General 
					 
					In the ponds and 
					swamps, one may find floating, rootless plants with 
					hundreds of tiny, balloon-like sacs attached to their green 
					branching stems. They are bladderworts, yet another kind of 
					carnivorous plants. These sacs in the water are 
					sophisticated miniature traps designed to provide animal 
					meals for these rootless floaters.  There are about 214 species of 
					bladderworts worldwide, occurring in
					practically every part of 
					the globe except in arid regions and 
					Antarctica. Some are terrestrial species found in 
					moist-to-wet, often acid soils, and in sphagnum moss 
					?? , while 
					others are aquatic, preferring to be floating freely in 
					quiet waters in bogs and lakes. Many terrestrial species 
					in the tropics are epiphytic. Some species exhibit 
					an intermediate lifestyle, capable of adapting to either 
					terrestrial or semi-aquatic habitats depending on the amount 
					of rainfall of the season. Of all the bladderworts in the 
					world, terrestrial species, by far, are
					the majority. (about 80%) Thirty-some 
					species of bladderworts are found in the 
					continental U.S.The bladderworts are perennial or 
					annual, consisting of long branching stems bearing numerous, 
					tiny, balloon-like sacs -- or bladders -- for which the 
					common name was given. The genus name Utricularia 
					is derived from the Latin word utriculus also 
					referring to a "small  bag". 
					Morphologically, 
					the arrangement of the vegetative organs of Utricularia is 
					quite peculiar in view of the general norm of the  
					flowering plants. Following Taylor (1989), the general 
					vegetative morphology of Utricularia is described as 
					follows:  
					Utricularia 
					does not have roots but in many (terrestrial) species 
					structures that resemble and function as roots exist. These 
					organs are termed rhizoids (Taylor 1989). A vertical
					stem is present only in a few species. In the 
					majority of 
					species, horizontal shoots called stolons form the 
					framework of the vegetative part of the plant (Taylor). Some 
					aquatic species produce air shoots, arising from the 
					stolons. The true function of this structure is not well 
					understood.  
					Utricularia has the green, 
					flat, sometimes feather-like organs that perform 
					photosynthesis. There are some disputes as to what to call 
					them (whether these 
					organs should properly be called leaves). "It seems reasonable to me", 
					Taylor (1989) writes, "that plants which have, in 
					defiance of the general rule, no radicle or roots and no 
					true cotyledons, may be allowed to have leaves which also 
					disobey the rules... "  
					In some 
					aquatic species, a dense cluster of leaves is formed at the 
					growth point of the stolon at the end of the growing season. 
					This is (called) a turion, a term for hiberncula (winter buds) 
					often used for aquatic species. Turions are resistant to low 
					temperature and desiccation.
					 
					The traps of all 
					known carnivorous plants are considered (to be) a modified leaf, 
					morphologically speaking, with Utricularia being the 
					only exception. 
					Positioning of the 
					traps in Utricularia is highly diverse in 
					relation to the other vegetative organs of the plant, and this 
					offers significant taxonomic characters (Taylor). 
					 
					In some species the 
					traps arise only from the stem or the peduncle base, while 
					in some others the trap appears at the tip of the leaf. In 
					terrestrial species, the traps may arise on leaves, stolons, 
					and also on rhizoids. In many aquatic species, the traps 
					are formed laterally on the leaves or leaf segments (Taylor).  
					 
					Inflorescence  
					Inflorescence is 
					always racemorse (Taylor 1989), meaning the flower scape..... 
					Corolla is always 
					two-lipped. (bilabiate) without an obvious tube (unlike 
					Pinguicula). The upper lip is small er than the lower. The 
					lower lip of the corolla always terminates in a spur, as in 
					Pinguicula. The expanded part (limb) of the lower lip has 
					(at its base) a palate. The front end of the lower lip ??? 
					is either entire or lobed, depending on the species. 
					The calyx of 
					Utricularia is always two-lobed, except in subgenus 
					Polypompholyx which has a four-lobed calyx. The calyx lobes, 
					in many cases, provide useful taxonomic characters in 
					species where the corolla morphology is often variable 
					(Taylor). In cases where corolla shapes have a high degree 
					of variation, the calyx lobes are, of all the floral organs, 
					probably the most useful in providing taxonomic character, 
					according to Taylor (1989).  
					Many species 
					produce a reduced-size flower that self-pollinates without 
					opening. This form of flower is called cleistogamous. In 
					some species, cleistogamous flowers coexist with normal, 
					open, chasmogamous flowers. In a terrestrial U.S. species,
					Utricularia subulata, cleistogamy is quite common. In an 
					aquatic species Utricularia geminiscapa, a short-stemmed 
					cleistogamous flower is produced underwater along with an 
					open, chasmogamous flower that protrudes above the water 
					surface.     
					The bladder 
					interior ?? contains two kinds of glands, bifids (two-armed 
					glands) and quadrifids (four-armed glands).... the relative 
					disposition of the arms of the quadrifids provides 
					diagnostic taxonomic character...  
					  
					
					
					 The bladderworts present a rather 
					unique morphology. First of all, the plants are entirely 
					rootless -- completely giving up the normal plant way of 
					obtaining nutrients from the root system. Also, the 
					distinction between stem and leaf is often vague, especially 
					in the aquatic species. The trapping mechanism, the bladder, 
					is a modified leaf or a leaf division morphologically, in 
					general conformity with all trapping structures found in 
					carnivorous plants of other genera. The inner surface of the 
					trap (bladder) corresponds to the upper surface of the leaf 
					division that it represents. ??????/ 
					The terrestrial species extends its 
					white stems in the damp soils from which arise green leaves 
					and slender flower stems above the ground. Numerous white 
					bladders are attached to the stem. In aquatic species, 
					branching stems and bladders are also greenish, indicating 
					photosynthetic in function, and the leaves are often 
					feathery and thread-like. During the growing season, aquatic 
					species float near the surface of the still waters with only 
					the flower scapes protruding above the water surface. 
					
					Inflorescence 
					
					
					
					 The flowers are generally quite 
					colorful and showy for both terrestrial and aquatic species, 
					especially when seen in masses. During the flowering, which 
					occurs from spring to late summer in the U.S. species, one 
					often finds the ponds covered with bright yellow or purple 
					corollas. This seems to be the only time these small , 
					obscure plants choose to announce their existence to the 
					rest of the world. Yellow is the most prevalent flower color 
					for this genus though white to purple or bluish flowers are 
					also common often with yellow or reddish markings. 
					
					
					 Although most of the U.S. species are 
					relatively small  and less noticeable than other 
					carnivorous plant species, sometimes even in flower, some 
					South American terrestrial species are massive in size, with 
					leaves reaching one foot (1 m??) or more in length. 
					Their 
					flower scapes may attain the height of 1m. Some of these 
					flowers compete with those of orchids in their beauty 
					and elegance. 
					
					  
					
					
					  
					
					Suction Trap 
					
					The traps range in size from 5 mm at the largest end to a 
					microscopic 0.3 mm. These sacs are highly sophisticated 
					mechanical traps capable of 
					catching tiny water animals with amazing efficacy. 
					The traps come complete with 
					a self-resetting mechanism  The 
					typical prey for these miniature traps includes insect 
					larvae (esp. those of mosquitoes), aquatic worms, water 
					ticks, and other tiny swimmers sharing the same habitat. 
					
					
					 The basic structure and function of 
					the traps are common for all species of bladderworts, 
					though there are some clear differences 
					between 
					terrestrial and aquatic species. Each trap has some antenna-like 
					hairs on one side of the trap opposite the attaching stem. 
					These hairs are non-irritable (non-sensitive) and are 
					considered ornamental in nature, contributing to attract 
					tiny animal prey to the trap entrance located just below the 
					base of the hairs. The hairs may also serve to protect the 
					entrance from flowing debris in the water. The lower half of 
					the entrance is a semi-circular valve -- or a door -- 
					hinged at the upper semi-circular arc, with the free edge of 
					the door tightly sealed in contact with 
					the firm collar of the 
					lower opening of the entrance, called threshold. The door 
					opens only inwardly. When the trap is set, with the door 
					sealed watertight, the pressure inside the trap is kept 
					lower than the outside. This happens because the water is 
					constantly being pumped out of the trap interior by glands 
					scattered all over the trap wall. Because of this pressure 
					differential, when the trap is viewed from above, the walls 
					are warped inwards and appear concave. 
					On the lower part of the outer surface 
					of the door grow tiny, stiff hairs. 
					
					 These hairs function as 
					a trigger lever. When a small  water animal, probably seeking 
					a shelter in the bladderwort jungle or perhaps lured by 
					nectar secretion, touches one of these levers, a delicate 
					mechanical latch of the door is broken. Giving in 
					to the outside water pressure, the door
					swings open inwardly, causing 
					the water animal to be sucked into the trap along with a bladderful of rushing water. The elastic trap bulges with an 
					in-rush of water, with the side walls of the trap popping up 
					in a normal convex shape. The door swings shut in a fraction of a 
					second, closing the entrance once again. All this happens in 
					an astonishing 1/30 to 1/40 of a 
					second. Once trapped inside, there is no hope left for the 
					prey. 
					Over a period of thirty minutes to an 
					hour, the trap mechanism is automatically set 
					again in 
					preparation for the next catch. This resetting is the result 
					of continuous pumping of water out of the trap interior.  
					
					
					Water Pumping 
					Mechanism 
					On 
					the inner surface of the trap are found numerous glands with 
					four projections known as quadrifids. 
					
					 These quadrifids are 
					believed to be responsible for absorbing water from 
					the trap 
					interior. Tiny spherical glands seen on the outer surface of 
					the trap are known to engage in active transport of ions 
					from the trap interior. The osmotic pressure (gradient) 
					built up by the ion movement generates the outward flow of 
					water from the trap interior. 
					Digestion 
					During a period of several days,  
					trapped animals are digested and absorbed by quadrifids, 
					and the nutrients are carried away to the rest of the plant. The digestive 
					enzymes, believed to be secreted from the quadrifids, 
					have been 
					detected inside the trap -- at least in younger traps. 
					After the first prey is captured, the bacterial actions are 
					seen to dominate in the digestive process
					and the trap often appears dark purple 
					in color. In an animal-rich environment, it is 
					not unusual for each trap 
					to capture several water 
					animals, filling the tiny trap completely.  
					
					Mechanical Delicacy 
					
					As we have seen, the water tightness 
					is essential for the proper function of 
					the suction-trap mechanism of 
					the bladderworts. Let us take a look at the mechanical 
					subtlety of this structure. Referring to this elaborate 
					trap structure, F. E. Lloyd, who contributed 
					immensely to our present knowledge of the
					trap mechanism, 
					comments, " ... But most to be wondered at are the traps 
					which present an astounding degree of mechanical delicacy 
					depending on a fineness of structure scarcely equaled 
					elsewhere in the plant kingdom."   
					
					
					 The surface of the threshold -- 
					against which the door edge rests -- is covered with a 
					pavement epithelium of sessile glands secreting mucilage. 
					There is a slight depression on the middle of the pavement 
					where the cells are most densely packed. The middle of the 
					free edge of the door -- which is strengthened with by dense 
					cells to make a firm edge -- rests in the pavement 
					depression. A slight change in 
					the door posture when the trap 
					becomes fully set is reflected in the trigger lever position 
					which is now more erect. Note that only the center of the 
					door edge impinges tightly on the pavement depression, with 
					the other portions of the free edge of the door merely lying 
					flat against the pavement, leaving chinks through which 
					the water can enter. This water leakage is prevented by the cuticular membrane attached to the outer edge of the 
					pavement, running completely across the threshold. This thin 
					but firm membranous tissue is called velum, which serves as 
					the second valve of the trap entrance, covering the outer 
					edge of the free margin of the door. That is, when the door 
					swings back right after springing the trap, it pushes 
					against the velum. Mucilage secreted from the stalked glands 
					on the threshold near the door rest also helps  
					seal the door further to prevent water leakage. This keeps the door 
					water-tight against the increasing outside pressure as the 
					water is continuously being pumped out from within the trap. 
					As long as this delicate mechanical balance of the door 
					latch is undisturbed, the door remains sealed in spite of 
					the mounting pressure outside.  
					Suppose a tiny water animal touches the tip 
					of a trigger lever. 
					By the lever action,
					the door edge 
					is pulled out from the pavement depression, thus unlatching 
					the door lock. Giving in to the outside water pressure, the 
					door flaps open inwardly. As expected from its mechanical 
					structure, the downward push of the lever is most effective in 
					triggering the trap.  
					Besides pumping of water by 
					the wall 
					glands -- which is physiological in nature -- setting, 
					tripping, and resetting of the trap are purely mechanical 
					phenomena, unrelated to growth movement. Therefore, one 
					bladder can repeat the trapping action without any 
					biological growth limitation. Some observer counted 14 times 
					of resetting of the bladderwort trap and this is not the 
					limit.   |