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The botanical term parthenocarpy refers to the development of the ovary of a flower into a fruit without fertilization. [The biological term parthenogenesis refers to the development of an egg without fertilization.] Fruits that develop parthenocarpically are typically seedless. Some seedless fruits come from sterile triploid plants, with three sets of chromosomes rather than two. The triploid seeds are obtained by crossing a fertile tetraploid (4n) plant with a diploid (2n) plant. When you buy seedless watermelon seeds, you get two kinds of seeds, one for the fertile diploid plant and one for the sterile triploid. The triploid seeds are larger, and both types of seeds are planted in the same vicinity. Male flowers of the diploid plant provide the pollen which pollinates (but does not fertilize) the sterile triploid plant. The act of pollination induces fruit development without fertilization, thus the triploid watermelon fruits develop parthenocarpically and are seedless. Most bananas purchased at your local supermarket came from sterile triploid hybrids. The fruits developed parthenocarpically and are seedless.
The cultivated banana is often listed in botanical references as Musa x paradisiaca (Musaceae), although it is actually a complex hybrid derived from two diploid Asian species, M. acuminata and M. balbisiana. Common cultivated bananas are usually triploid (3n) with three sets of chromosomes. [Note: The word "set" is defined here as one haploid set of chromosomes.] If A represents one haploid set of chromosomes from diploid M. acuminata (AA) and B represents one haploid set of chromosomes from diploid M. balbisiana (BB), then hybrid bananas have three sets of chromosomes represented by AAB, ABB or another 3-letter (triploid) combination of A's and B's. Like seedless watermelons and red grapes, bananas are sterile and do not produce mature seeds. [Sometimes you can find aborted ovules inside the fruit that appear like tiny black dots.] In the formation of gametes during normal meiosis, homologous chromosomes must pair up with each other during synapsis of prophase I. Like other odd polyploids (with 3 sets of chromosomes), bananas are sterile and seedless because one set of chromosomes (A or B) has no homologous set to pair up with during synapsis of meiosis. Therefore meiosis does not proceed normally, and viable gametes (sex cells) are not produced. Since banana fruits (technically berrylike ripened ovaries) develop without fertilization they are termed parthenocarpic. Without viable seeds, banana plants must be propagated vegetatively (asexually) by planting corms, pieces of corms or sucker sprouts.
Parthenocarpy can be induced by growth hormones such as gibberellic acid (GA3) in which the ovaries mature without fertilization. Grape cultivars such as 'Thompson Seedless' are treated with gibberellic acid to order to produce larger fruits with longer internodes. The bunches have wider spaces between the grapes and better air circulation, reducing their susceptibility to fungal diseases and rotting within the bunch. Contrary to some references, 'Thompson Seedless' grapes are not parthenocarpic because fertilization does occur, but the ovules fail to develop into seeds within the maturing fruit. In cultivated figs, parthenocarpy generally refers to the development of the ovaries of female flowers within the syconium into drupelets without fertilization. The syconium is the structure that you typically associate with an edible fig fruit; however, it is really a flask-shaped structure lined on the inside with numerous unisexual flowers. The actual botanical fruits (called drupelets) develop within the syconium. Since the entire syconium enlarges and ripens into a juicy, sweet morsel, it is often referred to as a fruit. The female flowers are pollinated by a tiny female fig wasp that enters the syconium through a pore called the ostiole. According to W.B. Storey (Advances in Fruit Breeding, 1975), there are 2 genetically determined forms of parthenocarpy: stimulative and vegetative. Stimulative parthenocarpy involves the insertion of the wasp's ovipositor down the stylar canal into the ovary of short style flowers. It can also be induced by blowing air into the syconium, or by spraying the syconium with a plant growth regulator. The mature drupelets may contain a wasp (if an egg was laid in the ovary) or it may be empty. Vegetative parthenocarpy involves the formation of drupelets without any external stimulation, and is responsible for the hollow drupelets inside common figs such as "black mission," "kadota," and "brown turkey." [Some authors use the term parthenocarpy to describe the ripening of seedless fig syconia on the tree without any pollination or fertilization.]
The mule is a hybrid between a female horse or mare (2n=64) and a male donkey or jackass (2n=62). Since the mare contributes 32 chromosomes in her egg and the jackass contributes 31 chromosomes in his sperm, the mule has a diploid number of 63. Male and female mules are typically sterile because the horse and donkey chromosomes differ in number and they are not homologous. Therefore, the horse and donkey chromosome doublets fail to properly pair up with each other during synapsis of meiosis I. In fact, one horse doublet lacks a donkey doublet to pair up with. By the way, if the mother is a donkey or jennyass and the father is a stallion, the resulting sterile hybrid is called a hinny. The mule is an unusual animal because it has an odd number of chromosomes (2n=63) that is not divisible by two. The haploid number of a mule is not 31.5 because you can't have half of a chromosome in the gametes. But there is another way to get an animal or plant with an odd number of chromosomes called aneuploidy. If the chromosome number in the sperm or egg is more or less than the normal number of chromosomes in a haploid set, the resulting offspring (called an aneuploid) may have a chromosome number that is not exactly diploid (2n). Trisomy (2n+1) occurs when an individual has an extra copy of a chromosome. Examples of trisomy in people are Down's syndrome (three #21 chromosomes) and several sex chromosome aneuploidies caused by extra X or Y chromosomes, including Klinefelter's Syndrome (XXY), Trisomy X Syndrome (XXX) and the XYY Male Syndrome (XYY). Monosomy (2n-1) is caused by an individual missing one chromosome. Turner's Syndrome is a human female who received only one X chromosome from one parent and no X or Y chromosome from the other parent. Usually the cause of aneuploidy is nondisjunction during meiosis I or meiosis II, in which the sperm or egg carries extra or fewer chromosomes. In animals, autosomal monosomies and trisomies (abnormal numbers of autosomes) are usually detrimental and often fatal. In duckweeds (Family Lemnaceae), Mr. Wolffia's favorite plant family, the number of chromosomes in one haploid (1n) set is 10; however, polyploidy is common in the family, including 3n=30, 4n=40, 5n=50, 6n=60, 7n=70, and 8n=80. There are also good examples of aneuploidy in the duckweeds Landoltia punctata, Lemna minuta and L. minor, including adult (sporophyte) individuals with 36, 42, 43 and 44 chromosomes. Aneuploid duckweeds often lack vigor and are sterile. Odd polyploids such as 3n, 5n and 7n are also sterile. Can you guess why?
According to John Roach (National Geographic News 16 May 2006), DNA evidence confirms a natural hybrid between a female polar bear and a male grizzly. Apparently, grizzly bears have migrated north into Canada's western Arctic, and occasionally enter the range of polar bears. The male hybrid's white fur was interspersed with brown patches. It also had long claws, a concave facial profile, and a humped back--all grizzly characteristics. Unfortunately, this unique hybrid bear was killed by a hunter who thought it was a polar bear when he pulled the trigger. The common name for this hybrid is "polargrizz." I have not substantiated this remarkable discovery in a peer-reviewd scientific journal.
The citrus family (Rutaceae) contains some of the world's most delicious fruits, including numerous hybrid crosses between species. The popular tangelo grown in San Diego County is a hybrid produced by crossing a grapefruit (C. x paradisi) with a tangerine (C. reticulata). The grapefruit is sometimes called a pomelo, and this explains the blending (portmanteau word) of tangerine and pomelo. Actually, the grapefruit is a hybrid produced by crossing the shaddock or pummelo (Citrus maxima) with a sweet orange (C. sinensis). The shaddock is a large, thick-skinned, tropical citrus fruit up to six inches (15 cm) in diameter that is occasionally sold in supermarkets.
According to Apples: A Catalog of International Varieties by Tom Burford, there are 17,000 varieties of apples! Most of the apples grown commercially are probably diploid (2n), although there are many triploid varieties. For example, 'Gravenstein' apples are triploid with a chromosome number of 51 (3n=51). They are produced by the union of a diploid egg (2n=34) and a haploid sperm (n=17). This is accomplished by crossing a tetraploid plant (4n=68) with an ordinary diploid plant (2n=34). Because the triploid (3n) varieties are sterile, they must be propagated by grafting, where the scions of choice cultivars are grafted to hardy, pest-resistant root stalks. Apples are mentioned throughout most of recorded human history. The generic name Malus is derived from the Latin word malus or bad, referring to Eve picking an apple in the Garden of Eden; however, some biblical scholars think the fig, and not the apple, was the forbidden fruit picked by Eve. One of the earliest records of any fruit eaten by people of the Middle East is the common fig (Ficus carica). Remnants of figs have been found in archeological excavations dating back to the Neolithic era, about 1000 years before Moses. The fig is also the first tree mentioned in the Bible in the story of Adam and Eve. There are some scholars who think the apricot is a more likely candidate because it was an abundant fruit (along with figs) in the ancient Palestine area. Other interesting tales about apples include Johnny Appleseed, William Tell, Sir Isaac Newton, and Apple Computers.
The rose family (Rosaceae) includes many economically-important fruit trees known as stone fruits in the genus Prunus. Each species has many vamed cultivated varieties (cultivars). Botanists have moved some of these species into separate genera, including Amygdalus (peach) and Armeniaca (apricot). Some examples of stone fruits are fuzzy-skinned peaches (P. persica syn. Amygdalus persica), smooth-skinned peaches called nectarines (another variety of P. persica), plums (P. domestica), apricots (P. armeniaca syn. Armeniaca vulgaris), and cherries (P. avium and P. cerasus). Like apples and pears, there are hundreds of cultivated varieties. These fruits are technically referred to as drupes because they consist of an outer skin or exocarp, a thick, fleshy middle layer or mesocarp, and a hard, woody layer (endocarp) surrounding the seed. The part of these fruits that is eaten by people is the mesocarp layer and also the exocarp if you don't bother to peel them. The woody endocarp layer protects the seed and probably aids in the dispersal of drupaceous fruits by hungry herbivores. In wild plants with drupes, the seeds can pass through the entire digestive system of grazing animals and be planted in new locations. The almond (Prunus amygdalus syn. Amygdalus communis) is also a drupe with a green exocarp and thin mesocarp surrounding the pit. When you crack open an almond to get the seed, you are actually cracking open the endocarp layer. Some species of Prunus have been artificially crossed to produce some unusual hybrids. The peachcot (Prunus persica x P. armeniaca) is a hybrid between the peach and apricot; the cherrycot (P. besseyi x P. armeniaca) is a hybrid between the cherry and apricot; the plumcot (P. domestica x P. armeniaca) is a hybrid between the plum and apricot. Some of these hybrids have many different named cultivars, depending on which varieties of stone fruits have been crossed together. In addition, hybrids often retain more characteristics of one parent and are given special names. For example, some cultivars of plumcots are called "pluots" because these resemble plums more than apricots. Plumcots called "apriums" resemble apricots more than plums.
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3. Some Generalizations About The Duckweed Family The duckweed family is well represented in western North America with nearly half of the world's species. The plant body of duckweeds is quite unlike other flowering plants because it does not have stems or leaves. It represents the ultimate in reduction of an entire vascular plant. The terms "frond" and "thallus" are sometimes used in the literature, but these terms are not appropriate because the plant body of duckweeds is not homologous to the fronds of ferns or the body of fungi and algae. Although the body of duckweeds does have paired guard cells and stomata on its upper surface and superficially resembles a leaf (particularly the flattened duckweeds Spirodela, Landoltia and Lemna), it is morphologically and embryonically completely different. In Spirodela, Landoltia and Lemna it is a flattened structure with slender, hairlike roots on the underside. Spirodela and Landoltia are unique among duckweeds because of a minute, membranous scalelike leaf (prophyllum) enveloping the dorsal and ventral surfaces of the basal end. In Spirodela polyrrhiza the prophyllum is visible on young plants (fugacious in older plants) and on overwintering turions. This basal portion and its connecting stalk correspond to a condensed shoot that has become greatly reduced through evolution. Landoltia has a reduced prophyllum that perishes in full grown plants. A prophyllum is lacking in Lemna, Wolffia and Wolffiella. The latter two genera have been reduced through evolution to minute, rootless spheres or flattened ribbons. Wolffia has a minute globose or ovoid body one mm long or less. In Wolffiella the thalluslike body is transparent and flattened, with the free ends often curved downward in the water.
Although all species of Lemna have a basal root sheath near the attachment node, two species in section Alatae (L. aequinoctialis and L. perpusilla) have a distinctive root sheath with 2 lateral wing-like appendages.
Elongated tracts of cells called nerves are present in Lemna, Landoltia and Spirodela. They originate at the node (point of root attachment) and extend through the plant body toward the distal (apical) region. A similar tract of elongated cells (called the costa) can be seen in the triagular budding pouch of Wolffiella. The position of the coasta in relation to the budding pouch is an important characteristic used to separate W. lingulata from W. oblonga. Tracts of elongated cells also extend through the center of the roots of Lemna, Landoltia and Spirodela. Nerves and tracts of elongated cells may serve to transport minerals and sugars, similar to the function of veins. In some species of Lemna, Landoltia and Spirodela, the elongated cells of nerves contain tracheids with ring-shaped or spiral-shaped thickenings in the walls (annular tracheids). These elongated cells are not called veins because the plant bodies of duckweeds are not homologous to leaves. 4. Cladogram Of The Duckweed Family Different genes within the nucleus and cytoplasmic organelles (chloroplast and mitochondria) can be used to construct phylogenetic trees called cladograms. One gene in the nucleolus codes for the smaller subunit of the ribosome. The gene is called SSU rDNA or small subunit ribosomal DNA. Base sequences from this gene are sometimes used to compare taxa at the species level. Chloroplast DNA, including the protein-coding rbcL gene, is often used at the family level to show the relationships between genera and species within the family. Introns are also used to construct family trees. Introns are sections of messenger RNA that are removed prior to translation at the ribosome. Most botanists consider the Lemnaceae to be closely related to the arum family (Araceae), and comparative chloroplast DNA studies have confirmed this taxonomic affinity (Duvall, et al. Annals of the Missouri Botanical Garden Vol. 80, 1993). In fact, several authorities have proposed some drastic and significant changes in the classification of many traditional angiosperm families, including the placement of all duckweeds in the Araceae rather than the Lemnaceae. [See: Angiosperm Phylogeny Group. 1998. "An Ordinal Classification For The Families Of Flowering Plants." Annals of the Missouri Botanical Garden 85: 531-553; Judd, W., C. Campbell, T. Kellogg and P. Stevens. 2002. Plant Systematics: A Phylogenetic Approach. Sinauer Associates, Inc., Sunderland, MA. Some of these proposed changes are summarized in an article by E. Dean in Fremontia 30 (2): 3-12, 2003. If accepted by the botanical community, the incorporation of these changes into botany textbooks, floras, checklists and herbarium collections will be a formidable task. Computer-generated evolutionary trees or cladograms have been used to show the taxonomic relationships of duckweed species within the family. The cladograms are based on thousands of data characters, including morphology, anatomy, flavonoids, allozymes, and DNA sequences from chloroplast genes and introns. The branch (clade) length and position in the tree correspond to the number of character differences between taxa. The characters are numerically weighted according to their evolutionary importance. For example, a root would have a higher value than a papule. Cladograms are generated multiple times, and they don't always come out the same. The term "bootstrapping" refers to a cladogram or phylogenetic tree that comes out the same way out of a total number of times. For example, one thousand cladogram "trees" are generated and the same pattern comes out 900 times. This cladogram would have a bootstrap value of 90 percent. The following cladogram shows all the five genera and 38 species within the duckweed family (Lemnaceae). It was generated from DNA sequences of rbcL genes from all known members of the the family using the computer program PAUP:
Because of their degree of reduction, Landolt (1986) considers the two diminutive genera Wolffia and Wolffiella to be the most recently evolved offshoots in the phylogeny of this family. Wolffia has the fewest shared characters with the presumed ancestral Spirodela and is placed farthest away in an evolutionary tree (cladogram). The new genus Landoltia is morphologically intermediate between Lemna and Spirodela. According to D.H. Les & D.J. Crawford (Novon 9: 530-533, 1999), it represents an isolated clade distinct from both Lemna and Spirodela. DNA comparisons of all members of the Lemnaceae by Les, et al. (Systematic Botany 27 (2): 221-240, 2002) indicate that all five genera represent distinct clades. With the exception of Landoltia and a few changes in sections, the 38 taxa recognized in the study by Les et al. (2002) are remarkably consistent with those recognized as morphologically distinct by Landolt. Duckweeds Now Placed In The Arum Family (Araceae)
Limnobium: Floating Aquatic Superficially Resembling Spirodela
5. Controversies Over The Genus Landoltia Many traditional phylogenetic groupings of species within families and genera are not monophyletic and are inconsistent with modern cladistical analyses based on DNA. In other words, the groupings are paraphyletic or polyphyletic, and do not show all species within a group descending from a common ancestor. Monophyly is the natural evolutionary pattern in which all species are descended from a common ancestor. In order to have consistent computer-generated, monophyletic cladograms, it is sometimes necessary to change paraphyletic and polyphyletic groupings by moving species into different genera, and by moving genera into different families. Many of the taxonomic revisions in the Jepson Manual 2nd Edition (2012) were made in order to have consistent monophyletic groupings. This is why Spirodela punctata was placed in the genus Landoltia and why the Lemnaceae was placed in the familiy Araceae.
In 1999, D.H. Les and D.J. Crawford proposed the new genus Landoltia containing one species L. punctata, formerly Spirodela punctata. This species is morphologically intermediate between Lemna and Spirodela. According to Les & Crawford, it represents an isolated clade distinct from both Lemna and Spirodela. Without this change, the genus Spirodela would be paraphyletc.
According to Professor Dr. Elias Landolt (personal communication, 2001), the creation of the new genus Landoltia is not necessary based on a purely morphological point of view; however, based on DNA and enzymatic studies, the change is warranted in order to form phylogenetically consistent taxa. The inclusion of a fifth genus Landoltia appears to be necessary based upon a comprehensive analysis of the Lemnaceae by D.H. Les, D.J. Crawford, E. Landolt, J.D. Gabel, and R.T. Kimball (2002). More that 4,700 characters were studied, including data from morphology and anatomy, flavonoids, allozymes, and DNA sequences from chloroplast genes (rbcL, matK) and introns (trnK, rpl16). The Angiosperm Phylogeny Group (APG) has proposed some significant changes in the classification of many traditional angiosperm families, including the placement of all duckweeds in the Araceae rather than the Lemnaceae. Nomenclatural changes are cited under the APG II system (2003) and superceeded by APG III system (2009). These changes are based on computer-generated evolutionary trees or cladograms. Thousands of data characters have been used, including morphology, anatomy, flavonoids, allozymes, and DNA sequences from chloroplast genes and introns. The Jepson Manual Second Edition (2012) essentially follows the changes summarized in the following reference by W.T. Judd, et al. 2008. Since the genus Landoltia was proposed by D.H. Les and D.J. Cawford in 1999, several classic papers on the phylogeny of the duckweed subfamily (Lemnoideae) and other aroids (Araceae) have used the name Landoltia. In my opinion, the name Landoltia is warranted because it is consistent with the objectives of the Jepson Manual 2nd Edition (2012) based on phylogenetic studies using plastid DNA.
Lemna punctata G.F.W. Meyer This was Meyer's original name based on the type specimen collected along the Essequibo River, Guyana, South America in 1818. Unfortunately, Meyer's original type specimen was lost. Spirodela punctata (G.F.W. Meyer) Thompson C.H. Thompson placed this species in the genus Spirodela in 1898. Since the type specimen was lost, he based the new name on a specimen from the 1938-1842 Wilkes Expedition, labeled Orange Harbor, Tierra del Fuego. According to Landolt (1986), Thompson neotypified this species in his 1898 publication. Landoltia punctata (G.F.W. Meyer) Les & D.J. Crawford In 1999, D.H. Les and D.J. Crawford placed this species in the genus Landoltia based on DNA evidence. Re-Neotypification Of G.F.W. Meyer's 1818 Type Specimen Of "Lemna punctata" Note: This is a complicated taxonomic subject involving many articles from the International Code of Nomenclature For Algae, Fungi, and Plants (Melbourne Code) 2011: Available on-line at: http://www.iapt-taxon.org/nomen/main.php. An argument for replacing the names Landoltia punctata and Spirodela punctata with the previous name Spirodela oligorrhiza has been made by Daniel B. Ward (2011). In order to make sure we are referring to the same species, Ward has suggested calling this "Lesser Greater Duckweed" to avoid confusing it with the larger species of Spirodela (S. polyrrhiza & S. intermedia) called "Greater Duckweeds." In this article I will simply call it LG Duckweed instead of Lesser Greater Duckweed. Ward's proposal involves the re-neotypification of G.F.W.Meyer's 1818 type specimen named Lemna punctata which was apparently lost. Ward also proposed as the new type a different species that we know today as Spirodela intermedia.
In July 2012, I received an e-mail message from Dr. Thomas Rosatti, editor of the revised Jepson Manual (2nd Edition), asking my opinion on Ward's retypification. Since C.H. Thompson already neotypified this species as Spirodela punctata in 1898, Ward's retypification should really be a "re-neotypification." Since I wrote the section on duckweeds (subfamily Lemnoideae), adopting Ward's re-neotypification would result in changes to several related species. In July 2012, I stated my opposition to Ward's proposal on my on-line Lemnoideae page on Wayne's Word. I also included a two-paragraph e-mail message from Dr. Elias Landolt, Zurich stating his opposition to the proposed re-neotypification (see below). This quotation can be verified on the Internet Archive Wayback Machine dated 8 September 2012. Spirodela punctata (Meyer) Thompson was named by C.H. Thompson in 1898 based on a collection from the 1938-1842 Wilkes Expedition, labeled Orange Harbor, Tierra del Fuego. Whether this collection actually came from the tip of South America is debatable. The parenthetical author G.F.W. Meyer described this species earlier as Lemna punctata from a type specimen collected in Guyana, South America in 1818. Unfortunately, Meyer's original type specimen was lost. According to Ward (2011), LG Duckweed does not occur in the areas where these collections were made: The Tierra del Fuego collection was mislabeled and the Guyana collection was not LG Duckweed. Futhermore, he states that the only native Spirodela in South America is S. intermedia. Since Meyer's type specimen was lost, Ward re-neotypified the species as Lemna punctata G.F.W. Meyer and he designated S. intermedia as the type. Thompson's binomial is still Spirodela punctata (Meyer) Thompson; however, this no longer refers to LG Duckweed. It is now the correct binomial for the South American Spirodela intermedia. The correct name for LG Duckweed now becomes Spirodela oligorrhiza (Kurz) Hegelmaier, a name published by Hegelmeier in 1868. Hegelmeier apparently never saw the South American specimens discussed above, so his name is probably based on the true LG Duckweed. Ward's 2011 neotypification will make Landoltia a synonym of Spirodela and no longer available for the intended LG Duckweed. The restoration of separate generic status for LG Duckweed now known as Spirodela oligorrhiza (Kurz) Hegelm. will require the creation of a new genus name. The binomial Spirodela punctata (Meyer) Thompson will now refer to the South American species known as Spirodela intermedia W. Koch. By neotypification the name Landoltia becomes a synonym of Spirodela intermedia. Quoted E-Mail Message From Dr. Elias Landolt According to E. Landolt (Personal Communication, 2012), the name change proposed by Ward is untenable. This quotation can be verified on the Internet Archive Wayback Machine dated 8 September 2012.
"I can understand that Thompson choose a new type for Lemna punctata. The correctness of his decision is not disputed. I checked the neotype the collection of Wilkes from copies in four different Herbara. It is clearly the species which is now called "punctata". It is not important if the material was collected in Orange Harbor or somewhere else. Because it is not possible and will probably never be possible to decide the identity of Lemna punctata with certainity it is not advisable to change the correctly published neotype of Thompson. If we change the type of L. punctata again we will have a terrible chaos in nomenclature. Therefore I am not following the proposal of Ward."
My objection to Ward's proposed neotypification is based on two primary points. (1) He is re-neotypifying Meyer's lost type specimen with the name Lemna punctata; however, he is using Spirodela intermedia as the type. It is impossible to know with 100% certainty which species Meyer was describing under the name Lemna punctata back in 1818. It could have been the "LG Duckweed" that we know as Landoltia punctata (Spirodela punctata = Spirodela oligorrhiza), or it could have been another species of Spirodela such as S. intermedia. Why complicate this taxonomy based on speculation. (2) Cladistical analysis has clearly shown that Spirodela punctata belongs in a separate genus (Landoltia), otherwise the grouping of Spirodela with 3 species is paraphyletic. The trend in modern floras such as the Jepson Manual Second Edition (2012) is for consistent monophyletic groupings. A Review Of Ward's Proposed Re-Neotypification Ward's re-neotypification of Lemna punctata has been reviewed by J.H. Wiersema of the USDA Agricultural Research Service, National Germplasm Resources Laboratory, Beltsville, Maryland.:
6. An Updated Key To The Duckweed Family
Depending on the genus, daughter plants are produced vegetatively in 2 lateral, flattened, budding pouches (Spirodela, Landoltia & Lemna), a flattened, triangular budding pouch at the basal end (Wolffiella), or a funnel-shaped budding pouch at the basal end (Wolffia). Each plant produces up to a dozen daughter plants during its lifetime of 1-2 (or more) months. The daughter plants repeat the budding history of their clonal parents, resulting in exponential growth. It has been estimated that the Indian Wolffia microscopica (Griff.) Kurz may reproduce by budding every 30 hours under optimal growing conditions. At the end of 4 months this would result in about 1 nonillion plants (1 followed by 30 zeros) occupying a total volume roughly equivalent to the planet earth. This astronomical vegetative growth and the ability of some species to grow in stagnant, polluted water is why some duckweeds are well suited for water reclamation. Some species not only thrive on manure-rich water, but can be fed back to livestock, thus completing the recycling process. In addition, some species (such as Wolffia) are a potential source of food for humans because they contain about 40 percent protein (dry weight) and are equivalent to soybeans in their amino acid content (with high levels of all essential amino acids except methionine). Although flowers are rarely observed in some species, all duckweeds bloom and reproduce sexually; however, some populations in small ponds may be clones of each other and not able to produce viable seeds. Since the flowers are typically protogynous with the stigma receptive before the anther is mature, the plants must be cross pollinated by genetically different individuals with mature pollen-bearing anthers in synchronization with the receptive stigmas. During the summer months, 2 stamens (androecium) and one pistil (gynoecium), all enclosed in a membranous saclike spathe, appear within budding pouches at the edge of the plant body in Spirodela, Landoltia and Lemna. In Wolffiella and Wolffia, a minute floral cavity develops on the upper side of the plant body containing a single stamen and pistil (not enclosed by a spathe). The tiny bisexual flowers have no sepals or petals, and are barely discernible without magnification. Because of the sweet (sugary) stigmatic secretions and spiny pollen grains (covered with minute protuberances), there is evidence that certain species may be pollinated by insects. In fact, Lemnaceae pollen has been detected on flies, aphids, mites, small spiders, and honey bees on the surface of dense duckweed layers. With floral sex organs projecting from the surface or lateral budding pouches, many duckweed species may be contact-pollinated as flowering individuals bump together or become piled up in windrows along the edges of ponds and lakes. 7. Identification Of Morphologically Similar Species Lemna minuta vs. L. valdiviana Since flowers and fruits are rarely observed, most taxonomic keys to the Lemnaceae are based on relatively few diagnostic vegetative characteristics that may vary under different environmental conditions. This often makes precise identification of some species difficult, or in some cases, practically impossible. All North American species have been separated by their flavonoid spot patterns using two-dimensional paper chromatography [see McClure & Alston (1966), Amer. J. Bot. 53: 849-860]. It should be noted that flavonoid chemistry is not always reliable for taxon distinction because chromatographic patterns may be influenced by environmental factors [see Ball, Beal & Flecker (1967), Brittonia 19: 273-279]. In addition, R. Scogin of RSA and J.L. Platt of OSU studied two-dimensional chromatography on clonal populations of Lemna minuta Kunth from San Diego County and came up with patterns identical with McClure & Alston's L. valdiviana Phil. According to Landolt (1987), the original clones of L. valdiviana studied by McClure & Alston may have actually been L. minuta. During the past century, the taxonomy of L. minuta Kunth has been complicated by different names used by different authors. Several of the synonyms commonly found in the literature include L. valdiviana var. minima Hegelm., L. minima Phil. ex Hegelm. and L. minuscula Herter. James L. Reveal (Taxon 19: 328-329, 1990) neotypified the oldest name L. minuta Kunth and cleared up some of the confusion and controversy about this widespread species.
Veins (Nerves) and Air Spaces
Note: Sometimes placing difficult species in an observation dish and examining them over several days can be helpful. Digital images can also bring out subtle differences. The following duckweeds were photographed through a dissecting microscope using a Sony digital camera with backlighting:
Using Dorsal Row Of Papules To Separate Lemna turionifera From L. minor Another difficult group of duckweeds is Lemna turionifera and L. minor. L. turionifera has three main veins and is superficially similar to L. minor and nongibbous L. gibba. It differs from L. minor and L. gibba in having a row of 3-7 minute papules along the midline of the dorsal surface. It also differs from L. minor by developing reddish anthocyanin on its underside, starting in the region around the root. What really sets this species apart from other duckweeds is the presence of rootless, overwintering turions in the fall months. These are referred to as "winter buds" in the Jepson Manual of California Plants (1996). L. turionifera appears to be more common than L. minor in San Diego County. It generally replaces L. gibba in the higher elevations. Unfortunately, reddish anthocyanin and turions are not always present, so you must rely on the row of papules along the midline of dorsal surface. This can be difficult to see, especially on dried herbarium specimens. Ideally, herbarium specimens should include field notes on the presence of a dorsal row of papules and reddish anthocyanin on the ventral surface. With some practice, these traits can be observed with a hand lens.
8. Importance Of Backlighting For Duckweed Identification When identifying duckweed species (especially Lemna, Landoltia and Spirodela), it is very important to view the plant bodies with backlighting (substage illumination) in order to see the number and the extent of the nerves. With a good 10x hand lens this can be accomplished by holding the plant body up against the bright sky. Backlighting is also crucial in order to see the tract of elongated cells (costa) in the budding pouch of Wolffiella. The position of the costa within the triangular budding pouch is very important in order to distinguish between W. lingulata and W. oblonga.
In these times of high technology, as botanical research moves toward a molecular emphasis, it is very important to have specimens verified by a taxonomist. It is also imperative to have carefully prepared voucher specimens on file in a nationally recognized herbarium. Modern molecular techniques, such as DNA sequencing, may lead to a better understanding of these fascinating species. 9. Photoperiodism In The Duckweed Family Although some duckweed species superficially resemble each other, they may have significantly different biochemical patterns, such as an entirely different photoperiodism in response to day length (hours of darkness). During the hours of daylight the protein leaf pigment called phytochrome 730 (P-730) is formed. During the hours of darkness P-730 is slowly converted into phytochrome 660 (P-660). In short-day plants P-730 inhibits flowering. Short-day plants typically need about 15 hours of darkness to convert all the P-730 present at sundown into P-660. In these plants, P-660 stimulates the release of the essential flower stimulant "florigen" which induces flowering. The P-660 pigment is very sensitive to specific wavelengths of light, and a flash of light during the 15 hours of darkness can instantaneously convert all the P-660 back into P-730. Lemna aequinoctialis is clearly a short-day plant because it requires 16 hours of darkness (8 hours of light) to flower. The closely related L. perpusilla is also a short-day species that exhibits maximum flowering with 13-18 hours of darkness, and no flowering with 9 hours of darkness (15 hours of light). These species will generally not bloom during the longest days of summer or in a pond next to a bright street light. Long-day plants require 15 hours of daylight and 9 hours of darkness in order to flower. In these plants P-730 stimulates the release of florigen and subsequent flowering. If the nights are long enough to convert all the P-730 into P-660, no florigen will be released and flowering will not occur. Lemna gibba is a long day plant that flowers with 9 hours of darkness. This species typically flowers during the longest days of summer. It will generally not flower with 12 hours of darkness, such as at the equator or during the vernal equinox, because the nights are too long. The physiology of these long-day and short-day species of duckweeds can definitely affect their range and potential for flowering and seed production. Exactly how some duckweed species are dispersed and how they survive intermittent streams and ponds that dry up during summer is an enigma. Being carried from pond to pond on the feet of water fowl (tucked neatly under the ducks' bodies during flight), probably explains the distribution of some species. In the southeastern United States there are records of wolffia plant bodies being carried by a tornado, and they have even been reported in the water of melted hailstones! Some species have been carried by rivers and streams, and in the shipment of fish and aquarium cultures. Professor Dr. Elias Landolt (1997) discusses some of the ways duckweeds survive dry conditions (Bulletin of the Geobotanical Institute ETH, Stiftung Rubel 63). Seeds of all Lemnaceae investigated so far tolerate drying for at least a few months to several years; however, seeds are rarely produced by clonal populations of some species. Although vegetative plant bodies are unable to withstand desiccation for more than a few hours, they may survive days (or weeks) embedded in wet mud and debris. According to Dan Richards (The Distributional Ecology Of Duckweeds (Lemnaceae) In Local Populations Of Northern California, MA Thesis, Humboldt State University, 1989), vegetative plants of two species survived up to six hours of desiccation (out of water). The two species tested by Richards (1989), Lemna minor and Landoltia punctata, had a much higher survival percentage when they were in large clumps compared to individually dried plants. Richard's experiments clearly show that these species could easily be carried short distances by migratory water fowl. Species that do not readily form seeds can also survive weeks or months of drought as turions, especially if the turions are imbedded in mud, silt and debris. This is especially true of the minute turions of Wolffia species. According to Landolt (1997), the South African Wolffia cylindracea may survive seasonally dry ponds for at least 16 months if the minute turions are firmly imbedded in clayey soil. 10. Axenic Culture Of Duckweeds In Nutrient Agar
11. Control of Duckweed Blooms In Ponds and Reservoirs One of the most common questions received at this site is how to control population explosions or "blooms" of duckweeds in which ponds, lakes and reservoirs become covered with a thick green layer of Lemna, Spirodela, Landoltia and Wolffia. Lemnaceae blooms typically occur in waters rich in nutrients, especially phosphorus and/or nitrogen. The nutrients originate from pollution from excessive use of fertilizers or possibly by an imbalance in the populations of fish or water fowl resulting in excessive nitrogenous waste products in the water. The recirculation of nitrogen and phosphorus from the cycle of growth and decomposition of duckweeds may also contribute to the high levels of these elements. Destroying the duckweed layer with herbicides does not solve the problem of excess nutrients in the water. In addition, the chemical herbicides may be toxic to the animal life, either directly or through biological magnification. Because of the exponential growth rate of Lemnaceae, herbicides must be used repeatedly (perhaps several times a year). Ideally, it is best to eliminate the influx of concentrated nitrates and phosphates into the water and avoid the use of concentrated fertilizers. The manual or mechanical removal of the duckweed cover can also remove a lot of the nitrogen and phosphorus nutrients. The duckweed mats can be composted and used as "green manure." They can also be fed to livestock, rabbits, poultry and fish. It has been estimated that 10 acres of duckweeds could theoretically supply 60 percent of the nutritional needs of 100 dairy cows, the manure of which could be recycled to provide fertilizer for the thriving duckweeds. According to R.M. Harvey and J.L. Fox, 1973 ("Nutrient Removal Using Lemna minor," J. Water Poll. Control Fed. 45: 1928-1938), one hectare of water area is sufficient to raise 4000-7000 chickens and ducks during a vegetation period. And according to E. Rejmankova, 1981. ("On The Production Ecology of Duckweeds," Intern. Workshop on Aquatic Macrophytes, Illmitz, Austria), one hectare of Lemnaceae cover is sufficient to produce protein for 480 ducks during the warm season. The utilization of duckweeds as food for animals is summarized by E. Landolt and R. Kandeler, pages 382-389 in Veroff. Geobot. Inst. ETH, Stiftung Rubel 95 "The Family of Lemnaceae: A Monographic Study" Vol. 2, 1987. An extensive bibliography of Lemnaceae is also given on pages 414-580. The following 3 classic papers discuss duckweed use in aquaculture:
Stopping the inflow of nutrients and the repetitive removal of the duckweed layer will greatly reduce the growth of duckweeds. Since water fowl and most fish feed on the duckweeds, they can help control the exponential population growth of these plants. In addition, Lemnaceae have a positive effect in eutrophic water because they remove ammonia which is toxic to fish in high concentrations. In general, Lemnaceae are very sensitive to herbicides. In fact, duckweeds are often used to test the toxicity of herbicides and to detect the presence of herbicides in water. According to Professor Dr. E. Landolt (pages 161-170 in Veroff. Geobot. Inst. ETH, Stiftung Rubel 95 "The Family of Lemnaceae: A Monographic Study" Vol. 2, 1987), heterocyclic compounds (e.g. 6-methylpurin), urea derivatives, and quaternary ammonium compounds (e.g. diquat and paraquat) are the most toxic substances for Lemnaceae. Some algicides, including PH 40:62 are extremely toxic to some species of Lemna. Some of these products are available from agricultural supply companies depending on federal, state or local regulations. They should be used with extreme caution and under very careful supervision. It would be advisable to consult with your city or county weed/mosquito abatement department before attempting any large herbicidal control project. Biological control using ducks, fish, turtles and crustaceans (water shrimp, crayfish, ostracods, freshwater prawns, daphnia, amphipods, etc.) may also help to control duckweed populations. There are a number of species of freshwater fish that eat duckweeds to supplement their diets, including grass carp (Ctenopharyngodon idella), channel catfish (Ictalurus punctatus), common carp (Cyprinus carpio), common mullet (Mugil cephalis), goldfish (Carassius auratus), and Tilapia (Sarotherodon), including S. mossambicus, S. hornorum, and S. nilotica. Duckweeds are also eaten by Pacu (Colossoma bidens), a freshwater fish native to the Amazon River. Some of these fish species may be available through aquafarm distributors or local county and state agencies. One aquaculture company in southern California was raising tilapia for local seafood restaurants.
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Plant bodies minute and rootless, with granular or mealy texture when rubbed between fingers of hands; generally globoid to ovoid-ellipsoid or cylindrical (flat-topped in some species); 0.4-1.3 mm long and 0.2-1.0 mm wide, floating on or partially below water surface; veins 0; pale transparent green throughout or with dark green dorsal surface; some species punctate with brown pigment cells in epidermis (visible on dead plants of W. borealis & W. brasiliensis); solitary or with smaller daughter plant attached at basal end; single, funnel-shaped budding pouch at basal end; daughter plants produced in basal budding pouch (in most species, some daughter plants may sink to bottom and function as overwintering turions); parenchyma without druse or raphide crystals of calcium oxalate; one bisexual flower produced inside dorsal floral cavity, consisting of a single pistil and single stamen (some authorities consider this to be an inflorescence with 2 unisexual flowers); pistil situated nearest the basal budding pouch; anther unilocular and apically dehiscent along pigmented line; ovary unilocular with one orthotropous ovule; utricle globose and slightly compressed, bearing 1 globose-ovoid, smooth seed with distinct conical operculum (seed may be slightly reticulate but not longitudinally ribbed); size and shape of plant body important for species identification (ideally under 10-20X magnification); at least 9 spp. worldwide, especially warm temperate and tropical regions; J.F. Wolff, German botanist and physician, 1778-1806; Armstrong, W.P. & R.F. Thorne (1984), Madrono 31: 172-179; Armstrong, W.P. (1989), Madrono 36: 283-285; Armstrong, W.P. (1985), Fremontia 13: 11-14.
All text material & images on these pages copyright © W.P. Armstrong Page 4Images of Lemnaceae in Western North AmericaLemna - Spirodela - Landoltia - Wolffia - Wolffiella - General
All text material & images on these pages copyright © W.P. Armstrong |