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Indeed I suppose they would never have made use even of this had not the English introd[u]ced vessels to put their wine in which were rather too large to be carried by hand, as they used to do every thing else. A speech of their late governeur is recorded here, which shews in what light they are lookd upon even by the Portugese, themselves I beleive far behind all the rest of Europe, except possibly the Spaniards : it was very fortunate said he that this Island was not Eden in which Adam and Eve dwelt before the fall, for had it been so the inhabitants here would never have been induc'd to put on Cloaths; so much are they resolvd in every particular to follow exactly the paths of their forefathers.
Indeed were the people here only tolerably industrious, there is scarcely any Luxury which might [not] be produc'd that either Europe or the Indies afford, owing to the great difference of Climate observable in ascending the hills; this we experien[c]d in a visit to Dr Heberden, who lives about two miles from the town, we left the Thermometer when we set out at 74 and found it there at Indeed the hills produce almost spontaneously vast plenty of Wallnutts, chestnutts, and apples, but in the town you find some few plants natives of both the Indies, whose flourishing state put it out of all doubt that were they taken any care of they might have any quantity of them.
Of these I mention some: the Banana tree, Musa sapientum Linn. These without mentioning any more seem very sufficient to shew that the tenderest plants might be cultivated here without any trouble; yet the indolence of the inhabitants is so great, that even that is too much for them; indeed the policy of the English here is to hinder them as much as possible from growing any thing themselves except what they find their account in taking in exchange for Corn, tho the people might with much Less trouble and expence grow the corn themselves.
What corn grows here, which indeed is not much, is of a most excellent quality, Large graind, and very fine; their meat also is very good, mutton, pork, and beef more especialy, of which what we had on board the ship was agreed by all of us to be very little inferior to our own; tho we Englishmen value ourselves not a little on our peculiar excellence in that production.
The fat of this was white like the fat of mutton, yet the meat Brown, and coarse graind as ours, tho much smaller. The town of Fonchiale is situated at the Bottom of the Bay, very ill Built, tho larger than the size of the Island seems to deserve. The houses of the bettermost people are in general large but those of the poorer sort very small, and the streets very narrow and uncommonly ill pavd.
The Churches here have abundance of ornaments, cheifly bad pictures and figures of their favourite saints in lac'd cloaths; the Convent of the Franciscans indeed which we went to See had very little ornament; but the neatness with which those fathers kept everything was well worthy of commendation, especialy their infirmary, the contrivance of which deserves to be taken particular notice of; it was a long room, on one side of which were windows, and an altar for the convenience of administering the sacrament to the sick; on the other were the wards, each just capable of containing a bed, and lind with white duch tiles; to every one of these was a door communicating with a gallery which ran paralel to the great room, so that any of the sick might be supplied with whatever they wanted without disturbing their neighbours.
In this Convent was a curiosity of a very singular nature; a small chapel whose whole lining, wainscote, and ceiling, was intirely compos'd of human bones, two large thigh bones across, and a skull in each of the openings. Among these was a very singular anatomical curiosity, a skull in which one side of the Lower jaw was perfectly and very firmly fastned to the upper by an ossification, so that the man whoever he was must have livd some time without being able to open his mouth, indeed it was plain on the other side that a hole had been made by beating out his teeth, and in some measure damaging his Jaw bone, by which alone he must have receivd his nourishment.
I must not leave these good fathers without mentioning a thing which does great credit to their civility, and at the same time shews that they are not bigots to their religion: we visited them on Thursday Even just before their supper time; they made many apologies that they could not ask us to sup, not being prepard; but said they, if you will come tomorrow, notwithstanding it is fast with us, we will have a turkey roasted for you.
There are here, beside friarys, 3 or 4 houses of nunns. To one of these Sa'nta Clara we went, and indeed the ladies did us the honour to express great pleasure in seeing us there; they had heard that we were great Philosophers, and expected much from us, one of the first questions that they askd was, when it would thunder; they then desird to know if we could put them in a way of finding water in their convent, which it seems they were in want of; but notwishstanding our answers to these questions were not quite so much to the purpose as they expected, they did not at all cease their civilities, for while we stayd, which was about half an hour, I am sure there was not the fraction of a second in which their tongues did not go at an uncommonly nimble rate.
It remains now that I should say something of the Island in general, and then take my leave of Madeira till some other opportunity offers of visiting it again, for the climate is so fine that any man might wish it was in his power to live here under the benefits of English laws and liberty. The hills here are very high, much higher than any one would imagine, Pico Ruevo the highest is ft which is much higher than any land that has been measured in Great Britain; indeed as I hinted before the whole Island has probably been the production of a Volcano, notwishstanding which its fertility is amazing, all the sides of the hills are coverd with vines to a certain hight, above which are woods of chestnut and pine of immense extent; and above them forests of wild timber of kinds not known in Europe, which amply supply the inhabitants with whatever they may want.
Among these some there were whose flowers we were not able to procure and consequently could not settle their Genera, particularly those calld by the Portugese Mirmulano and Pao branco , both which, and especialy the first, from the Beauty of their leaves promise to be a great ornament to our European gardens.
The inhabitants here are supposd to be about 80,; and from the town of Fonchiale its custom house I mean the King of Portugal receives pounds a year, after having paid the Governor and all expences of every kind, which may serve to shew in some degree the consequence which this little Island is of to the crown of Portugal; was it in the hands of any other people in the world its value might easily be doubled, from the excellence of its climate capable of bearing any kind of crop, a circumstance which the Portugese do not make the least advantage of.
This Evening every thing being ready for sea, we went on board, and at 8 o'Clock got under way with a very light breeze. Still almost calm, which gave us an opportunity of taking with the casting nett a most beautifull species of Medusa, of a colour equaling if not exceeding the finest ultramarine; it was describd and calld Medusa azurea.
This morn wind foul, saw however some rocks call'd in the old charts Salvages which lay to the northward of the Canarys. No land in sight this morn, towards noon almost calm, many fish were about the ship, but our fishermen could not contrive to catch any of them. This morn we were calld up very early to see the pike of Teneriffe, which now for the first time appeard at a vast distance much above the clouds I mean those which form a bank near the Horizon ; the hill itself was so faint, that no man who was not used to the appearance of land at a great distance could tell it from a cloud, it soon however appeard something clearer and a sketch was made of it.
While we were engagd in looking at the hill a fish was taken which was describ'd and called Scomber serpens ; the seamen said they had never seen such a one before except the first lieutenant, who rememberd to have taken one before just about these Islands; Sr Hans Sloane in his Passage out to Jamaica also took one of these fish which he gives a figure of, Vol.
The Pike continued in sight almost all day, tho sometimes obscurd by the clouds; at sunset however its appearance was most truely elegant, the rays of the sun remaining upon it sometime after it was set and the other land quite Black, and giving it a warmth of colour not to be express'd by painting. This Morn the Pike appeard very plain and immensely above the clouds, as may well be imagin'd by its hight which Dr Heberden of Madeira who has been himself upon it communicated to us, 15, feet.
The Dr also says that tho there is no eruption of visible fire from it, yet heat issues from the chinks near the top so strongly that a person who putts his hand upon these is scalded; from him we receivd among many other favours some salt which he supposes to be the true natron or nitrum of the ancients, and some native sulphur exceedingly pure, both which he collected himself on the top of the mountain, where large quantities, especialy of the salt, are found on the surface of the Earth.
Wind continued to blow much as it had done so we were sure we were well in the trade; now for the first time we saw plenty of flying fish, whose bea[u]ty especialy when seen from the cabbin windows is beyond imagination, their sides shining like burnishd silver; when seen from the Deck they do not appear to such advantage as their backs are then presented to the view, which are dark colourd. Went as usual and as we expect to go these next two months; flying fish are in great plenty about the ship.
About one today we crossd the tropick, the night most intolerably hot, the Thermometer standing all night at 78 in the cabbin tho every window was open. About one this morn a flying fish was brought into the cabbin, the first that had been taken; it flew aboard, I suppose chasd by some other fish, or maybe merely because he did not see the ship; at breakfast another was brought, which had flown into Mr Green the Astronomers Cabbin.
This whole day we saild at the rate of 7 knotts, sometimes a fathom or two more the wind being rather stronger than it usualy is in the trade. Wind rather slackend; three birds were today about the ship, a swallow, to all appearance the same as our European one, and two motacillas, about night fall one of the latter was taken; about 11 a shoal of Porpoises came about the ship, and the fisgig was soon thrown into one of them but would not hold.
This morn calm; employd in drawing and describing the bird taken yesterday, calld it Motacilla avida ; while the drawing was in hand it became very familiar, so much so that we had a brace made for it in hope to keep it alive; as flies were in amazing abundance onboard the ship we had no fear of plentiful supply of provision.
About noon a young shark was seen from the Cabbin windows following the ship, who immediately took a bait and was caught on board; he provd to be the Squalus Charcharias of Linn[aeus] and assisted us in clearing up much confusion which almost all authors had made about that species; with him came on board 4 sucking fish, echineis remora Linn. Notwistanding it was twelve O'Clock before the shark was taken, we made shift to have a part of him stewd for dinner, and very good meat he was, at least in the opinion of Dr Solander and myself, tho some of the Seamen did not seem to be fond of him, probably from some prejudice founded on the species sometimes feeding on human flesh.
This Morn at day break made the Island of Bonavista, one of the Cape Verde Islands: Mr Buchan employd in taking views of the land; Mr Parkinson busy in finishing the sketches made of the shark yesterday. This Evening the other Motacilla avida was brought to us, it differd scarce at all from the first taken, except that it was something larger; his head however gave us some good, by supplying us with near twenty specimens of ticks, which differd but little from the acarus vicinus Linn; it was however described and calld Motacilla.
This Morn Bonetos were in great plenty about the ship, we were calld up early to see one that had been struck, found it to be the Scomber Pelamis Linn. After having examind and drawn the animal we proceeded to disect him, and in the course of the operation were much pleasd by the infinite strength we observd in every part of him, specialy the stomack, the coats of which were uncommonly strong especialy about the sp[h]incter, or extremity by which the digested meat is discharged; this I suppose is intended to crush and render usefull the scales and bones of fishes which this animal must continualy swalow without seperating them from the flesh.
From the inside of its scales we took a small animal who seemd to be a louse if I may so call it as it certainly stuck to him and preyd upon the Juices which it extra[c]ted by sucktion, probably much to his disquiet; it provd to be monoculus piscinus Linn. In the inside of the fish were also found two animals which preyd upon him, one in his very flesh tho near the membrane which covers the intestines, Fasciola Pelamines Mss, the other in the stomach sipunculus piscium Mss.
This morn two swallows were about the ship, tho we must now be 60 Leagues at least from any land, at night one of them is taken and proved to be Hirundo domestica Linn.
This morn the other swallow was found dead upon the deck; now for the first time we have lost the trade, and expected calm and squally weather till we shall cross the line. The trade had now lasted us pretty free from squalls or calms these [] days it has been in general between [] but ever since we have been in it the air has been uncommonly damp, every thing more than usualy liable to mould, and all Iron work to rust, the air has seldom been Clear, but a haize in it which was even perceiveable to the human frame.
Today quite calm, I went out in a boat and took dagysa strumosa , medusa porpita , the same which we before calld asurea, mimus volutator , and cimex [] who runns upon the water here in the same manner as C.
Lacustris does on our ponds in England. Towards even two small fish were taken under the stern, they were following a shirt which was towing and showd not the least signs of fear, so that they were taken with a landing net without the least difficulty.
Balistes monoceros Linn. Weather pretty good, at night a squall with Lightning and rain, another swallow came to the ship today and was taken with the snippers as soon as he went to roost. Blew fresh this morn with heavy rain; towards noon five swallows came on board and were taken at roosting time, and provd like all we have taken before to be H. This morn calm; went out in the boat and took what is calld by the Seamen a Portugese man of war, Holothuria Physalis Linn; also Medusa velella L.
The Holothuria proved to be one of the most bea[u]tifull sights I had ever seen, it consisted of a small bladder in shape much like the air bladder of fishes, from the bottom of which descended a number of strings of bright blue and red, some three or four feet in length which if touchd stung the person who touchd them in the same manner as nettles, only much stronger: on the top of this Bladder was a membrane which he turnd either one way or the other as the wind blew to receive it, this was veind with pink in an uncommonly beautifull manner, in short the whole together was one of the most beautifull sights I have seen among the mollusca, tho many of them are beautifull.
The floating shells H. Janthina and violacea from their particularity deserve also to be mentiond, they are found floating on the top of the water by means of a small cluster of Bubbles filld with air, which are composd of a tenacious slimey substance, not easily parting with its contents; these keep him suspended on the surface of the water and serve as a hiding for his Eggs, and it is probable that he never goes down to the bottom, or willingly comes near any shore, as his shell is of so brittle a construction that few fresh water snails are so thin.
Every shell contains within it about a teaspoonfull of Liquid, which it easily discharges on being touched, this is of a most beautifull red purple colour and easily dies linnen clothes; it may be well worth inquiry whether or not this is the purpura of the ancients as the shell is certainly found in the Mediterranean.
We have not yet taken a sufficient quantity of the shells to try the experiment, probably we shall do soon. Procellaria oceanica differs very little from P. This morn a shark calld us out of our bedds, and was soon hookd, but as soon broke his hold and went off: at noon went out in the boat but found nothing on the surface of the water; on returning home however found on the sterm of the ship two new species of Lepas vittata and midas , they were both sticking to the bottom in company with the anatifera , of which there was great abundance.
After dinner calld upon deck by another shark, who had been lately wounded by a harpoon, but he was two cunning after his misfortune to bite at our baits, which we much Lamented as he had sucking fish upon him that were quite white, probably a species not yet describd. Went out in the boat today, took plenty of Helix Janthina and some few of violacea , shot the black toed gull of Penn. It had not yet been describd according to Linnaeus's system, so calld it Larus crepidatus ; its food here seems to be cheifly Helixes which appeard probable at least, on account of its dung being of a lively red colour, much like that which was procurd from the shells.
I was drove home from this excursion by a very heavy squall of rain, which intirely wetted me through long before it was possible to return to the ship, however I receivd no other harm from the ducking than the present inconvenience of being so thoroughly wet. The remainder of today was very squally, with much rain; indeed it has been so ever since we lost the trade, and the people who have been here before say that it is generaly so in these latitudes; I can liken it to nothing so much as April in England, when it is very showery, the weather is never certain for two hours, or indeed half the time, tho calms spend much the greatest part of our time in idleness.
Today much like yesterday, very squally; saw a dolphin, and admired the infinite beauty of his colour as he swam in the water, but in vain, he would not give us even a chance of taking him. A shark, squalus carcharias Linn. The pilot fish Gasterosteus ductor Linn. The blubbers taken today were Beroe Labiata and Marsupialis Mss, the first of which made a pretty appearance in the water, by reason of its swimmers, which line its sides like fringes, and are of a changeable fire colour; Callirhoe bivia Mss, the most lifeless lump of Jelly I have seen, it scarcely seems to be possessd of life but for one or two motions we saw it make.
Calm this Morn; a shark was taken, but not one pilot fish attended him, which is rather uncommon as they seldom are without a shoal of from ten to twenty. At noon I went in the boat, and took the Sallee man Phyllodoce velella Linn. We saw several of the latter today, and observd many small fish under their tentacula who seemd to shelter there as if with its stings it could defend them from large enemies.
Ventur'd out today, but found the surface of the water so ruffled that nothing at all floated upon it, I had the good fortune however to see a bird of the shearwater kind which I shot, and it provd to be not describd; it was about as large as the common but differd from it in being whiter, especialy about the face: calld it Procellaria crepidata , as its feet were like the gulls shot last week, black without but white near the leggs. A large shoal of fish were all this day under the shipp's stern, playing about, but refusing to take bait; we however contrivd to take one of them with a fish gigg, which provd not describd; it was in make and appearance like a Carp, weighing near two pounds, its sides were ornamented with narrow yellow lines and its finns almost intirely coverd with scales: calld it Chaetodon cyprinaceus.
A fine breeze of wind started up last night which held us all day, so I found it impossible to go out in the boat; tonight however to make these 24 hours not intirely unprofitable I had the opportunity of seeing a Phenomenon I had never before met with, a lunar rainbow which appeard about ten O'Clock very faint and almost or quite without colour, so that it could be tracd by little More than an appearance which lookd like shade on a cloud.
This morn went out in the boat but caught no one thing, I had never been before so unfortunate. Wind continued to blow fresh so we had little doubt of the reality of yesterdays hopes. This evening trying as I have often foolishly no doubt done to exercise myself by playing tricks with two ropes in the Cabbin I got a fall which hurt me a good deal and alarmd me more, as the blow was on my head, and two hours after it I was taken with sickness at my stomack which made me fear some ill consequence.
Today thank God I was much better and easd of all apprehensions, the wind continuing fair and I had given over all thoughts of boat expeditions for some time at least.
Trade continues. Today the cat killd our bird M. Avida who had lived with us ever since the 29th of Septr intirely on the flies which he caught for himself; he was hearty and in high health so that probably he might have livd a great while longer had fate been more kind. Trade had got more to the Southward that it usualy had been, which was unlucky for me as I proposd to the Captain to touch for part of a day at least at the Island of Ferdinand Norronha, which he had no objection to if we could fetch it: that however seemd very uncertain.
This Evening we saw 6 or 7 large fish of the whale kind which the Seamen calld Grampuses tho I think they were very different from the fish commonly so calld; they were however Certainly of the whale kind and blew throug[h] two? They had heads smaller and rounder than those fish in general have and very low back finns and very small tails; thus much was all that I could see as they never came within two cables lengh of the ship.
Trade today was still more to the Southward, almost due South, so that we tackd and stood to the eastward lest we should fall in with the coast of Brazil to the Northward of Cape Frio. Wind today as fair as we could wish, ship layd up so well that it renewd our hopes of touching at the Island.
About noon today we experiencd what the Seamen call a white squall, that is a gust of wind which came upon us quite unawares, unattended with a cloud as squalls in general are and therefore took us quite unprepard; it was however very slight so no ill consequence ensued except Mr Parkinson and his potts going to leward, which diverted us more than it hurt him.
This morn about 8 O'Clock crossed the Aequinoctial line in about 33 degrees West Longitude from Greenwich, at the rate of four knotts which our seamen said was an uncommonly good breeze, the Thermometer standing at The Thermometers used in this voyage are two of Mr Birds making after Farenheights scale, which seldom differ above a degree from each other and that not till they are as high as 80, in which case the medium between the two instruments is set down.
This Evening the ceremony of ducking the ships company was performed as always customary on crossing the line, when those who have crossd it before Claim a right of ducking all that have not, the whole of the ceremony I shall describe. About dinner time a list was brought into the cabbin containing the names of every body and thing aboard the ship, in which the dogs and catts were not forgot; to this was affixd a petition, sign'd 'the ships company,' desiring leave to examine every body in that List that it might be know[n] whether or not they had crossd the line before.
This was immediately granted; every body was then calld upon the quarter deck and examind by one of the lieutenants who had crossd, he markd every name either to be duckd or let off according as their qualifications directed. Captn Cooke and Doctor Solander were on the Black list, as were my self my servants and doggs, which I was oblig'd to compound for by giving the Duckers a certain quantity of Brandy for which they willingly excusd us the ceremony. Many of the Men however chose to be duckd rather than give up 4 days allowance of wine which was the price fixd upon, and as for the boys they are always duckd of course; so that about 21 underwent the ceremony which was performd thus:.
A block was made fast to the end of the Main Yard and a long line reved through it, to which three Cross peices of wood were fastned, one of which was put between the leggs of the man who was to be duckd and to this he was tyed very fast, another was for him to hold in his hands and the third was over his head least the rope should be hoisted too near the block and by that means the man be hurt.
When he was fas[t]ned upon this machine the Boatswain gave the command by his whistle and the man was hoisted up as high as the cross peice over his head would allow, when another signal was made and immediately the rope was let go and his own weight carried him down, he was then immediately hoisted up again and three times served in this manner which was every mans allowance.
Thus ended the diversion of the day, for the ducking lasted till almost night, and sufficiently diverting it certainly was to see the different faces that were made on this occasion, some grinning and exulting in their hardiness whilst others were almost suffocated and came up ready enough to have compounded after the first or second duck, had such proceeding been allowable. It is now time that I should say something of the climate and degree of heat since crossing the tropick, as we have been for some time within the bounds which were supposd by the ancients to be uninhabitable on account of their heat.
Almost immediately on crossing the tropick the air became sensibly much damper than usual, tho not materialy hotter, the thermometer then in general stood from 80 to The nearer we approachd to the calms still the damper every thing grew, this was perceivable even to the human body and very much so, but more remarkably upon all kinds of furniture: every thing made of Iron rusted so fast that the knives in peoples pockets became almost useless and the razors in cases not free.
All kinds of Leather became mouldy, Portfolios and truncks coverd with black leather were almost white, soon after this mould adheerd to almost every thing, all the books in my Library became mouldy so that they were obligd to be wiped to preserve them.
About this time we came into the calms which we met with earlier than usual; the thermometer was then at 83 and we sufferd from the heat and damp together.
Bathing however kept me in perfect health, tho many of the ship[s] company were ill of bilious complaints which however were but of short duration.
This continued till we got the S. The air during the whole time sin[c]e we crossed the tropick and indeed sometime before has been nearly of the same temperature throughout the 24 hours, the Thermometer seldom rising above a degree during the time the sun is above the horizon. The windows of the cabbin have been open without once being shut ever since we left Madeira. Last night and today the weather has been squally, wind rather fresh but keeping very much to the Southward; great plenty of flying fish have been about the ship few or none of which have been seen since we left the N.
Fine weather but Wind rather too much to the Southward. We are today nearly in the latitude of the Island of Fernand de Norhona, so that I am not without hopes of making it if rightly laid down; night however put an end to our hopes for the present at least, and left us in no very agreable situation as shoals and foul ground is laid down all round the Islands. Fine breeze today, our hopes of seeing the Island were again renewd but without success, so at night we judge ourselves to be past it and that the longitude is wrong laid down.
Wind East very pleasant, we now gave up all thoughts of the Island. This Evening the sea appeard uncommonly bea[u]tifull, flashes of light coming from it perfectly resembling small flashes of lightning, and these so frequent that sometimes 8 or ten were visible at the same moment; the seamen were divided in their acco[u]nts some assuring us that it proceeded from fish who made the light by agitating the salt water, as they calld it, in their darting at their prey, while others said that they had often seen them and knew them to be nothing but blubbers Medusas.
This made us very Eager to procure some of them, which at last we did one by the help of the landing net. They prov'd to be a species of Medusa which when brought on board appeard like metal violently heated, emitting a white light; on the surface of this animal a small Lepas was fixd exactly the colour of it, which was almost transparent not unlike thin starch in which a small quantity of blue is disolv'd.
This Morn employd in Examining the things caught last night, which being taken by the light of our lamps for the wind which blows in at the windows always open will not suffer us to burn candles we could hardly then distinguish into genera, much less into species, had the good fortune to find that they were all quite new.
Calld them Medusa pellucens , Lepas pellucens ,[], Clio [], Cancer fulgens and Cancer amplectens , but we had the misfortune to loose two more species of Crabbs overboard by the tumbling of a glass overboard in which they were containd. In the Evening the Sea was lighted in the same manner as it was last night only not near so strongly; we renewd however our endeavours to take some of the light carriers, not without success as two new species of Crabbs were taken one of which was very singular.
Nothing to be done today, found however that the crabbs taken yesterday were both new, calld them vitreus and Crassicornis. A shoal of small fish were today under our stern who attended the ship for some time; she had however too much way through the water for our instruments so we could not take any of them.
This day was quite void of Events, the wind however was very fair and we now approachd the place where we were next to refresh ourselves apace. This morn the sun was immediately over our heads notwiths[t]anding which the Thermometer was no higher than Since we left the calms under the line the weather has grown cooler by gradual degrees, now we reckon it quite moderate after having felt the heat of 83 so lately.
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Combining optogenetic stimulation and fMRI to validate a multivariate dynamical systems model for estimating causal brain interactions. Irrespective of the technical approach used, rod phagocytosis exhibits a marked morning rise in activity in every species examined: eg. In rats, this daily rhythm is established by one month of age, and is irrespective of the lighting conditions during development [ ].
The timing and magnitude of cone shedding are more variable, since in many species [ , , — ] there is a distinct peak temporally removed from that of rods, occurring during the early night time. It has been suggested, based on these inversed rod and cone peaks, that phagocytic activity may be linked to PR physiology, rod shedding occurring at the beginning of the light period in which rod PR are less solicited, and cone shedding taking place during their resting period darkness [ ].
However, in several species rod and cone OS shedding occur simultaneously and maximally just after light onset [ , , ]. These differences cannot easily be explained by differences in behaviour, since both groups are diurnally active, but indicate that timing of phagocytosis is not simply related to visual activity. It is noteworthy that maximal rod phagosome numbers exceed by tenfold those of cones in mammalian species in which both have been compared [ , ].
This may reflect a different turnover rate of rods versus cones, since rod membranes may be renewed more rapidly than cones. Responses of phagocytosis to changes in ambient lighting have been widely studied. La Vail [ ] showed that a morning shedding peak was retained in rats placed in permanent darkness DD.
A subsequent study examined the effects of multiple lighting paradigms on the shedding response in albino rats [ ],and obtained similar resultsn DD, persisting even after two weeks of such conditions.
By contrast, the phagocytic response in Rana pipiens seems to be purely light driven, since it disappears in DD conditions [ ]. Effects of constant light LL have also been investigated, demonstrating that the shedding peak is abolished rapidly [ ].
In the Nile Rat, LL induced complete loss in rhythmicity of rod and cone PR phagocytosis, which instead showed a broadly uniform activity [ ]. In addition cone but not rod PR phagocytosis was strongly up-regulated, perhaps linked to the constant activation of the former by ambient light levels.
Indeed rod phagosome activity dropped to comparable levels to that of cones, fitting broadly with previous observations in rats [ ] which concluded that prolonged maintenance in LL abolishes the burst of shedding.
On the other hand, these findings contrast sharply with previous studies on nocturnal mice [ ] which showed no changes. Recent studies using mouse mutants have identified specific components of the circadian phagocytic pathway.
Knockout mice further develop age-related autofluorescent inclusions within the RPE, and exhibit age-related decline in visual responses. Subsequent studies highlighted the role of MFG-E8 as a candidate PR-bound ligand for this integrin, and revealed that mice with a gene deletion for this protein also showed uniform uptake profiles, although there was no age-related accumulation of lipofuscin [ ].
There are so far no data linking specific clock genes to retinal phagocytosis, or whether similar mechanisms underlie cone phagocytosis. The RCS rat is a commonly used animal model of inherited retinal degeneration. Known since 70 years, and well described nearly 50 years ago [ ], it was long ago realised through studies using chimeric animals derived from combinations of control and affected embryos that the disease state correlated with RPE rather than neural retina [ ].
Tissue culture studies demonstrated that RCS RPE were defective in OS internalisation, although they were still capable of binding normally [ , ]. But the difficulties associated with performing rat genome searches hindered identification of the mutation responsible.
This was finally achieved ten years ago, when the laboratory of Vollrath showed through positional cloning that the orphan tyrosine kinase receptor c-mer or mertk was a strong candidate [ ]. This was proven beyond doubt when the same group showed that replacement of a normal mertk gene into RCS RPE partially rescued the disease phenotype in vivo [ ]. Consistent with the degeneration observed in the rat and mouse [ ] models, mutations in the human MERTK gene are responsible for a sub-group of early-onset retinitis pigmentosa characterized by severe rod-cone dystrophy [ ].
Mertk mutations have also been linked with a rare retinal dystrophy [ ] and severe rod-cone dystrophy [ ]. The membrane renewal process engaged in by PR cells involves the adjacent cell monolayer — the RPE — in the disposal of spent OS membrane components.
The consequence of the phagocytotic load born by the RPE is apparent from the abundant lipofuscin Fig. For most non-dividing cell types, lipofuscin probably originates internally by autophagy [ — ]. Several bisretinoids of RPE lipofuscin have been identified; these compounds include the founding members of 3 families A2E, A2-DHP-PE and all- trans -retinal dimer various isomers and photo-oxidized forms and conjugated and unconjugated members in the case of the all- trans -retinal dimer series [ , — ].
It has been shown by a number of studies that the bisretinoid lipofuscin of RPE exerts adverse effects on RPE, acting both as a photosensitizer to damage cellular organelles [ , ] and exhibiting detergent properties which destroy membranes [ ].
In vivo , the regional lipofuscin content correlates directly with adjacent PR death [ ]. The lipofuscin that is amassed by RPE cells is the source of fundus autofluorescence, the natural autofluorescence [ ] that is imaged clinically by confocal scanning laser ophthalmoscopy cSLO [ 77 ] and fundus camera-based system [ ]. Given that the autofluorescent pigments of RPE lipofuscin are produced by reactions of all- trans -retinaldehyde it is not surprising that when the cis - and all- trans -retinal chromophores fail to be generated via the visual cycle, as in early-onset retinal dystrophy associated with mutations in RPE65, fundus autofluorescence is not present [ ].
While the accumulation of lipofuscin pigments is characteristic of normal RPE, this autofluorescent material is amassed in particular abundance in retinal disorders associated with mutations in the ABCA4 gene Fig.
These approaches include ABCA4 gene replacement [ , ] and systemic administration of compounds that limit the visual cycle [ — ]. There is also considerable interest in the role of RPE lipofuscin in age-related macular degeneration.
Complement activation that is insufficiently regulated is considered to underlie the susceptibility to age-related macular degeneration that occurs with certain genetic variants in complement factors [ 35 — 38 ] Section 4. In this regard it is notable that photooxidation products of A2E and all- trans -retinal dimer can activate complement while depletion of factor B reduces complement activation as does an inhibitor of C3 [ , ].
Complement activation triggered by photooxidation products of RPE bisretinoid lipofuscin could generate chronic inflammatory processes that gradually predispose the macula to disease. In mutant mice lacking the retinol dehydrogenase Rdh8, clearance of all-trans-retinal from OS following photobleaching, is significantly delayed and the mice exhibit several fold increases in A2E relative to wild-type mice [ ].
Another retinol dehydrogenase RDH12 is expressed in inner segments of PR where it may be involving in the reduction of not only all- trans -retinal but other damaging compounds [ , ]; a null mutation in this gene also results in elevated levels of A2E [ ]. In humans, mutations in RDH12, cause a severe form of autosomal recessive retinal degeneration characterized by early-onset and progressive rod and cone degeneration [ , ].
Another juvenile-onset form of maculopathy with dominant inheritance is associated with enhanced RPE lipofuscin accumulation [ — ]. The ELOVL4 protein serves to synthesize C28 and C30 saturated fatty acids and in the synthesis of CC38 very long chain polyunsaturated fatty acids, the latter being abundant in retina [ ].
In the healthy developed eye, RPE do not undergo mitosis; however, under some conditions, including complicated cases of retinal detachment [ ] and retinitis pigmentosa [ ], RPE cells both proliferate and migrate. When the the neural retina detaches from RPE, the latter cells can be dispersed into the vitreous cavity where they undergo a epithelium-to-mesenchyme transition to myofibroblasts that proliferate on the surface of the retina and via their contractile properties can re-detach a repaired retinal detachment [ , ].
Known as proliferative vitreoretinopathy, this condition is a result of the phenotypic regression of the cells that follows from a loss of junctional complexes [ 45 ]. Mutations in several genes expressed in RPE lead to retinal degeneration. It is thus significant that RPE cells are also an attractive target for gene therapy in part because the cells are readily accessible.
Another rapidly growing area of research involves cell-based therapies aimed at the replacement of defective RPE. Success in this area could be curative for conditions such as AMD. RPE cell proliferation in the form of PVR is the major cause of failure of retinal detachment surgery and yet this condition remains difficult to treat. Improved understanding of RPE biology may be key here.
Curr Mol Med. Author manuscript; available in PMC Aug 4. Hicks , 2 and C. Hamel 3. Author information Copyright and License information Disclaimer. Copyright notice. The publisher's final edited version of this article is available free at Curr Mol Med. Abstract Retinal pigment epithelial cells RPE constitute a simple layer of cuboidal cells that are strategically situated behind the photoreceptor PR cells.
Keywords: Retinal pigment epithelium, visual cycle, phagocytosis, melanin, retinal degeneration, transport, lipofuscin, retinoid. Open in a separate window. RPE Cells on the Frontline Poised at the interface of choroid and neural retina, RPE cells are responsible for the trafficking of nutrients and metabolic endproducts and serve to maintain ion balance in the subretinal space [ 57 ]. RPE Contributions to the DC ERG and the EOG As a consequence of the polarized distribution of ion channels and transport proteins on the apical and basolateral membranes of the cell [ 67 , 68 ], the RPE generates a spontaneous transepithelial potential TEP 5—15 mV that is primarily responsible for the cornea-positive potential difference that can be recorded across the eye in the dark; this voltage difference increases with a light stimulus [ 63 , 67 , 68 ].
The visual cycle Schematic of the visual cycle in rod photoreceptor outer segments and retinal pigment epithelium. Molecular Actors in the Rod Visual Cycle After rhodopsin decay, all- trans retinal is released into the lumen of the rod outer segment disc where it can combine with membrane phosphatidylethanolamine PE to form N-retinylidene-phosphatidylethanolamine N- ret -PE.
Cone Visual Cycle There is another visual cycle which does not go through RPE but rather cycles retinoids through cones and Muller cells [ ].
Genetic Diseases Associated with RPE Proteins of the Visual Cycle Impairment of some RPE genes involved in the visual cycle leads to a spectrum of conditions ranging from stationary visual defects to very severe forms of progressive retinal dystrophies.
Fundi of patients with various conditions related to cycle visual genes A, B : Left eye of a patient with Leber congenital amaurosis caused by mutations in RPE65 at 13 A and 22 B years of age. Cellular and Molecular Aspects of Phagocytosis Membrane-Bound Receptors The early studies by Hall and colleagues suggested that PR phagocytosis involved a ligand-receptor mediated interaction, since in vitro analyses demonstrated that OS binding and internalisation by RPE monolayers were temperature dependent, saturable and specific [ , ].
Potential Ligands The TAM family members were originally classified as orphan receptors since their natural ligands were not known [ ]. Intracellular Pathways Activation of different membrane-bound receptors triggers a variety of downstream signalling pathways, which play either direct or indirect roles in the phagocytic process Fig. Physiological and Integrated Aspects Compared to the important advances in identification of molecular components of the phagocytic process, less progress has been made in recent years concerning the general environmental and physiological control of PR phagocytosis, and the integration of the phagocytic process within the overall retinal metabolism.
RPE Lipofuscin in Retinal Disorders and Disease Models The lipofuscin that is amassed by RPE cells is the source of fundus autofluorescence, the natural autofluorescence [ ] that is imaged clinically by confocal scanning laser ophthalmoscopy cSLO [ 77 ] and fundus camera-based system [ ]. Bok D, Hall MO. The role of the pigment epithelium in the etiology of inherited retinal dystrophy in the rat. J Cell Biol. The relationship of primate foveal cones to the pigment epithelium.
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