Chapter XII: Section II: Algæ (3)

Some genera, as for example Dasya, have slender, often elegantly branched threads, while such genera as Amansia and Odonthalia have instead a flat and pinnatifid frond. The latter, which has a very conspicuous cellular reticulation, is a genus of high latitudes, but is common on some parts of the Scotch and North American coasts. The British seas are rich in many genera of this order, and analogous forms occur in the southern hemisphere, where there are at least twenty-three genera. Many are remarkable for their singularity of structure: the Claudea for example, which is one of the most elegant of the Algæ, has a cancellated frond and is the ornament of warm seas; the Amansia and Leveillea which are distinguished by the beautiful reticulation of their fronds caused by large hexagonal cells; and the Dictyurus, in which the net forms a spiral web round the principal stem. Fig. 27 shows a portion of the network of Dictyurus purpurascens magnified. All the genera of this order possess free areolate hollow conceptacles perforated above, and containing nuclei, from the base of which short tufts of threads arise, each bearing a large obovate spore at its apex. The tetraspores are arranged in series either within the frond, or in distinct pod-like receptacles called stichidia. Fig. 28 shows the Polyzonia cuneifolia with its tetraspores arranged in rows in their pod-like stichidia, together with the areolated conceptacle and spores, all highly magnified. The antheridia differ in form in the different genera. In the Dasya they assume that of pods full of cells, in which the motile particles are generated; in the Rytiphlæa tinctoria the antheridia resemble those of the Dasya except in being elliptical, and in the Rytiphlæa pinastroides they are cellular bodies, without any investing membranes, clothed with delicate hairs.

The form of the Rhodosperms, as well as the limits of the species, like those of other Algæ, are affected by many circumstances known and unknown, such as the depth, temperature, saltness, and currents in the water. The Gelidium corneum varies to such an extent that its forms may not only be considered as distinct species, but even as belonging to different genera. The Delesseria alata is sometimes destitute of its margin, and then its midribs alone being left, it has the form of the Delesseria angustissima. Several species of the florid Algæ, which in their natural state have the tips of their fronds even and straight, occasionally produce hooked and clasping tips.

Brackish water is often a cause of change. The Irish moss, Chondrus crispus, when exposed to the fresh water of an estuary acquires great breadth and thickness, while at low water mark it is thin and has narrow forked branches, and there are many intermediate forms. The fruit rarely varies with these changes; its disposition and intimate structure, as well as that of the frond, are the points of prime importance for the determination of genera and species in the Algæ.

The MELANOSPERMEÆ, or Melanosperms, are olive-green Algæ, sometimes inclining to brown. They have fewer species than the Rhodosperms, but the individuals exceed in abundance and in magnitude all the other Algæ.

These large Melanospermous Algæ, which form marine forests in both hemispheres, are excessively strong and tough on the exterior but of a looser texture within, so that the cells of their tissue are of different sizes and forms, according to the degree of pressure. The stems and branches are more dense than the leaves. This highest order, however, has small and delicate Algæ united to the largest by many intermediate forms. The Melanosperms are either monœcious or diœcious, and bear their olive-green spores in cases, that is cysts, variously disposed on the plants. Many have two kinds of zoospores differing in nothing but size; they are produced in different organs; in some species both are fertile, in others only one, and, in these cases, the other is therefore supposed to be a fertilizing body, but however that may be, there are certainly antherozoids in this group of Algæ, especially in the order Fucaceæ.

The Ectocarpeæ have many representatives on our coasts, all of which are tufts of articulated threads from one to eighteen inches long, branched or simple. They are generally soft, some so flaccid that they cling together, but sometimes they are firm and stiff. The cysts which are attached to these threads have various forms; they are spherical, siliquose (that is, like long pods), or of other shapes, according to the species; but whatever form they may assume, they are filled with a dense endochrome. Besides these they have active granules contained in other distinct organs. M. Thuret has decided beyond a doubt that the latter are small zoospores, and it is presumed that the endochrome in the cysts is resolved into zoospores, but of a different order, as in the Ulvas. These two organs are for the most part situated on different individuals; in Ectocarpus pusillus (fig. 29 _b_) they are on the same. The different forms of fruit carpels are represented magnified in fig. 29.

The Ectocarpeæ contain little or no gelatine, whereas the genera of the group Chordariæ have soft gelatinous fronds of many forms, either incrustations, convex lumps, or tubers, like the Leathesia so common on our coasts; small plants as the Mesogloias, which have soft slippery filiform stems beset with myriads of moniliferous worm-like branches; or lastly the Chorda filum, a simple unbranched slimy cylindrical cord, varying from a quarter of an inch to the thickness of a pencil, and from one to twenty or even forty feet in length in deep water. The cord is tubular, divided into chambers by transverse partitions, formed of interlaced vertical and horizontal articulated threads. It tapers at each extremity, and the exterior, which is brown, is clothed with pellucid hairs. Vertical spores are immersed throughout the whole surface of the cord, and Dr. Harvey says that, mixed with these, there are numerous narrow, elliptical, transversely striated cells, which according to M. Thuret produce zoospores. Each plant rises solitary from its own little disc, but as the Chorda filum is a social plant, vast assemblies of it cover extensive areas of sand and mud, and form dense thickets in our northern seas. There are bands of it in the North Sea 15 to 20 miles long, and more than 600 feet wide; there is a submarine forest of it in Skapta Bay, Orkney; and in passing through the sounds of the western islands, as between Kerrera and the mainland, there are others. The long cords always lean in the direction of the tide, and must oscillate between two zones of rest, one at the turn of the flood, and another at the turn of the ebb. When dried the people use them for fishing lines. In the Chordaria divaricata both kinds of spore cysts are external, and give rise to zoospores.

In the preceding divisions of the Melanosperms the fronds consist of articulated threads; in the succeeding divisions the fronds are inarticulate. The latter comprise four very remarkable groups, of which the Dictyoteæ are distinguished by a leathery or membranous frond, sometimes cylindrical, but mostly flat, the surface of which is reticulated and sprinkled with groups or little patches of naked spores or cysts. The endochrome in the cysts is sometimes quadripartite, or even divided into eight parts. In one of the genera only, anything like antheridia have been found. The zoospores produced from the quadripartite endochrome are large, of a dark colour, and have two lateral cilia, while the bodies in the filiform much divided antheridia seated variously in the tufted threads are far more minute and pale, but with similar cilia. This order obtains its maximum of development in the tropical and subtropical regions; several species are found in the Mediterranean, while a few occur on our coasts, and on those of North America.[43]

The genus Dictyota begins the zonarioid group, whose structure is very curious. Every band (lacinia) of the frond terminates in a single cell, by the constant division of which at the lower side, the other cells of the frond are formed, the terminal cell of the frond being thus continually pushed onwards. Hence it results that the longitudinal lines of superficial cells converge, thus affording a ready method of ascertaining the genus in default of fructification. When a new centre of growth is to be made, that is, when the frond is to become forked, the terminal cell divides longitudinally and then each half-cell grows according to its own law. Fig. 30 shows the tip of the frond of the Dictyota dichotoma magnified; the cells on its surface are square, and the interior of each has a spiral structure.

The Padina Pavonia, or Peacock’s-tail laver of our southern coast, and those of North America and the Mediterranean, is sometimes included in the genus Zonaria. The species is remarkable for its wedge-shaped fronds, which are olive green shaded with rust colour, and, when in fruit, they are striped across with dark concentric zones, which are merely lines of spores immersed in the frond and seen through its transparent superficial membrane. Each zone is ornamented with a fringe of orange-coloured hairs. Parallel to, or rather concentric with, the spores, is a row of articulated threads, which bear so strong a resemblance to the antheridia of the Cutleria that a similarity of function is suspected by Mr. Berkeley. Species of Zonaria, Padina, and Haliseris, which is the most highly developed of the Dictyoteæ, are most abundant in tropical and low latitudes.

The Cutleria multifida is a small plant not exceeding eight inches in length, of an olive green varied with rusty tints. The frond is a flat ribless expansion many times variously slit in the upper part. It is beautifully marked by prominent dot-like tufts of fructification scattered over both sides of the frond, and grows on rocks and shells in from four to fifteen fathoms water.[44]

The great Laminariæ form the principal part of those vast submarine forests which encircle the globe in the arctic and antarctic oceans. None of these gigantic Algæ are to be met with in low latitudes, but there are several smaller species. The Laminaria debilis of the Mediterranean is not more than five inches high, and we have some ribbon-shaped species also of small size. Besides, many small individuals of the large species grow on our coasts at low water mark or below it; but the largest individuals are only found at depths suited to their size, so that the great Laminaria, or tangle forests, extend from low water mark to a depth of fifteen fathoms.

The fronds of these Algæ are for the most part leathery and of a fibro-cellular consistence. The Laminaria bulbosa is the largest of our sea weeds. Mr. Berkeley says that individuals are sometimes found which are a sufficient load for a man to carry. A flat stem, often more than a foot long, rises with a twist from a round hollow bulb a foot in diameter, throwing out numerous stout fibrous roots below; the stem is bordered by a thin wavy membrane, whence these plants are commonly called sea furbelows. At the top of the stalk there is a broad leafy expansion cut into straps or segments, twelve or more feet long, and from one to two feet wide.

The Laminaria digitata, commonly called the great tangle, oar weed, or sea girdle, has a fibrous root, a stem six or more feet long, with a wide expansion at its top cut into very long narrow segments. The fronds of some Laminariæ are deciduous; the stem increases in size year by year, a new frond springing from the apex and replacing the old one, which at last separates from the point of junction with the new frond, to which it is attached till the latter has attained its natural form and dimensions.

The Laminaria saccharina, called the devil’s apron on our northern coasts, is of a greenish olive when young, brownish when old. It has a fibrous root, a stem several feet long, ending in a flat ribless ribbon-like expansion, always very much longer than the stem, and terminating in a point. The margin of the frond is even, but wavy or puckered.

‘The fruit of these three great Laminariæ is imbedded here and there in the surface of the frond, thickening it and forming cloudy patches.’[45] It consists of thick club-shaped perpendicular cells in which the endochrome is ultimately divided into four parts. This is certainly the case in the Laminaria bulbosa, and also in the Alaria Pylaii, a species of which latter genus, the Alaria esculenta of our own coasts, is a much esteemed British dulse.

Abundance of colossal Algæ are found in the North Pacific, about the Kurile and Aleutian Islands, and along the deeply indented and channel-furrowed northwestern coast of America. The Nereocystis Lutkeana forms dense forests in Norfolk Bay, and all about Sitka. Its stem resembles whipcord, and is sometimes 300 feet long. It is exceedingly slender at the top, where it terminates in an enormous air-bladder six or seven feet long, and about four feet and a half in diameter at its widest part, the lower extremity passing into the stem. This huge air-vessel, which is the usual seat of the sea otter, is crowned with a tuft of twin leaves mostly rising on five stalks. These leaves, which are membranous and lanceolate when young, and from one to two feet long and two inches broad at the centre, are only marked with a few faint nerves, but they ultimately split lengthwise, cover a large space, and attain a length of twenty-seven or thirty feet, or even more. The growth of the Nereocystis must be enormously rapid, since it is an annual, and must therefore develop its whole gigantic proportions in one summer.[46] Boats cannot pass through the floating masses of this plant, whose stem is used for fishing lines, and whose cylindrical air-vessel serves as a siphon for pumping water out of boats.

The Thalassiophyllum Clathrus is also an inhabitant of the Russian coast of North America. It is about six feet high, very bushy and branched, each branch bearing a broad leaf at its extremity which unfolds spirally, and by this gradual development produces the stem with its branches and lateral divisions. A spiral border wound round the stem indicates the growth of the frond, which presents a large convex bent lamina without nerves, or a leaf of which one-half is wanting. Numerous long narrow perforations, arranged in a radiating form, give it the appearance of a cut fan.

The Macrocystis pyrifera and the Laminaria radiata are the most remarkable of marine plants, for their gigantic size and the extent of their range. They are met with on the antarctic coasts two degrees nearer the pole than any other vegetable, except the Diatomaceæ. The stem of the Macrocystis is slender, smooth, round, and slimy, rising from a fibrous root, like other Laminariæ, and bearing at its tip a lanceolate or oblong lanceolate frond. This frond divides at the base; the fissures extend upwards so as to form two petioles, each of which swells into an oblong or pyriform air-vessel. Another fissure is formed in a similar way a little above, and so on, till a single frond may at the same time have eight or ten fissures, each of which will ultimately gain the common apex. The margins of the fissures are at first perfectly smooth, but they soon become ciliated like the outer edge. The continuity with the fibrous base is at last broken, and the divisions of the leaves going on indefinitely, the whole reaches the length of some hundred feet, forming enormous floating masses which are wafted by the waves hundreds of miles from their origin. Fructification only takes place in young plants; consequently in such as are still attached to their native rocks. Even in that youthful state, Mr. Darwin mentions that such is the buoyancy of this powerful weed, that there is scarcely a loose block of stone on the coasts of Cape Horn that is not buoyed up by it.[47] The Macrocystis is native on the shores of the Atlantic, from Cape Horn to 43° S. latitude; but on the Pacific coast, according to Dr. Hooker, it extends to the river San Francisco in California, and perhaps to Kamschatka. The plant is reproduced by pyriform cells, full of endochrome, in nearly parallel rows imbedded in the fronds.

The rocky coasts of the Falkland Islands are covered with a vast growth of the gigantic Macrocystis mixed with forests of the arborescent Lessonia, which forms large dichotomous trees with a stem from eight to ten feet high and a foot in diameter. The leaves are two or three feet long, drooping from the forked branches like weeping willows. In the Lessonia nigrescens the quadripartite endochrome, ultimately resolved into spores, is contained in thickened club-shaped cells springing vertically between the surfaces of the frond.

A transverse section of the stem of many of the larger sea weeds presents zones, formed period by period, corresponding with the development of the laminæ, roots, and branches. The stem of the Lessonia bears a strong analogy to that of dicotyledons in having rings of growth, though there is a great difference. As increase in Lessonia takes place by the constant division of a flat leaf, the basilar portion of which becomes the petiole and ultimately swells into a branch, the stems have always a more or less elliptical form, and their section exhibits an elliptical core. This form of the core is not however peculiar, but exists in other Algæ. It is probable that the Lessoniæ, although attaining so large a size, are really of rapid growth.[48]

The Ecklonia is essentially a southern genus, though one species ascends to Spain and the Canaries. The frond is pinnatifid, the segments arising from the evolution of marginal teeth. The stem of the Ecklonia buccinalis, which is three or four inches thick and strongly inflated above, exhibits rings of growth with an orbicular central pith.

The group of the Fucaceæ exhibits the highest structure of all the olive-green Algæ, and forms a large portion of the sea weeds on our coasts, but they abound more in individuals than in the number of genera and species. A few have cylindrical stems and branches swelling out at intervals into large oblong inflated air-vessels, which gives them buoyancy in the water. The rest have a flat, ribbon-like stem, and for the most part dichotomous branches with a decided midrib, but no air-vessels, because they chiefly grow at half-tide level, and are exposed twice every twenty-four hours. The most common of our fuci, the Fucus vesiculosus, or bladder-wrack, has a midrib with air-vessels, generally in pairs on each side of it, formed by the inflation of the frond; these vessels, however, are frequently wanting, for it is the most variable in form and most widely spread of the Fuci. The fructification of this group is contained in large clavate receptacles or expansions of an orange or greenish yellow colour situated at the extremities or borders of the branches.

MM. Thuret and Decaisne discovered, by microscopic investigation, that the fuci have a truly sexual fructification, consisting of male and female cells inclosed in these receptacles. In the common Fucus vesiculosus it was found that the male and female cells are either in different individuals, or in different conceptacles on the same individual; whilst in the Fucus platycarpus, both the male and female cells were found to be contained in a globular cavity enclosed in the flattened receptacles which grow at the extremities of the branches. The cavity is lined with jointed hair-like filaments formed of cells, some of which are so long as to project through a pore on the surface of the receptacle in a spreading brush (see fig. 31, where the whole is highly magnified). Towards maturity, the cells of some of these filaments assume an ovoid form; the white viscous, granular matter in their interior acquires an orange hue, and is divided into a multitude of hyaline particles, each having an orange spot and two cilia of unequal lengths, which enable these spermatozoids to swim with great vivacity in the water as soon as they are set free by the rupture of the cell in which they are inclosed. Besides these, dark olive-green female cells, of a large pyriform shape, are fixed to the walls of the same cavity by very short stems; their contents spontaneously divide into eight spore cells, never more; each contains a colourless viscous liquid, which is mixed with protein and yellow-green matter, and is inclosed in a double coat. ‘The coats are united at the base, and when the spores are ready for dispersion, the inner coat bursts through the apex of the outer one, dragging with it a portion of the latter in the form of a little peduncle. The immediate covering of the spores at length bursts, and they are set free.’[49] In Fucus serratus, vesiculatus, and nodosus, swarms of spermatozoids are produced, but M. Thuret has proved by experiment that they never come to anything of themselves, and the unfertilized spores perish.

When a fertilized spore begins to grow, it assumes a pear shape, and sends out from its narrow end filaments or footstalks containing solid yellow grains at their extremities, where a hook or claw is formed by which it fixes itself to rocks or stones. The spore then divides itself into four equal cells of a brown colour, and by the continued subdivision of these into four, the plant increases in size, and assumes a form corresponding to the genus and species of the spore. Dr. Carpenter mentions that in the Fucaceæ there is also a multiplication by zoospores. These bodies are produced within certain of the cells that form the superficial layer of the frond, and swim about freely for a time in the water after their emission, until they fix themselves and begin to grow; but these are merely gemmæ.

All the Fucaceæ are tough leathery plants. This is even characteristic of the genus Cystoseira, various species of which may be seen on our coasts at low water mark, or in the tide pools. They are little shrub-like and somewhat thorny plants, not more than three feet high, with a cylindrical stem and many branches, near the extremities of which there are inflated air-vessels, sometimes two or three together; in some species they are lower down. Long spiny conceptacles are situated at the tips of the branches, but the endochrome does not divide in the germ cells as it does in the Fuci, so that each cell produces but one spore.

‘Throughout all latitudes the two divisions of Fucaceæ—Fucoideæ and Cystoseireæ, form the prevailing marine vegetation to which the name of sea-weed is commonly applied, and the different genera so arrange themselves as to present, with a few exceptions, a most harmonious assemblage.’ ‘None of these approach the tropics; the Fucoideæ abound towards the poles, and there attain their greatest bulk, diminishing rapidly towards the equator, and ceasing some degrees from the line itself; while the immense genus Sargassum finds its maximum in lower latitudes and under the equator itself. In the opposite cold and frigid zones the waters are inhabited by certain genera of Fucoideæ, which are in a great measure representatives of one another.’[50] The huge D’Urvillæa and the Sarcophycus in the Antarctic Ocean represent the Himanthalia and Fucus proper in the north, and the Cystoseireæ and Halidrys of the northern seas are represented by the Blossevillea and Scytothalia in the southern.

The frond of the Himanthalia lorea is a knob about an inch high, somewhat like a small mushroom; by degrees the top of the knob sinks in, and the frond becomes cup-shaped. In the second year of growth it throws out from its centre strap-shaped receptacles from two to three feet long and the sixth of an inch wide; they are slimy, forked, and entirely covered with fruit. The true frond sometimes becomes hollow and swells into a bladder. This singular plant, which grows on our coasts, extends from Norway to Spain. In the D’Urvillæa, its representative in the southern hemisphere, the frond and receptacle are united, for the plant, which is of large dimensions, has dichotomous fronds ten feet long, and an inch or more in breadth. Their surface is ornamented with large cavities like a honeycomb, and the fruit imbedded within them consists of antheridia and club-shaped germ cells with four spores in each. These plants form a large portion of the wrack and also of the living Algæ which surround the Falkland Islands and Cape Horn; and they extend to Western Chili, where the poorer class make a sweet mucilaginous soup of them. The Sarcophycus potatorum, the only species of its order, is nearly allied to the D’Urvillæa by the structure of its fruit, and is so named from pieces of its frond being used to carry water. Many other olive-green Algæ are peculiar to the southern hemisphere—among them the Hormoseira, in which the frond, at first even and filiform, becomes inflated so as to produce moniliform chains of vesicles, parts of which are at length rough with the apertures of the conceptacles; this plant has bladder-like air-vessels formed by swollen parts of the frond, like many of our Fucaceæ.

Those genera which have distinct organs containing air, as the Sargassum, of which there are numerous species, are either of low latitudes or tropical, but are sometimes drifted by currents to the extra-tropical shores. The Sargassum vulgare, however, grows on the rocks in the Mediterranean. The whole plant is of a translucent reddish brown; the stem has alternate branches, bearing lanceolate serrated leaves with a midrib, and generally dotted with dark pores. The air-vessels are small translucent round balls about the size of a currant, borne on flat stalks in the axils of the branches, and the spores are in conceptacles borne on the branchlets just above the air-vessel. In one variety of this most variable plant, the Uva di mare, the main stem ends in a loose bunch of these little air-balls.

The Sargassum bacciferum is often found in the Mediterranean, but only as a wanderer drifted in from the Atlantic, where masses of it, like floating meadows, occupy an area west of the Azores equal in extent to that of France, which has never changed its position since the time of Columbus, on account of the surrounding currents. Fields of it cover the seas near the Bahama Islands, and another permanent area of Sargassum of great extent occurs in the South Pacific. The Sargassum bacciferum is of a pale translucent olive colour, having branched stems, with lanceolate, midribbed, and serrated leaves, destitute of pores, and little stalked air-balls in the axils of the branches. The same individual continually produces new branches and leaves, and thus multiplies its species, but it never produces fruit; consequently its habits exactly resemble those of the Macrocystis, and as that plant becomes detached and floats after fructification, it is supposed that the Sargassum bacciferum may grow on rocks at the bottom of the Atlantic, between the parallels of forty degrees north and south of the equator, and when detached after fructification that it is uniformly drifted to particular spots which never vary. ‘Multiplication is so rapid in the floating beds of the Sargassum and Macrocystis, as to render fruit needless; and even the common Fucus vesiculosus occurs in the Mediterranean under a peculiar form consisting entirely of specimens derived from sea borne weed carried in by the current which sets in towards the Mediterranean from the Atlantic.’[51]

Kelp, the ashes of sea weeds, is the commercial source of iodine. Algæ growing in deep water contain most of that substance; consequently the kelp made at Guernsey, consisting chiefly of the ashes of Laminaria digitata, is richer in iodine than that made elsewhere.

Marine vegetation varies both horizontally and vertically with the depth, and it seems to be a general law throughout the ocean, that the light of the sun and vegetation cease together. It consequently depends upon the power of the sun, and the transparency of the water; so that different kinds of sea weeds affect different depths, where the weight of the water, and the quantity of light and heat, suit them best. One great marine zone lies between high and low water marks, and varies in species with the nature of the coasts, but exhibits similar phænomena throughout the northern hemisphere. In the British seas this zone does not extend deeper than thirty fathoms, but it is divided into two distinct provinces, one to the south and another to the north. The former includes the southern and eastern coasts of England, the southern and western coasts of Ireland, and both the Channels; while the northern flora is confined to the Scottish seas, and the adjacent coasts of England and Ireland. The second British zone begins at low water mark, and extends below it to a depth of from seven to fifteen fathoms. It contains the great tangle sea weeds or marine forests mixed with fuci, and is the abode of a host of animals. A coral-like sea weed is the last plant of this zone and the lowest in these seas, where it does not extend below the depth of sixty fathoms; but in the Mediterranean it is found at seventy or eighty fathoms, and is the lowest plant in that sea. The same law prevails in the Bay of Biscay, where one set of sea weeds is never found lower than twenty feet below the surface, another only in the zone between five and thirty feet, another between fifteen and thirty-five feet. In these two last zones they are most numerous; at a greater depth the kinds continue to vary, but their numbers decrease. The distribution in the Ægean sea was found by Professor E. Forbes to be perfectly similar, only that the vegetation is different and extends to a greater depth in the Mediterranean than in more northern seas. He also observed that sea weeds growing near the surface are more limited in their distribution than those that grow lower down, and that with regard to vegetation, depth corresponds with latitude, as height does on land. Thus the flora at great depths in warm seas is represented by kindred forms in higher latitudes. There is every reason to believe that the same laws of distribution prevail not only throughout the ocean, but in every sea.