaves plenty of room for spontaneity and adventure . there is no amount of photos , video , or stories that can prepare a person for the incredibly captivating country that is greece . it ' s a place that few people visit just once .
join italy gay travels on a not - to - be - missed tour of the worldwide gay opera capitals : milan , verona and venice . our gay opera tour will start from the fashion capital of italy , milan of course , where we will discover the amazing history of the city , enjoy its exuberant gay nightlife , and spend a night at la scala for an unforgettable opera night . a day trip exploring como and its lake is without a doubt a highlight on this tour .
a gay sicily tour like no other ! join italy gay travels on an amazing gay sicily tour . sicily is the biggest island in the mediterranean sea , rich in culture and home to important archaeological sites . yes , in this case bigger is better , as sicily packs in all the punches what all gay tour should offer . this exclusive gay sicily tour will focus on the eastern part of this amazing region , fast becoming a gay hotspot .
one of the most successful trainers in grand national history he is the only one to have had four wins with four different horses . one of only two greys to ever win the race , nicolaus silver , was trained by him , as were national winners e . s . b . , rag trade and of course gay trip .
hop on our gay tour of thailand and discover the exquisite land of smiles in style ! thailand conjures up images of spicy delicacies , majestic elephants , golden buddhas , vibrant nightlife and exotic hideaways - and we experience it all on this trip for gay men , their friends and family . learn about , and spend time with , the majestic asian elephant . live it up on phuket , thailand’s famous beach island .
on race day , gay trip was carrying the top weight of 11st 5lbs , and having never won beyond two and a half miles his chances were not that good . taaffe did a great job steering him to a 20 length victory at odds of 15 / 1 and in doing so provided the perfect end to his own racing career .
this gay group tour features our favourite part of spain — barcelona and south through madrid and andalucia . though distances travelled are not huge , this is a very diverse trip , from glitzy barcelona and its outstanding nearby beaches , to stately madrid with its marvelous galleries and shopping , to the fascinating moorish sites of cordoba , granada and sevilla .
as gay travelers , this trip gives us the ability to take in the middle east while enjoying the comfort of a relatively gay welcoming community . we ' ll immerse ourselves into the thriving lgbt scene , hike around the clifftop fortress of masada , cover ourselves in the nutrient - rich mud of the dead sea , explore the winding alleys of one of the most historically significant cities in the world among many more unforgettable experiences .
from colonial architecture to cuban cigars , this gay group trip gets up close and personal with the caribbean island’s distinct latin culture . it also happens to coincide with new year’s eve , so pack your fun pants and prepare to celebrate . throw in some friendly locals , a few old cars , and even a splash of salsa , and 2019 is off to a stellar start ! ring in 2019 on this gay people - to - people tour to cuba !
from colonial architecture to cuban cigars , this gay group trip gets up close and personal with the caribbean island’s distinct latin culture . it also happens to coincide with new year’s eve , so pack your fun pants and prepare to celebrate . throw in some friendly locals , a few old cars , and even a splash of salsa , and 2020 is off to a stellar start ! ring in 2020 on this gay people - to - people tour to cuba !
whether you ' re planning a road trip , currently on the road , or simply love the genre , here are some of our favorite road trip movies , all of which feature lgbt characters , themes , or flavors . taking us from the deserts of the american southwest to the flowery meadows of the south of france , and across the vast australian outback and to the secluded beaches of mexico , these movies show us some of the many ways in which traveling the open road can change us - and help us to see the world and ourselves in a different light .
this is a salubrious addition to our european walking series in region that lends itself beautifully to our leisurely style of day hiking from a hotel base . the trip also features walking tours in no fewer than six unesco world heritage towns , as well as the stunning unesco protected plitvice lakes region .
enjoy an amazing 7 - day gay trip experience in barcelona from july 28th to august 4th , getting to meet your hosts in exciting activities while you discover the wonders of of spain ! join ricky roman , josh moore , ian greene , jay alexander and other hot spanish influencers on their vacations . it ' s hot in barcelona during august , but our influencers are even hotter !
enjoy an amazing 7 - day gay trip experience in barcelona from august 4th to august 11th , getting to meet your hosts in exciting activities while you discover the wonders of of spain ! join alex mecum , jack hunter , kyle goffney & anthony forte and other hot spanish influencers on their vacations . it ' s hot in barcelona during august , but our influencers are even hotter !
enjoy an amazing 7 - day gay trip experience in madrid from september 29th to october 6th , getting to meet your hosts in exciting activities while you discover the wonders of the capital city of spain ! join jake ashford , beaux banks , salvatore martinez and brogan fletcher on their vacations . it’s the start of autumn – a great month to explore all that madrid has to offer .
join italy gay travels on an amazing gay bike tour in the southernmost tip of puglia , known as salento . fast becoming the gay destination of south italy , this area is rich in culture , food , awe inspiring landscapes , white sandy beaches and some gay nightlife for good measure ! our tour is set at an easy level that will suit most cyclists .
enjoy an amazing 7 - day gay trip experience in barcelona from september 1st to september 8th , getting to meet your hosts in exciting activities while you discover the wonders of of spain ! join ryan rose , niko wirachman , ryan carter , seth knight and other hot spanish influencers on their vacations . it ' s still hot in barcelona during september , but our influencers are even hotter !
the forces of nature that shaped patagonia remain on display to this day . this trip begins with a visit to perito moreno and upsala , two of the region’s most iconic glaciers . another day features a nature walk , a visit to estancia cristina ranch , and a boat ride on argentino lake .
an elegant gay group journey to selected historic homes & gardens in ireland , featuring accommodations in ireland’s best castle & manor house hotels . we will luxuriate at such famous hotels as waterford castle ; ashford castle and adare manor . dinner has been planned at the hotels every evening to take advantage of the excellent kitchens of these establishments . the sightseeing theme of our trip is historic houses and gardens .
the north atlantic’s azores , like hawaii , is a remote volcanic archipelago . this active gay group trip explores the crater of sete cidades ( one of the 7 wonders of portugal ) , bathes you in natural thermal pools , and includes a world - class whale watching excursion . you ' ll have the opportunity to explore four iconic islands including săo miguel , faial , pico and săo jorge .
peru is ready to blow your mind ! with breathtaking vistas in every direction , incredibly unique local culture , and an unlimited number of exciting activities along the way , peru is a perfectly enriching destination . our 12 - day gay trip through peru mixes our laid - back travel concept with peru ' s most unique and memorable sites to form an adventure that you ' ll be talking about forever .
join italy gay travels on an amazing tour of the italian gay capitals : rome , florence and turin . yes , take in all the key sights in one tour ! rome , florence and turin have one thing in common : they have all been capitals of italy since 1861 , the year italy was unified . the gay capitals of italy tour allows you to discover the lifestyles of the most fascinating gay hotspots in italy . you too can now join the unification on the gay capitals exclusive tour !
what ? a greek isles gay tour that ' s not a cruise ? ? ? on this very special trip , we travel by scheduled ferry through the beautiful cycladic island group and spend our nights on each unique island , with a perfect balance of organized activities and free time to soak it all up . greece is a country of beautiful contradictions , a constant journey in time from the present to the past and back again .
an exclusive and unforgettable gay naples , capri and the amalfi coast tour ! join italy gay travels on an amazing gay group tour of the coast around naples . this unforgettable holiday will commence with a gay naples tour , to discover this charming seaside city . from there we will make your way to the amalfi coast and to the island of capri , world famous for its beaches and the stunning faraglioni .
thailand has that exotic , mysterious , exciting appeal to it . our 12 - day adventure starts and finishes in the metropolis of bangkok , the capital city of thailand and home to asia ' s best food , nightlife , and shopping . the majority of this incredible trip will be spent on the islands and beaches of southern thailand .
gay games is an event for inclusion and respect of diversity . from 4 august to 12 august , 2018 , paris hosts the 10th edition of the gay games , the world’s largest sporting , cultural and festive event open to all ! held every four years for more than 32 years , gay games , a hymn to love , advocates a society for all diversities . it ' s the spirit of the gay games ! paris gay games 2018 rich program of festivities completes sporting and cultural programs . will you join us ?
join us for a gay tour that blends together the sophistication , heritage and promise of cuba’s capital with three days of exploring smaller towns that are rarely visited by foreigners . on this trip we will have a special emphasis on the arts of cuba , and the role of the lgbt community . we invite you to join us for a people - to - people exchange with leaders of social change in cuba and some of the country’s most creative artists .
nepal is a hiker’s paradise laden with ancient kingdoms , majestic mountains , and monasteries galore . our gay trip begins with two days exploring the busy streets of kathmandu , before we fly over the haunting himalayas en route to the everest region . our own hike reaches its own impressive heights with an overnight stay at everest base camp ( 5 , 364m ) , where we can bask in the majesty of the mother of the universe , that epic mountain herself .
our the best of vietnam & cambodia luxury small gay group trip is distinguished by the top hotels selected in three countries and ample time to leisurely visit each city . dine at the best restaurants and learn the troubled history of these countries . enjoy hanoi , ha long bay ( overnight luxury cruise ) , danang , hue , saigon & angkor wat . take optional post - tour extension to luang prabang , laos - february 27 - march 2 , 2019 .
glorious mountains , beautiful wines , and the west coast ' s biggest gay pride weekend . this out adventures official tenth - anniversary trip starts in calgary before taking you through the majestic rocky mountains with stops in banff , lake louise and jasper . then we head to the okanagan wine region , where things get extra tasty . you’ll see moose , deer , and bears – oh my ! we wrap up in vancouver just as pride gets underway . boom !
our 12 - day costa rica trip has been designed to maximize your experience here and have you begging to stay forever ! instead of rushing around and trying to visit as many places as possible , we ' ve specially chosen three locations , plus san jose that we know you ' ll fall in love with . costa rica is also known as the most gay - friendly country in central america . we can ' t wait to share this paradise with you !
our 8 - day gay trip through peru mixes our laid - back travel concept with peru ' s most unique and memorable sites to form an adventure that you ' ll be talking about forever . if you ' ve ever dreamed of soaking up the incredible mountaintop inca ruins of machu picchu , taking a scenic train journey through the peruvian andes , or walking in the footsteps of an ancient civilization , join us for an incredible 8 days you ' ll never forget .
patagonia’s wilderness is so renowned there’s an outdoor clothing brand named after it . this trip introduces you to the heaving perito moreno glacier , then heads for an eco - camp in torres del paine national park where we ' ll conquer some of the region ' s most rigorous treks . you ' ll be rewarded daily with unrivalled flora , fauna , and vistas .
gay pride frankfurt , in germany’s financial capital , is one of the country ' s liveliest gay pride parades – with lots of floats and a main stage with 3 - days of entertainment located in the city’s main square . the costumes are pretty outrageous and very inventive . the town’s gay bars come to life – hosting all sorts of parties for all tastes . gay pride frankfurt is also known as frankfurt csd - or ' christopher street day ' .
gay tour of spain . this gay holiday takes an intimate group of travelers to the most picturesque parts of spain . join us and explore the culture , history , and beauty of madrid , toledo , seville , cordoba , and the scenic spanish countryside between . spain is a rich collection of regions , each fiercely proud of its own identity . vacation with a congenial group of gay travelers to sample the cultural life of beautiful & gay friendly spain .
mexico city is the largest metropolis in the western hemisphere and is brimming with rich culture and awe - inspiring architecture . it can be hard to experience it all in one trip , so we’ve lined up plenty of big and hidden gems . we’ll visit ancient pyramids , take walks through interesting neighborhoods , enjoy a wide variety of arts , and have delicious food and drinks along the way .
essential japan is the ideal route for those who visit japan for the first time . we have selected the most iconic cities of the archipelago , so you can take home the true essence of japan . japan has a wide , varied history and is a hub for those who seek authentic culture in a world where that is hard to find . travel from big cities to mountain villages and everything in - between as you take in the rich , cultural heritage this amazing country has to offer on this gay trip .
an industrious yet luxurious wine country bike tour . can ' t decide whether to partake in a hardcore bike holiday , or an elegant wine country affair ? this chile gay trip offers both , for an unforgettable taste of south america ' s wild west . we kick off with a couple days in wine country , then visit atacama desert - the driest place on earth - with great bike rides and a trek thrown in . we ' ll also get a taste of santiago , chile ' s sensational urban center .
india ' s beauty and diversity will enrich you with her culture , stun you with her modernity , embrace you with her warmth , and overwhelm you with her hospitality . . . our gay group tour combines the ultimate luxury with seamless exploration of one of the world ' s most fascinating countries . you will stay in the absolute best hotels and palaces in delhi , jaipur , udaipur , hyderabad , a leopard safari , and jodhpur plus enjoy our pre - trip to nepal ! , march 10 - 12 , 2019
china seems to be on the minds of everyone lately . it ' s economy is booming , its major cities like shanghai are devoloping at a rapid pace , and the olympics in china are aguably more spectacular than any before . there is no better time to come to china , and zoom vacations has created a once - in - a - lifetime trip combining culture , history , glamour , authentic cuisine , and 5 star accommodations throughout , all with the gay sensibilty you ' ve come to expect from zoom .
you haven’t lived until you’ve experienced reykjavík on new year’s eve . pretty much everyone in the city takes a box of fireworks outside just before midnight to blow up the old year then party until dawn . there is nothing quite like it . there really isn’t . new year’s in iceland is the new ‘hot’ destination . we have created a six - night trip for a new year’s celebration in iceland !
a gay adventure tour in key west and the florida keys . from the bustling gay nightlife of miami beach , across the gorgeous islands of the florida keys , our multi - sport adventure will put you on the water , on your bike , and in the sky . enjoy excellent food & hospitality on the way to fabulous , sunny , friendly gay key west .
witness the great wildebeaet migration • lap it up in a luxury tented camp • spot lions in their natural habitat • take an excursion into the magnificent ngorongoro crater • feast on top - notch cuisine • pair this trip with our kilimanjaro climb , or continue your african adventure on a tailor - made extension . spot lions , elephants and buffalo , the elusive leopard in a wildlife enthusiast ' s dream , the serengeti
our 12 - day gay trip through peru mixes our laid - back travel concept with peru ' s most unique and memorable sites to form an adventure that you ' ll be talking about forever . whether you ' ve always dreamed of soaking up the incredible mountaintop inca ruins of machu picchu , taking a scenic train journey through the peruvian andes , or you ' re ready to discover off - the - beaten - track sites like the desert oasis of huacachina and the penguin - filled ballestas islands , peru is ready to blow your mind !
a gay travel turkey cultural tour . from istanbul to the ancient city of ephesus , and the beautiful turquoise coast to the mysterious landscape of the anatolian heartland . on this gay travel turkey tour we’ll explore the best of this fascinating crossroads of civilization . enjoy stunning city sites , archaeological wonders and a boat ride along the coast in a classic vessel on our gay tour of turkey .
rio ' s carnival is perhaps the world ' s most famous celebration . carnival is a spectacular time to come to rio de janeiro , especially for gay travelers ! there are several gay festivals including one in front of our hotel , and people joke that everyone seems a little gay during carnival ! during carnival , in addition to gay pool parties and large dance events , you can also join street festivals and parades all over the city ; and there is an attitude of good will and harmony that pervades everything .
our 12 - day trip through south africa encompases the very parts of south africa that makes it so well known ! think a safari in kruger national park full of lions , hippos , zebras , giraffes . think wine tasting in the the midst of the worlds best vineyards in the winelands . think cosmopolitan cape town where you can dive with great white sharks , summit iconic table mountain , and walk in the footsteps of incredible history .
a defining aspect of dallas ' character is the local lgbt community . the city of dallas has the sixth largest gay population in the u . s . and the largest gay population in the state of texas . approximately 120 , 000 lgbt people live in dallas .
rosa wies ' n - a very gay oktoberfest in munich ! while 6 million people from around the world getting together to drink beer and sing bavarian songs might not sound like a gay haven , for one special sunday during the legendary oktoberfest , it kinda is . while nothing bars homosexuals from attending any day or area of the festival , several events during the festival draw a largely gay , male crowd .
western cuba gay cycling tour . cuba , a country on the cusp of change , has become folkloric . travel with our gay group to penetrate its almost - mythical façade . cycling through the countryside we can truly experience cuba’s heartfelt music , redolent landscapes , and friendly people .
we ' ll stay within the walled town center of zadar , surrounded by all the ancient beauty . we ' ll island hop over to the cosmopolitan island town of hvar for a few days of gorgeous island life . we ' ll find out why dubrovnik was chosen as the set of so many game of thrones episodes . and we ' ll start and finish the trip in the capital and largest city of zagreb with plenty of unique museums and nightlife .
gay pride amsterdam is one of the best and most fun prides in the world . amsterdam pride is a citywide gay festival and attracts several hundred - thousand visitors each year and is one of the largest publicly held annual events in the netherlands . the peak of the festival is during the canal parade , a parade of boats of large variety . there are also street parties in the streets with concentration of gay bars .
our 12 days together in south africa will be surrounded by incredible wildlife , vistas , and opportunites to get out there and take part in some amazing activities ! some of your
must do
activities like a full day safari in kruger park and a full day of wine tasting in the winelands is included in this trip . a real experience in africa sparks not just all five senses , but also imagination and emotions quite unlike any other place you’ll experience .
join this gay scuba diving tour and plunge into the tropical caribbean waters of a gay - friendly island paradise ! discover a colorful world beneath the waves . far from the usual vacation spots , the dutch island of saba is known to scuba divers worldwide for its spectacular underwater landscapes .
do you love a parade ? we’ll be sallying forth to strut our stuff through three of the most picturesque and gay - friendly cities of europe : prague , dresden , and berlin . this gay group adventure comes to a rousing conclusion with gay pride festivities in berlin – the most decadent of the three cities ! prague is one of those magical places you never forget , dresden was once known as “florence on the elbe” .
join italy gay travels on an exclusive gay running & hiking tour in the southeasternmost tip of italy , puglia . spend a full week running or hiking along some of the most beautiful landscapes in italy . the gay running and hiking tour strides along stunning beaches and hectares of ancient olive groves . you set the pace , choose to either run or hike . our experienced guides are there to ensure that you have the best experience .
take this gay tour and immerse yourself in a marvelous discovery of two of the world ' s greatest cities , london and paris . uncover gay history from the renaissance through the modern age while touring england ' s famous gardens and museums and the glorious chateaux of the french loire valley countryside .
join tens of thousands of tourists from all over the world for the annual xlsior gay festival in mykonos , greece . the best djs in the world , the most beautiful mediterranean beaches , and the hottest guys come together on this serence grecian capital of gay vibrancy and pride . spend the days at the beach and the nights partying on the dance floor . no other place in the world loves the proud gay community as much as mykonos .
budapest is often called the paris of the east . its grand boulevards , sumptuous architecture , and focus on food are not to be rushed . then on to the alpine sweetheart , slovenia . spend two nights in private wooden huts where we explore local lakes , the marvellous alps and kilometres of caves . and finally , ljubljana offers a little taste of the big city . here we’ll enjoy top - notch cuisine , a lovely boat ride , and a day trip to the goriška brda wine region .
join italy gay travels for the luxury gay salento food and wine tour ! welcome to salento , the southeastern - most tip of puglia , the heel of the boot of italy . our luxury gay salento food and wine tour is your opportunity to explore the world - famous food in this amazing location , connecting ionian and adriatic seas . our tour combines the must - see sights of the area with a sophisticated exploration of the best food and wines of southern italy .
for 200 years , the kings of france outdid one another as they built palaces along the loire valley . bicycle to amboise , chenonceau ( where gay henri iii held drag parties ) , and azay - le - rideau . our gay french bike tours are the perfect way to travel between the castles of france .
a gay travel cultural tour visiting budapest , vienna , prague , and more ! during the height of the holy roman empire , central europe nurtured architectural and musical influences that continue to this day . enjoy our gay cultural tours with walking , concerts , a dinner cruise , rejuvenating baths , and ample time to explore
this intriguing country of gods and goddesses , pharaohs and fantasies will inspire and surprise you as it has done so to its travelers for centuries . but egypt is so much more than mythology and magic , and offers more to see and do than ruins and camel rides . zoom vacations trip to egypt brings out the rich history , culture and cuisine in addition to the things every visitor to the country must experience . highlight will likely be our 4 night cruise aboard the sanctuary zein nile chateau , perhaps the most elegant boat on the nile .
india . . . a country which will enchant you with her beauty and diversity , enrich you with her culture , stun you with her modernity , embrace you with her warmth , and overwhelm you with her hospitality . . . zoom vacations combines the ultimate luxury with seamless exploration of one of the world ' s most fascinating countries . you will stay in the absolute best hotels and palaces in delhi , agra , jaipur , udaipur and mumbai ( bombay ) . plus enjoy our pre - trip to nepal ! , september 21 - 23 , 2018
an exquisite gay cultural tour of italy with a congenial gay group exploring the amalfi coast , capri and the enchanting regions of basilicata and puglia . we ' ll delve into the history of pompeii , and delight in the natural splendor of the region , admiring the tenacity with which ancient residents settled the cliffs and islands of the mediterranean .
rendez - vous in paris in 2018 will be greatest sporting and cultural event , where we will share together some moments rich in emotion along with up to 15 , 000 attendees from over 70 countries . since 1982 , the gay games have brought together people from all over the world , with diversity , equality , solidarity , and respect . with sport a common theme for inclusion , gay games are open to all , young or old , athlete or artist , experienced or novice , gay or straight .
the biggest gay and lesbian celebration in bavaria is the munich christopher street day ! munich is the city of the oktoberfest , but munich has much more to offer - particularly for gays and lesbians . the gay and lesbian community energise munich during the city’s annual csd celebrations . the munich city centre explodes with drag queens , high heels and feather boas .
join italy gay travels for a culinary delight in puglia the south of italy . our authentic and extravagant gay puglia foodies tour is a once in the lifetime opportunity to explore the world - famous food in this amazing location . our tour combines the must - see sights of puglia with a sophisticated exploration of the food and wine specific to the region .
more than a week of party , lots of party and more than a week of fun , lots of fun . come to experience this amazing 8 days gay tour in three of the top 10 world gay hotspots : barcelona , sitges and ibiza . we will visit the best bars , clubs , saunas . . . but not everything is partying . we will also visit the best gay beaches , the best sunsets and we will taste the authentic spanish food . get ready for the best possible combination of pleasure and fun .
a gay travel italian hiking tour in tuscany . enjoy a week of fun on this gay hiking tour in italy . see florence and the fairy - tale village of san gimignano . trek through orchards , vineyards , and woodland on our way to radda to visit ancient stone villages , churches , farms , and etruscan bridges , arches and burial mounds .
tropout – the festival for the modern gay traveller – is finally ready to come to europe ! and what a place to start – malta is europe’s number one ranked destination for gay travellers . this is one party you won’t want to miss . we’ve created a festival that mixes the glamour of the european summer with prime party season – all in a location that will amaze you with its landscapes , history and culture . if you’re seeking an unforgettable mediterranean experience with likeminded gay guys , you’ve found it . welcome to tropout malta .
pinkish movement in asia ! be a witness on asia ' s first gay - focused flight from tokyo to pride parade in taiwan . don ' t miss it ! nowadays , taiwan is one of the hottest destinations in asia for lgbt travellers , because there are so many lgbt activities in taipei city . the people of taiwan is what makes it , not to mention the delicious food ! the biggest gay event in taiwan is taipei gay pride with more then 80 , 000 people from around the world attending this 4 day event .
named by out traveler magazine as one of the top 10 trips to change your life ! plus , our post - trip to lake titicaca ! for thousands of years , peru ' s extraordinary beauty has been both the backdrop of incredible cultural and spiritual expression , and a pallet for architectural artists . your journey will take you through the cultural heart of lima , charming cusco , and the valley of the incas . but perhaps what will impress and inspire you most is your visit to one of the world ' s most important archeological sights : the ancient mystical city of machu picchu .
what is the best place to visit if you’re interested in the gay past ? there is a lot of competition but greece often falls at the top . here ' s why .
a beautifully observed 1950s - period coming of age romance between a young shop girl and aspiring photographer , therese ( rooney mara ) , and an impossibly glamorous but tormented older woman , carol ( cate blanchett ) , from acclaimed director todd haynes . therese joins carol on an impulsive getaway , as they road - trip west from new york city with no destination planned ( although they don ' t stop there in the movie itself , the many gorgeous shots of both manhattan and chicago were in real life filmed in cincinnati ) . both blanchett and mara deliver knockout performances and form a cinematic couple for the ages .
this wonderfully relaxed and interesting gay group tour to sri lanka gives tour members the major tourist attractions of the country in comfort and in the company of other gays . it is a varied itinerary that includes historical sites , beaches , national wildlife parks and cultural experiences . plus the fun of the gay group with welcome and farewell parties and the opportunity of making new friends .
at club atlantis , we take over the entire 5 - star resort to create a relaxing and friendly all - gay beach vacation like no other . our all - inclusive resort includes deluxe accommodations , all meals , all drinks , sports , activities , entertainment , parties , gratuities , wireless internet , and so much more . it ' s the best gay vacation value on the planet !
known by locals as csd berlin ( christopher street day ) , and by the rest of us as gay pride berlin , offers one of the most eclectic pride celebrations in europe . on christopher street day , hundreds of thousands of people demonstrate for the rights of lgbt people with a great parade , colourful floats , and music . csd berlin is one of the biggest gay prides in europe .
join us for a quick holiday getaway to one of america’s most interesting cities : new orleans , louisiana . our gay new orleans holiday weekends are full of campy fun ! we’ll stay in the french quarter and explore the sultry gay history of the “big easy” , while enjoying a touch of southern decadence . attend a jazz brunch and get ready for mardi gras in an atmosphere of unique new orleans kitsch .
an active scotland gay adventure with stops in glasgow and edinburgh . the scottish highlands offer some of the most glorious hikes in the world . this 8 - day gay adventure features four hikes , a few isles , and time in both the capital of edinburgh and scotland’s largest city , glasgow . if you’re not a hardcore hiker , you can skip the most challenging day and spend it exploring brodick castle instead .
circuit festival is one of the biggest and most important gay festivals worldwide . taking place in the city of barcelona , circuit festival offers you ten days of themed parties as la leche , papa party or the water party , dj sets , artists and many other incredible activities . . . happy gay travel is the official travel agency for circuit gay festival 2018 , so we have tailored a complete ‘festival package’ deal which include an official wristband that grants access to all the parties of the circuit festival , including the water party , papa party and la leche and a choice from 2 * to 5 * accommodation 7 nights stay .
stroll through lima ' s vibrant plaza mayor , learn the secrets of the amazon jungle from a local guide , wake to the sounds of an amazonian animal orchestra , admire cuzco ' s stunning colonial architecture , glimpse terraced walls in the fertile sacred valley , stroll geometrically perfect streets in ollantaytambo , wander among the ancient ruins of machu picchu , discover colourful traditional textiles on our gay peru , amazon & machu picchu gay tour .
dance . sun . swim . the perfect weekend in the the most amazing location ! as the weekend comes in and the parties only get hotter , join tens of thousands of tourists from all over the world for the annual xlsior gay festival in mykonos , greece . the best djs in the world , the most beautiful mediterranean beaches , and the hottest guys come together on this serene grecian isle of gay vibrancy and pride .
the ultimate female buddy movie . while not an explicitly gay film , it is easy to see why the lgbt community champions this rollicking feminist story . thelma and louise ( played to perfection by geena davis and susan sarandon ) are two arkansas women who set out for a weekend trip in the mountains in louise ' s ' 66 ford thunderbird convertible . things quickly take a turn for the worse when a man thelma dances with at a roadhouse attempts to rape her . louise , in an effort to protect her friend , loses her temper and shoots and kills the man . in fear that no one will believe their claim of attempted rape , the two run from the law , driving along the open roads of the southwest , ultimately winding up at the grand canyon , in one of the most unforgettable finales in hollywood history . and who can forget brad pitt ' s star - making turn as the shirtless cowboy ?
costa rica is also known as the most gay - friendly country in central america , with a lively gay social scene in san jose and manuel antonio , and some of the friendliest local latin people you could ever meet ! this is a true latin experience from the people , to the salsa dancing , to the gallo pinto for breakfast . we genuinely love costa rica and can ' t wait to share this paradise with you !
we invite you to join our gay tanzania safari & explore of one of the most well - preserved wilderness areas on earth . tanzania has more land devoted to wildlife than anywhere else in the world . see what draws celebrities from around the world to spend days or even weeks in tanzania . walk on the wild side in tanzania ! this gay safari hits all the best spots , from lake manyara , to the expansive savannahs of the serengeti .
egypt has enchanted visitors throughout history and our gay egypt tour is no exception . join our small gay group us as we sail down the nile on the ss karim , a traditional paddle steamer built for kings . the riches of antiquity are our destination and our backdrop as we cruise through the land of the pharaohs . egypt , the wonder of travelers for thousands of years , calls us for a pilgrimage to the treasures of this historic land .
tropout koh samui – the official after white party gay travel experience – gives those in bangkok ( and elsewhere ) celebrating new year festivities the opportunity to start 2019 in the tropout style . so much more than a traditional ‘resort holiday’ , tropout gay resort week is filled with opportunities to chill , party & explore in a tropical paradise with guys from all over the globe in stylish settings . come join us in tropical koh samui and experience it all !
join italy gay travels on an exclusive gay yoga retreat in the southernmost tip of italy , known as puglia . we will spend a full week in a typical masseria ( farmstead ) set within acres of olive groves and nearby to stunning beaches . the farmstead is close to ostuni , known as the ‘white city’ as it sits perched on the rocks overlooking the adriatic sea with acres and acres of ancient roman olive groves as far as the eye can see .
12 - day all - inclusive peruvian adventure including an amazon river cruise and culminating with a comprehensive machu picchu experience . it ' s the perfect combination of inspiring places and rich history in the comfort of an all - gay environment .
13 - day customized luxury gay group tour of japan , starting in tokyo . explore some of japan’s top destinations , epic sites and off - the - beaten - track hidden gems . our gay group tour includes three nights in tokyo , one of the world’s most cutting - edge capitals . experience two nights at a traditional japanese ryokan , three nights in kyoto , the historic former capital , famous for its abundance of temples , gardens , and unesco world heritage status .
halloween & day of the dead with vacaya in mexico ! vacaya completely reinvents the all - lgbt all - inclusive resort experience with the 5 - star unico riviera maya . as the newest member of the leading hotels of the world , unico offers a level of luxury and inclusions never before seen in the world of all - gay travel . throw in halloween and mexico’s day of the dead and this exclusive 7 - day all - gay resort holiday experience is without compare !
the remote middle fork of the salmon river runs cold , clear , and quick , past soaring cliffs and soothing hot springs . join us for approximately 90 miles exhilarating gay whitewater rafting adventure . camp on large sandy beaches and and explore burbling side creeks and quiet hiking trails . our gay rafting adventure begins near the sawtooth mountains of idaho . the main salmon river is known as the
river of no return
, and runs cold and clear through vast , deep canyons and forests .
10 days egypt gay tour & nile river cruise from / to cairo , egypt . the eternally staggering sights of cairo and ancient egypt in top comfort . 4 nights in 5 - star hotels , plus 4 nights sailing the nile aboard the 40 - cabin ultra - luxury sanctuary sun boat iv in the cabin of your choice . this four night luxury nile gay cruise , beginning in luxor and ending in aswan , takes in the highlights of upper egypt on board the finest floating boutique hotel .
italy gay history tour from caesar to michelangelo and beyond . the ancient greeks , the ancient romans , think of julius caesar and nero , etc . , plus all those renaissance artists—donatello , botticelli , leonardo , michelangelo , caravaggio : did it ever occur to you how much italian gay history and art there is ? well , let us tell you : italy not only has some of the most beautiful cities , the most great art , one of the world’s top cuisines , some of the cutest guys .
spend a week cycling along ancient roman lanes , coastal drives and mountain paths . each night , relax at an authentic gay owned and operated agriturismo , enjoying a glass of wine from the villa ' s vineyards , gourmet meals , and a dip in the pool , with cute locals always stopping by to enjoy the festivities . during our gay italy bike tour in puglia , we ' ll have five biking days , designed in a ' roller coaster ' fashion , with light days in the beginning and end .
join this gay tour and witness the extremes of iceland : friendly viking descendants ; unspoiled lush wilderness ; fuming volcanic craters , geysers and natural hot springs ; rainbow - festooned waterfalls ; playful puffins , and a rich history including some of folklore ' s strangest sagas .
two teenage boys from mexico city , julio ( gael garcia bernal ) and tenoch ( diego luna ) , meet a beautiful spanish woman , luisa ( maribel verdu ) , and manage to convince her to join them on a road trip through the mountains , desert , and jungles of central mexico to a secluded pacific beach they claim no one else knows about . the film garnered international acclaim not only for its honest depiction of adolescent sexual relationships , but also for the story ' s deep thematic explorations of loss , death , intimacy , and the cultural and political realities of a burgeoning new era in mexico ' s complicated history . y tu mama tambien , along with a handful of other works by its creator ' s contemporaries , would go on to usher in a new wave of mexican cinema and establish writer - director alfonso cuaron as one of the great international talents of his generation .