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The dinoflagellates (Greek δῖνος dinos "whirling" and Latin flagellum "whip, scourge") are a large group of flagellate protists that constitute the phylum Dinoflagellata. Most are marine plankton, but they are common in fresh water habitats as well. Their populations are distributed depending on temperature, salinity, or depth. Many dinoflagellates are known to be photosynthetic, but a large fraction of these are in fact mixotrophic, combining photosynthesis with ingestion of prey (phagotrophy).[1] In terms of number of species, dinoflagellates form one of the largest groups of marine eukaryotes, although this group is substantially smaller than the diatoms.[2] Some species are endosymbionts of marine animals and play an important part in the biology of coral reefs. Other dinoflagellates are colorless predators on other protozoa, and a few forms are parasitic (see for example Oodinium, Pfiesteria). Some dinoflagellates produce resting stages, called dinoflagellate cysts or dinocysts, as part of their life cycles.
Dinoflagellates are considered to be protists, with their own division, Dinoflagellata.[3]
About 1,555 species of free-living marine dinoflagellates are currently described.[4] Another estimate suggests ca. 2,000 living species, of which more than 1,700 are marine (free-living as well as benthic) and about 220 are from freshwater.[5] The latest estimates suggest a total of 2,294 living dinoflagellate species, which includes marine, freshwater and parasitic dinoflagellates.[6]
An algal bloom of dinoflagellates can result in a visible coloration of the water colloquially known as red tide.
- 1 History
- 2 Classification
- 3 Identification
- 4 Morphology
- 5 Endosymbionts
- 6 Theca structure and formation
- 7 Habitats
- 8 Nutritional strategies
- 9 The dinoflagellate nucleus: Dinokaryon
- 10 Life cycle
- 11 Harmful algal blooms
- 12 Bioluminescence
- 13 Lipid and sterol production
- 14 Transport
- 15 Genomics
- 16 Evolutionary history
- 17 Examples
- 18 See also
- 19 References
- 20 Bibliography
- 21 External links
In 1753, the first modern dinoflagellates were described by Henry Baker as "Animalcules which cause the Sparkling Light in Sea Water",[7] and named by Otto Friedrich Müller in 1773.[8] The term derives from the Greek word δῖνος (dinos), meaning 'whirling,' and Latin flagellum, a diminutive term for a whip or scourge.
In the 1830s, the German microscopist Christian Gottfried Ehrenberg examined many water and plankton samples and proposed several dinoflagellate genera that are still used today including Peridinium, Prorocentrum and Dinophysis.[9]
These same dinoflagellates were first defined by Otto Bütschli in 1885 as the flagellate order Dinoflagellida.[10] Botanists treated them as a division of algae, named Pyrrophyta or Pyrrhophyta ("fire algae"; Greek pyrr(h)os, fire) after the bioluminescent forms, or Dinophyta. At various times, the cryptomonads, ebriids, and ellobiopsids have been included here, but only the last are now considered close relatives. Dinoflagellates have a known ability to transform from noncyst to cyst-forming strategies, which makes recreating their evolutionary history extremely difficult.
Dinoflagellates are protists which have been classified using both the International Code of Botanical Nomenclature (ICBN, now renamed as ICN) and the International Code of Zoological Nomenclature (ICZN). Approximately half of living dinoflagellate species are autotrophs possessing chloroplasts and half are non-photosynthesising heterotrophs. It is now widely accepted[citation needed] that the ICN should be used for their classification.
Most (but not all) dinoflagellates have a dinokaryon, described below (see: Life-cycle, below). Dinoflagellates with a dinokaryon are classified under Dinokaryota, while dinoflagellates without a dinokaryon are classified under Syndiniales.
Although classified as eukaryotes, the dinoflagellate nuclei are not characteristically eukaryotic, as they lack histones, nucleosomes and maintain continually condensed chromosomes during mitosis. The dinoflagellate nucleus was termed 'mesokaryotic' by Dodge (1966),[11] due to its possession of intermediate characteristics between the coiled DNA areas of prokaryotic bacteria and the well-defined eukaryotic nucleus. This group, however, does contain typically eukaryotic organelles, such as Golgi bodies, mitochondria and chloroplasts.[12]
Jakob Schiller (1931–1937) provided a description of all the species, both marine and freshwater, known at that time.[13] Later Alain Sournia (1973, 1978, 1982, 1990, 1993) listed the new taxonomic entries published after Schiller (1931–1937).[14][15][16][17][18] Sournia (1986) gave descriptions and illustrations of the marine genera of dinoflagellates, excluding information at the species level.[19] The latest index is written by Gómez.[6]
English-language taxonomic monographs covering large numbers of species are published for the Gulf of Mexico,[20] the Indian Ocean,[21] the British Isles,[22] the Mediterranean[23] and the North Sea.[24]
The main source for identification of freshwater dinoflagellates is the Süsswasser Flora.[25]
Calcofluor-white can be used to stain thecal plates in armoured dinoflagellates.[26]
Dinoflagellates are unicellular and possess two dissimilar flagella arising from the ventral cell side (dinokont flagellation). They have a ribbon-like transverse flagellum with multiple waves that beats to the cell's left, and a more conventional one, the longitudinal flagellum, that beats posteriorly.[27][28][29] The transverse flagellum is a wavy ribbon in which only the outer edge undulates from base to tip, due to the action of the axoneme which runs along it. The axonemal edge has simple hairs that can be of varying length. The flagellar movement produces forward propulsion and also a turning force. The longitudinal flagellum is relatively conventional in appearance, with few or no hairs. It beats with only one or two periods to its wave. The flagella lie in surface grooves: the transverse one in the cingulum and the longitudinal one in the sulcus, although its distal portion projects freely behind the cell. In dinoflagellate species with desmokont flagellation (e.g., Prorocentrum) the two flagella are differentiated as in dinokonts, but they are not associated with grooves.
Dinoflagellates have a complex cell covering called an amphiesma or cortex, composed of a series of membrances, flattened vesicles called alveolae (=amphiesmal vesicles) and related structures.[30][31] In armoured dinoflagellates, these support overlapping cellulose plates to create a sort of armor called the theca, as opposed to athecate dinoflagellates. These come in various shapes and arrangements, depending on the species and sometimes on the stage of the dinoflagellate. Conventionally, the term tabulation has been used to refer to this arrangement of thecal plates. The plate configuration can be denoted with the plate formula or tabulation formula. Fibrous extrusomes are also found in many forms. Together with various other structural and genetic details, this organization indicates a close relationship between the dinoflagellates, Apicomplexa, and ciliates, collectively referred to as the alveolates.[32]
Dinoflagellate tabulations can be grouped into six "tabulation types": gymnodinoid, suessoid, gonyaulacoid–peridinioid, nannoceratopsioid, dinophysioid and prorocentroid.
The chloroplasts in most photosynthetic dinoflagellates are bound by three membranes, suggesting they were probably derived from some ingested algae. Most photosynthetic species contain chlorophylls a and c2, the carotenoid beta-carotene, and a group of xanthophylls that appears to be unique to dinoflagellates, typically peridinin, dinoxanthin, and diadinoxanthin. These pigments give many dinoflagellates their typical goldenbrown color. However, the dinoflagellates Karenia brevis, Karenia mikimotoi and Karlodinium micrum have acquired other pigments through endosymbiosis, including fucoxanthin.[33] This suggests their chloroplasts were incorporated by several endosymbiotic events involving already colored or secondarily colorless forms. The discovery of plastids in Apicomplexa has led some to suggest they were inherited from an ancestor common to the two groups, but none of the more basal lines have them. All the same, the dinoflagellate cell consists of the more common organelles such as rough and smooth endoplasmic reticulum, Golgi apparatus, mitochondria, lipid and starch grains, and food vacuoles. Some have even been found with a light-sensitive organelle, the eyespot or stigma, or a larger nucleus containing a prominent nucleolus. The dinoflagellate Erythropsidium has the smallest known eye.[34]
Some athecate species have an internal skeleton consisting of two star-like siliceous elements that has an unknown function, and can be found as microfossils. Tappan[35] gave a survey of dinoflagellates with internal skeletons. This included the first detailed description of the pentasters in Actiniscus pentasterias, based on scanning electron microscopy. They are placed within the order Gymnodiniales, suborder Actiniscineae.[3]
All Zooxanthellae are dinoflagellates and most of them are members within the genus Symbiodinium.[36] The association betweenSymbiodinium and reef-building corals is widely known. However, endosymbiontic Zooxanthellae inhabit a great number of other invertebrates and protists, for example many sea anemones, jellyfish, nudibranchs, the giant clam Tridacna, as well as several species of radiolarians and foraminiferans.[37] Many extant dinoflagellates are parasites (here defined as organisms that eat their prey from the inside, i.e. endoparasites, or that remain attached to their prey for longer periods of time, i.e. ectoparasites). They can parasitize animal or protist hosts. Protoodinium, Crepidoodinium, Piscinoodinium and Blastodinium retain their plastids while feeding on their zooplanktonic or fish hosts. In most parasitic dinoflagellates the infective stage resembles a typical motile dinoflagellate cell.
Theca structure and formation
The formation of thecal plates has been studied in detail through ultrastructural studies.[31]
Dinoflagellates can occur in all aquatic environments: marine, brackish, and fresh water, including in snow or ice. They are also common in benthic environments and in sea ice.
Nutritional strategies
There are several nutritional strategies in dinoflagellates: phototrophy, mixotrophy and heterotrophy. Phototrophs can be photoautotrophs or auxotrophs. Mixotrophic dinoflagellates are photosynthetically active, but are also heterotrophic. Facultative mixotrophs, in which autotrophy or heterotrophy is sufficient for nutrition, are classified as amphitrophic. If both forms are required the organisms are mixotrophic sensu stricto. Some free-living dinoflagellates do not have chloroplasts but host a phototrophic endosymbiont. A few dinoflagellates may utilize alien chloroplasts (cleptochloroplasts), obtained from food (kleptoplasty). Some dinoflagellates may feed on other organisms as predators or parasites.[38]
Food inclusions contain bacteria, bluegreen algae, small dinoflagellates, diatoms, ciliates and other dinoflagellates.[39][40][41][42][43][44][45]
Mechanisms of capture and ingestion in dinoflagellates are quite diverse. Several dinoflagellates, both thecate (e.g. Ceratium hirundinella,;[44] Peridinium globulus,[42]) and nonthecate (e.g. Oxyrrhis marina,[40] Gymnodinium sp.[46] and Kofoidinium spp.[47]), draw prey to the sulcal region of the cell (either via water currents set up by the flagella or via pseudopodial extensions) and ingest the prey through the sulcus. In several Protoperidinium spp., e.g. Protoperidinium conicum, a large feeding veil — a pseudopod called the pallium — is extruded to capture prey which is subsequently digested extracellularly (=pallium-feeding).[48][49] Oblea, Zygabikodinium and Diplopsalis are the only other dinoflagellate genera known to use this particular feeding mechanism [49][50][51]). Katodinium (Gymnodinium) fungiforme, commonly found as a contaminant in algal or ciliate cultures, feeds by attaching to its prey and ingesting prey cytoplasm through an extensible peduncle.[52] The feeding mechanisms of the oceanic dinoflagellates remain unknown, although pseudopodial extensions were observed in Podolampas bipes.[53]
The dinoflagellate nucleus: Dinokaryon
Most dinoflagellates have a peculiar form of nucleus, called a dinokaryon, in which the chromosomes are attached to the nuclear membrane. These lack histones and remain condensed throughout interphase rather than just during mitosis, which is closed and involves a uniquely extranuclear mitotic spindle.[54] This sort of nucleus was once considered to be an intermediate between the nucleoid region of prokaryotes and the true nuclei of eukaryotes, so were termed mesokaryotic, but now are considered advanced rather than primitive traits.
Life cycle
Dinoflagellates have a haplontic life cycle — with the possible exception of Noctiluca and its relatives.[3] The life-cycle usually involves asexual reproduction by means of binary fission, either through desmoschisis or eleuteroschisis. More complex life cycles occur, more particularly with parasitic dinoflagellates. Sexual reproduction also occurs (e.g.[55]) and is only known in a small percentage of dinoflagellates.[56] This takes place by fusion of two individuals to form a zygote, which may remain mobile in typical dinoflagellate fashion and is then called a planozygote. This zygote may later form a resting stage or hypnozygote, which are called dinoflagellate cyst or dinocyst. After (or before) germination of the cyst, the hatchling undergoes meiosis to produce new haploid cells.
Harmful algal blooms
Dinoflagellates sometimes bloom in concentrations of more than a million cells per millilitre. Under such circumstances they can produce toxins (generally called dinotoxins) in quantities capable of killing fish and accumulating in filter feeders such as shellfish, which in turn may be passed on to people who eat them. This phenomenon is called a red tide, from the color the bloom imparts to the water. Some colorless dinoflagellates may also form toxic blooms, such as Pfiesteria. Some dinoflagellate blooms are not dangerous. Bluish flickers visible in ocean water at night often come from blooms of bioluminescent dinoflagellates, which emit short flashes of light when disturbed.
The same red tide mentioned above is more specifically produced when dinoflagellates are able to reproduce rapidly and copiously on account of the abundant nutrients in the water. Although the resulting red waves are an unusual sight, they contain toxins that not only affect all marine life in the ocean, but the people who consume them, as well.[57] A specific carrier is shellfish. This can introduce both nonfatal and fatal illnesses. One such poison is saxitoxin, a powerful paralytic neurotoxin. Human inputs of phosphate further encourage these red tides, so there is a strong interest in learning more about dinoflagellates, from both medical and economic perspectives. The ecology of harmful algal blooms is extensively studied.[58]
At night, water can have an appearance of sparkling light due to the bioluminescence of dinoflagellates.[59][60]
More than 18 genera of dinoflagellates are bioluminescent,[61] and the majority of them emit a blue-green light.[62]
These species contain scintillons, individual cytoplasmic bodies (ca. 0.5 µm in diameter) distributed mainly in the cortical region of the cell, outpockets of the main cell vacuole. They contain dinoflagellate luciferase, the main enzyme involved in dinoflagellate bioluminescence, and luciferin, a chlorophyll-derived tetrapyrrole ring that acts as the substrate to the light-producing reaction. The luminescence occurs as a brief (0.1 sec) blue flash (max 476 nm) when stimulated, usually by mechanical disturbance. Therefore, when mechanically stimulated—by boat, swimming or waves, for example—a blue sparkling light can be seen emanating from the sea surface.[63]
Dinoflagellate bioluminescence is controlled by circadian rhythms and only occurs at night (e.g.,[64]). Luminescent and nonluminescent strains can occur in the same species. The number of scintillans is higher during night than during day, and breaks down during the end of the night, at the time of maximal bioluminescence.[65]
The luciferin-luciferase reaction responsible for the bioluminescence is pH sensitive.[63] When the pH drops, luciferase changes its shape, allowing luciferin, more specifically tetrapyrrole, to bind.[63] Dinoflagellates can use bioluminescence as a defense mechanism. They can startle their predators by their flashing light or they can ward off potential predators by an indirect effect such as the burglar alarm. The bioluminescence attracts attention to the dinoflagellate, and hence to the predator which this makes more vulnerable to predation from higher trophic levels.[63]
Bioluminescent dinoflagellate ecosystem bays are among the rarest and most fragile,[66] with the most famous ones being: the Bioluminescent Bay in La Parguera, Lajas, Puerto Rico; Mosquito Bay in Vieques, Puerto Rico; and Las cabezas de San Juan Reserva Natural Fajardo, Puerto Rico. There is also a bioluminescent lagoon near Montego Bay, Jamaica.
Lipid and sterol production
Dinoflagellates produce characteristic lipids and sterols.[67] One of these sterols is typical of dinoflagellates and is called dinosterol.
Dinoflagellate theca can sink rapidly to the seafloor in marine snow.[68]
One of the most striking features is the large amount of cellular DNA that dinoflagellates contain. Most eukaryotic algae contain on average about 0.54 pg DNA/cell, whereas estimates of dinoflagellate DNA content range from 3–250 pg/cell,[54] corresponding to approximately 3000–215 000 Mb (in comparison, the haploid human genome is 3180 Mb and hexaploid Triticum wheat is 16 000 Mb). It has been suggested that polyploidy or polyteny may account for this large cellular DNA content,[69] but studies of DNA reassociation kinetics do not support this hypothesis.
In addition to their disproportionately large genomes, dinoflagellate nuclei are unique in their morphology, regulation, and composition.
The dinoflagellates share an unusual mitochondrial genome organisation with their relatives, the Apicomplexa.[70] Both groups have very reduced mitochondrial genomes (~6 kilobases in the Apicomplexa). The genes on the dinoflagellate genomes have undergone a number of reorganisations, including massive genome amplification and recombination which have resulted in multiple copies of each gene and gene fragments linked in numerous combinations. Loss of the standard stop codons, trans-splicing of mRNAs for the mRNA of cox3 and extensive RNA editing recoding of most genes has occurred. The reasons for this transformation are unknown.
The DNA of the plastid in the peridinin-containing dinoflagellates is contained in a series of mini circles.[71] Each mini circle contains one or two polypeptide genes. The genes for these polypeptides are chloroplast specific because their homologs from other photosynthetic eukaryotes are exclusively encoded in the chloroplast genome. Within each mini circle there is a distinguishable 'core' region. Genes are always in the same orientation with respect to this core region.
In terms of DNA barcoding, ITS sequences can be used to identify species,[72] where a genetic distance of genetic distance of p≥0.04 can be used to delimit species.[73]
Evolutionary history
Dinoflagellates are mainly represented as fossils by fossil dinocysts, which have a long geological record with lowest occurrences during the mid-Triassic.,[74] whilst geochemical markers suggest a presence to the Early Cambrian.[75]
There is some evidence that dinosteroids in many Paleozoic and Precambrian rocks might be the product of ancestral dinoflagellates (protodinoflagellates).[76][77]
Molecular phylogenetics shows that dinoflagellates are grouped with ciliates and apicomplexans (=Sporozoa) in a well-supported clade, the alveolates. The closest relatives to dinokaryotic dinoflagellates appear to be apicomplexans, Perkinsus, Parvilucifera, syndinians and Oxyrrhis.[78] Molecular phylogenies are similar to phylogenies based on morphology.[79]
The earliest stages of dinoflagellate evolution appear to be dominated by parasitic lineages, such as perkinsids and syndinians (e.g. Amoebophrya and Hematodinium).[80][81][82][83]
Dinoflagellate evolution has been summarized into five principal organizational types: prorocentroid, dinophysoid, gonyaulacoid, peridinioid and gymnodinoid.[84] The transitions of marine species into freshwater have been infrequent events during the diversification of dinoflagellates and in most cases have not occurred recently, possibly as late as the Cretaceous.[85]
Recently, the "living fossil" Dapsilidinium pastielsii was found inhabiting the Indo-Pacific Warm Pool, which served as a refuge for thermophilic dinoflagellates.[86]
- Alexandrium
- Ceratium
- Gonyaulax
- Gymnodinium
- Lingulodinium polyedrum
- Noctiluca (sea ghost or "fire of sea")
- Pfiesteria
- Symbiodinium (Zooxanthella, a coral endosymbiont)
See also
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- ^ Beam, Carl A.; Himes, Marion. Ch. 8: Dinoflagellate genetics. pp. 263–298. In Spector 1984
- ^ Jackson CJ, Gornik SG, Waller RF; Gornik; Waller (2012). "The mitochondrial genome and transcriptome of the basal dinoflagellate Hematodinium sp.: character evolution within the highly derived mitochondrial genomes of dinoflagellates". Genome Biol Evol 4 (1): 59–72. PMC 3268668. PMID 22113794. doi:10.1093/gbe/evr122.
- ^ Laatsch T, Zauner S, Stoebe-Maier B, Kowallik KV, Maier UG; Zauner; Stoebe-Maier; Kowallik; Maier (July 2004). "Plastid-derived single gene minicircles of the dinoflagellate Ceratium horridum are localized in the nucleus". Mol. Biol. Evol. 21 (7): 1318–22. PMID 15034134. doi:10.1093/molbev/msh127.
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- ^ Litaker RW, Vandersea MW, Kibler SR, Reece KS, Stokes NA, Lutzoni FM, Yonish BA, West MA, Black MND, Tester PA; Vandersea; Kibler; Reece; Stokes; Lutzoni; Yonish; West; Black; Tester (April 2007). "Recognizing dinoflagellate species using ITS rDNA sequences". J. Phycol. 43 (2): 344–355. doi:10.1111/j.1529-8817.2007.00320.x.
- ^ MacRae RA, Fensome RA, Williams GL; Fensome; Williams (1996). "Fossil dinoflagellate diversity, originations, and extinctions and their significance". Can. J. Bot. 74 (11): 1687–94. ISSN 0008-4026. doi:10.1139/b96-205.
- ^ Moldowan JM, Talyzina NM; Talyzina (August 1998). "Biogeochemical evidence for dinoflagellate ancestors in the early cambrian". Science 281 (5380): 1168–70. Bibcode:1998Sci...281.1168M. PMID 9712575. doi:10.1126/science.281.5380.1168 (inactive 2015-01-12).
- ^ Moldowan JM, Dahl JEP, Jacobson SR, Huizinga BJ, Fago FJ, Shetty R, Watt DS, Peters KE; Dahl; Jacobson; Huizinga; Fago; Shetty; Watt; Peters (February 1996). "Chemostratigraphic reconstruction of biofacies: molecular evidence linking cyst-forming dinoflagellates with Pre-Triassic ancestors". Geology 24 (2): 159–162. Bibcode:1996Geo....24..159M. doi:10.1130/0091-7613(1996)024<0159:CROBME>2.3.CO;2.
- ^ Talyzina NM, Moldowan JM, Johannisson A, Fago FJ; Moldowan; Johannisson; Fago (January 2000). "Affinities of early Cambrian acritarchs studied by using microscopy, fluorescence flow cytometry and biomarkers". Rev. Palaeobot. Palynol. 108 (1–2): 37–53. doi:10.1016/S0034-6667(99)00032-9.
- ^ Saldarriaga J, Taylor MFJR, Cavalier-Smith T, Menden-Deuer S, Keeling PJ; "Max" Taylor; Cavalier-Smith; Menden-Deuer; Keeling (2004). "Molecular data and the evolutionary history of dinoflagellates". Eur J Protistol 40 (1): 85–111. doi:10.1016/j.ejop.2003.11.003.
- ^ Fensome RA, Saldarriaga JF, Taylor FJR; Saldarriaga; Taylor (1999). "Dinoflagellate phylogeny revisited: reconciling morphological and molecular based phylogenies". Grana 38 (2–3): 66–80. doi:10.1080/00173139908559216.
- ^ Gunderson JH, Goss SH, Coats DW; Goss; Coats (1999). "The phylogenetic position of Amoebophrya sp. infecting Gymnodinium sanguineum". J. Eukaryot. Microbiol. 46 (2): 194–7. PMID 10361739. doi:10.1111/j.1550-7408.1999.tb04603.x.
Gunderson JH, John SA, Boman WC, Coats DW; John; Boman; Coats (2002). "Multiple strains of the parasitic dinoflagellate Amoebophrya exist in Chesapeake Bay". J. Eukaryot. Microbiol. 49 (6): 469–74. PMID 12503682. doi:10.1111/j.1550-7408.2002.tb00230.x.
- ^ López-García P, Rodríguez-Valera F, Pedrós-Alió C, Moreira D; Rodríguez-Valera; Pedrós-Alió; Moreira (February 2001). "Unexpected diversity of small eukaryotes in deep-sea Antarctic plankton". Nature 409 (6820): 603–7. PMID 11214316. doi:10.1038/35054537.
- ^ Moon-van der Staay SY, De Wachter R, Vaulot D; De Wachter; Vaulot (February 2001). "Oceanic 18S rDNA sequences from picoplankton reveal unsuspected eukaryotic diversity". Nature 409 (6820): 607–10. PMID 11214317. doi:10.1038/35054541.
- ^ Saldarriaga JF, Taylor FJ, Keeling PJ, Cavalier-Smith T; Taylor; Keeling; Cavalier-Smith (September 2001). "Dinoflagellate nuclear SSU rRNA phylogeny suggests multiple plastid losses and replacements". J. Mol. Evol. 53 (3): 204–13. PMID 11523007. doi:10.1007/s002390010210.
- ^ Taylor FJ (1980). "On dinoflagellate evolution". BioSystems 13 (1–2): 65–108. PMID 7002229. doi:10.1016/0303-2647(80)90006-4.
- ^ Logares R, Shalchian-Tabrizi K, Boltovskoy A, Rengefors K; Shalchian-Tabrizi; Boltovskoy; Rengefors (December 2007). "Extensive dinoflagellate phylogenies indicate infrequent marine-freshwater transitions". Mol. Phylogenet. Evol. 45 (3): 887–903. PMID 17928239. doi:10.1016/j.ympev.2007.08.005.
- ^ Mertens KN, Takano Y, Head MJ, Matsuoka K; Takano; Head; Matsuoka (2014). "Living fossils in the Indo-Pacific warm pool: A refuge for thermophilic dinoflagellates during glaciations". Geology 42 (6): 531–534. Bibcode:2014Geo....42..531M. doi:10.1130/G35456.1.
- Spector, D.L. (1984). Dinoflagellates. Academic Press. ISBN 978-0-323-13813-0.
- Taylor, F.J.R. (1987). The Biology of Dinoflagellates. Botanical monographs 21. Blackwell Scientific. ISBN 0632009152.
External links
|Wikimedia Commons has media related to Dinoflagellata.
- International Society for the Study of Harmful Algae
- Classic dinoflagellate monographs
- Japanese dinoflagellate site
- Noctiluca scintillans — Guide to the Marine Zooplankton of south eastern Australia, Tasmanian Aquaculture & Fisheries Institute
- Tree of Life Dinoflagellates
- Centre of Excellence for Dinophyte Taxonomy CEDiT
- Dinoflagellates
- Judson O (5 January 2010). "A Tale of Two Flagella". New York Times.


text:
0034 965 020 584
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11 September 2015
As the 14th anniversary of the 9/11 terror attacks comes around, everyone can recall where they were when it happened. It is an occasion to pause and reflect, and remember the victims and heroes from the atrocious event.
Whilst many lives were lost, some were saved by the help of others and evacuation equipment at the World Trade Centre on September 11, 2001.
John Abruzzo, 41, a staff accountant, was working at his computer on the 69th floor of the World Trade Centre when the first hijacked plane crashed into the skyscraper.
John, who is a C 5-6 quadriplegic, relies on an electric wheelchair for mobility and remembers the day vividly. He recalls how he made his way to the hallway where his colleague was there with an Evac+chair waiting to assist him. He was transferred into the rescue device which is a universal evacuation solution for smooth stairway descent during an emergency situation without the need of great physical strength or lifting. Ten of his fellow workers stayed with him and took turns to take him down the 69 sets of stairs.
On floor 40 a dangerous thick cloud of smoke meant the team had to cut across to a stairwell on the other side of the building which was made possible by wheeling John in the Evac+chair. At the 20th floor, they heard an ear-splitting thunder that emanated from the second tower as it was hit.
Eventually they made it to the lobby where Abruzzo had to be heaved over large pieces of fallen debris. Since chunks of boulder-sized concrete made their exit impassable, Abruzzo, still harnessed to the Evac+chair, was hoisted and evacuated through a smashed window and out onto the sidewalk.
As he looked back, ground zero was behind him. An ambulance rushed Abruzzo for medical attention. He lived to tell the tale thanks to the Evac+Chair and the brave people that stayed behind to help him down the stairs.
To hear Johns full story click the link below.
20 April 2018
Evac+Chair International surveyed visitors at this year's Health & Safety Event at the NEC with surprising results. Of the visitors questioned, 55% do not include terrorism within their emergency evacuation plans.[Read Article]
28 March 2018
Evac+Chair will again be showcasing its world-leading emergency stair-descent solutions at this year's Health and Safety Event, taking place at Birmingham's NEC from 10 - 12 April 2018, stand number HS120. [Read Article]
10 October 2017
A guide to Fire Safety Risk Assessments for educational premises. [Read Article]
08 September 2017
Evac+Chair launches guide to emergency evacuation[Read Article]
20 July 2017
PGA European Tour, promoter of this year's Aberdeen Asset Management Scottish Open golf tournament, has 'paired' up with Evac+Chair International by employing the use of their stairway evacuation chairs which will enable mobility impaired guests easy access and egress in the event of an emergency situation.[Read Article]
04 July 2017
Shifting demographics and social awareness of issues such as disability are bringing greater diversity into the modern workforce. Accessibility is something that every organisation must consider. That brings specific implications for those responsible for health and safety practices within a building. Gerard Wallace, Managing Director of Evac+Chair looks at the changing workforce profile and the key issue of ensuring safety for everyone in the event of a building's evacuation.[Read Article]
07 June 2017
Product design experts at Birmingham City University have teamed up with Evac+Chair International - makers of the world's most popular stairway evacuation chair - to develop the first-ever stretcher interface device (SID) for neonates.[Read Article]

subset: commoncrawl


text:
Allah gives the example of Maryam (as), mother of Prophet 'Isa (as), as a servant devoted only to Him. Before she was born, Maryam (as) was dedicated to Allah by her mother with prayers that she would be a good servant and free from all worldly attachments:
Remember when the wife of 'Imran said, "My Lord, I have pledged to You what is in my womb, devoting it to Your service. Please accept my prayer. You are the All-Hearing, the All-Knowing." (Surah Al 'Imran, 35)
Other verses in the Qur'an, show that Allah accepted her mother's prayer about Maryam (as). We are told that she was morally pure and virtuous:
Her Lord accepted her with approval and made her grow in health and beauty… (Surah Al 'Imran, 37)
Allah chose her and gave her great honor:
And when the angels said, "Maryam, Allah has chosen you and purified you. He has chosen you over all other women." (Surah Al 'Imran, 42)
Moreover, we are told that Allah commanded her to be sincerely submissive and obedient to Him.
[The angels said,] "Maryam, obey your Lord and prostrate and bow with those who bow." (Surah Al 'Imran, 43)
At a certain point in her life, Maryam (as) left her family and moved east.
Mention Maryam in the Book, how she withdrew from her people to an eastern place. (Surah Maryam, 16)
There she lived alone and received the marvelous message of the birth of Prophet 'Isa (as). This good news was brought to her by Jibrail.
When the angels said, "Maryam, your Lord gives you good news of a Word from Him. His name is the Messiah, Isa, son of Maryam, of high esteem in this world and the Hereafter, and one of those brought near." (Surah Al 'Imran, 45)
By a miracle from Allah, Maryam (as) became pregnant untouched by any man and gave birth to Prophet 'Isa (as) in a remote place. (Surah Maryam, 20) Maryam (as) humbly accepted the fate that Allah had prepared for her and, after the birth, trusted in Allah and took refuge in Him against all the accusations of the people. The people did not understand all the miraculous events that preceded and followed the birth of Prophet 'Isa (as), and the Qur'an tells us about the hurtful calumny directed against her.
She brought him to her people, carrying him. They said, "Maryam! You have done an unthinkable thing! Sister of Harun, your father was not an evil man, nor was your mother an unchaste woman!" (Surah Maryam, 27-28)
Another verse tells us that the people rejected Allah and brought many false accusations against Maryam (as):
And [We punished them] on account of their disbelief, and their utterance of a monstrous slander against Maryam. (Surat an-Nisa', 156)
When Allah wanted Maryam (as) to take Prophet 'Isa (as) and return to her people; without any concern for what they might think about her or the accusations they would make, she obeyed His command. Even though she found herself in a situation difficult to explain, she did exactly what was expected of her. She reacted to the accusations against her as becomes a Muslim. By obeying Allah's command and being unconcerned about what people thought of her, she showed her moral character to be quite different from those ignorant people who disregarded Allah and made deities out of their fellow men.
With these examples in the Qur'an from the life of Maryam (as), Allah has given us an important message. Maryam (as) was tested in this world with wondrous events. In a very difficult environment that demanded a lot of patience, she endured the onslaught of others' accusations. As a result of all this, Allah made Prophet 'Isa (as) speak while he was still in the cradle and clear his mother of all the accusations against her.
Just like Maryam (as), everyone sincerely attached to Allah is under His protection. Whatever people may say or think about him, a person who practices the morality of the Qur'an is not concerned, because just as in the case of Maryam (as), the important thing is where he stands in the sight of Allah.


text: 'But Ruff's saga hasn't reached a conclusion yet. The Sabres remain in limbo as the NHL looks for prospective buyers. No one has placed a bid. It's a tenuous situation for Ruff, who is entering the final season of his contract. He has no idea who his future boss will be, or whether he will be wanted.'


text:
While Skype on Windows Phone 8 still falls short of the more seamless experience on Android, the chaps at Microsoft and Skype are at least working their tails off to improve the Windows Phone experience. Such is the case with the steady stream of monthly updates to the popular VOIP service, with the last occurring on July 9th.
Version 188.8.131.52 should be going live in the Store right now for users, meaning if you follow the link below you can force update the app instead of waiting 12 hours for your Store tile to refresh. More interestingly, Skype actually included a changelog with the update and it's more than the usual "bug fixes".
Version 188.8.131.52 brings the following new experience to Windows Phone:
- It's now even easier to find who you're looking for, simply filter your People list to only show Skype contacts.
- See even more favorite contacts at a glance with the new and improved layout.
- General fixes and speed improvements.
The Skype blog goes on to describe those changes in more detail, noting:
- People List Filter – We received feedback from users that they'd like to see Windows Phone contacts and Skype contacts split out in the Skype People List. So, in 2.9 we've added a filter so you can see either all your contacts, just Skype contacts, or only available contacts.
- Home Screen – The Skype home screen has been improved so that users can now pivot through app functions. The Favorite Contact tiles are now smaller, fitting three across on the screen so it is faster for you to contact the people you care about most.
Certainly the filter and new layout are the most interesting of the bunch. Indeed we've always been mixed on the old layout because while it was bright and bold, it was also a bit cramped at times. Regarding "general fixes and speed improvements", we'll never turn those down as the lag to getting into Skype has always been a downer on Windows Phone. Unfortunately, Video Messages are still missing.
You can pick up Skype for Windows Phone 8 version 2.9.0129 here in the Store, scan the QR code below or swipe right in our app. Are things better? Let us know in comments.
Source: Skype Blog; Via: Plaffo


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Windsurfing GPS units?
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Joined: 11 Jun 2005
Posts: 215
Location: SE Michigan
Posted: Fri Apr 12, 2013 9:18 pm
Post subject: Windsurfing GPS units?
Anyone know what the latest is in windsurfing GPS units? Is it still the GT31?
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Joined: 23 Aug 2001
Posts: 1068
Posted: Sat Apr 13, 2013 9:28 am
Post subject:
So it seems. There's a
GPS receiver with 10x
the sampling frequency (10 Hz) out there that is supposedly a lot more accurate, but that's for do-it-yourself projects so far. It would be nice if someone would make this into a commercially available GT-31 replacement. The 1 Hz sampling in the GT-31 is a bit too slow for accurate jibe analysis.
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Joined: 23 Jul 2008
Posts: 766
Location: Seattle, Wa
Posted: Tue Apr 16, 2013 11:41 pm
Post subject:
I understand that the GT-31 might be the approved unit for actual speed-sailing events.
But I have a Garmin GPS unit that you wear like a wristwatch, that works great for me. All I care about, is the top max speed for "that session"... and the avg. speed for the sesh, and distance traveled, etc. And in addition, if I want more than those quick stats I read off the Garmin unit... I can download the data onto the computer. And then transfer that data, to a site like Endomondo.com to see a map of where I sailed (which is interesting if you actually sail somewhere, besides back and forth). And endomondo also has other data and speed stats on the map you get, that can be interesting.
This is technology from 2 to 5 years ago (I think)... but is plenty good enough for me.
What are the new tech systems that you guys use... say, from smart phones. What data do you get... is it accurate... can you get maps and other stats on the computer with it ???
How do you do it... is it easy for the non-tech type?
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Joined: 11 Jun 2005
Posts: 215
Location: SE Michigan
Posted: Thu Apr 18, 2013 6:04 pm
Post subject:
In the past, the wristwatch style Garmins were not surviving the splash. Is yours really waterproof?
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Joined: 23 Jul 2008
Posts: 766
Location: Seattle, Wa
Posted: Thu Apr 18, 2013 10:47 pm
Post subject:
Hi Brian,
Mine is the Garmin Foretrex 201 - (I don't think they're still making 'em).
I bought it approx. 4 years ago... AND... they probably came out approx. 4 years before that. So, it's not current start of the art technology. Which is WHY I'm asking others about using new smart ph apps (or whatever the new tech is).
But regarding my Foretrex 201, you asked a question -
Well, I've been really happy with it... and wish they were still being produced.
I had heard a couple reports, about "not the best water-proofing". But I've used it quite a bit in more than 4 years... all windsurfing... many hi-speed catapults, smashing into the water, which must but GREAT water force onto the unit... And it's still waterproof. All functions still work, like new.
Most of my sailing is in freshwater... but I've also used it in saltwater many times. And when I do, I always rinse it well when I get home. I figure the possible corrosion from being soaked in saltwater (there are some exposed metal contact points where you attach the charger) would not be good. Anyway, it shows no wear from all my freshwater use. And because of the rinse I give it after being in saltwater... it still works and looks brand new.
Then only issue, happened last summer after another hi speed launch in the Gorge... one side-pin that holds the unit to the Velcro/webbing watch-band broke. It popped right out. I still need to get a new pin, which is available from a watch store. But in the meantime, I've just wrapped a rubber band around it... and no worries.
Like I said... I wish they still made 'em... some store may have some somewhere... I've never looked, as mine's fine.
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Joined: 25 Apr 2010
Posts: 23
Posted: Fri Apr 19, 2013 12:40 am
Post subject:
I attach my Garmin 301 to the mast right above the boom. It the safest place wrt any kind of falls and mast rarely goes deep in water in case of a swim.
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Joined: 23 Jul 2008
Posts: 766
Location: Seattle, Wa
Posted: Fri Apr 19, 2013 4:02 pm
Post subject:
almineev wrote:
I attach my Garmin 301 to the mast right above the boom. It the safest place wrt any kind of falls and mast rarely goes deep in water in case of a swim.
Hey Alex, how's it going
This thread, got me shopping and researching a bit...
So - Alex do you have the Foretrex 301... or the Forerunner 301 ??
Garmin makes both... and they are similar in what they look like, and their dimensions... and their features.
And for windsurfing, they would both tell me what I want to know, just the same as my good old Foretrex 201 does.
The biggest difference for us windsurfers is that the (new) Foretrex 201 has replaceable AAA batteries... and the Forerunner 301 has a built-in rechargeable battery.
THIS "might be" a concern to windsurfers, because although both units have the same water proof rating (min performance is 30 min submerged, 1 meter)... how long will the Foretrex hold up compared to the Forerunner?
My concern, is the removable door to access the AAA batteries in the Foretrex. Because over time, it seems the water proof-ness of that door (which is removed and replaced every 12 - 18 hours of battery life) will not be as water proof as the Forerunner.
The current Forerunner with the built-in rechargeable battery has a sealed case at the factory. The battery is not designed to ever be removed.
THIS is the same design as my old Foretrex 201 (built-in permanent battery)and it works great. Water-proof for several years of hi-speed crashes... charges great, lasts about the same time, at least 12 hours.
Anyway, I question whether the style with AAA batteries and a removable door will be as water proof as the style with a non-removable door.
What do all think...
Greg -
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You are currently browsing the tag archive for the 'smartphone' tag.
UPDATE: Friends pointed out Ustream or other streaming apps are best for making sure your video gets saved. My picks are aimed at those with mobile data caps, which make streaming a less-viable option.
In hectic settings like the Zuccotti Park police riots on Occupy Wall Street earlier this week, those gathering photos and video from the scene face dangers including arrests, violence, threat of having equipment lost, stolen or broken and, in some instances, having cops delete your footage.
While none of those are welcome outcomes, having your footage deleted is the most heinous to plenty of journalists. But for those shooting with smartphones, the Google+ app and Apple's Photostream part of its new iCloud services can offer a way to save your photos on the fly in case someone tries to delete them.
Both services allow you to automatically upload your footage to the Web immediately after you shoot it. Google+ saves it to your personal album on its service called "photos from your phone." Google+ also gives you the option to have it shared on your G+ page immediately, or to keep it to yourself in that album until you're ready to release it at your convenience.
To activate the automatic upload, turn it on in the photo portion of the app's settings. And don't ignore those "When to upload photos" (Wi-Fi only or Wi-Fi and mobile networks) and "on battery" options to ensure that your photos get uploaded ASAP. If you're being accosted in a park, it's doubtful you'll be able to make it to a Wi-Fi hotspot or an electrical outlet in time to save your shots.
Apple's iCloud for iOS devices serves a similar purpose with Photostream, but the automatic upload doesn't work for videos; they must be uploaded manually. Since I don't have an iPhone, I'll leave the explanation to somebody with one. Also make note that Photostream saves only your 1,000 most recent photos and they expire after 30 days.
The big thing with each service is to make sure that the automatic upload features are on before you go out shooting. I can understand not wanting all your photos uploaded to G+ all the time, so make sure to toy with the options before you go out.
It's important if you find yourself in a situation where someone's trying to delete your photos that you limit their access to your phone, buying it time to upload your shots. One of the easiest ways to hold somebody up is to lock your screen with a PIN. Don't forget you have the right to remain silent, including remaining mum on your unlock PIN. Your upload will go through unless the officer removes your battery or you're in an area with shoddy service. Also don't tell him that the footage is in the process of uploading. Then he'll be even more likely to pull the battery. I know it's fun to gloat when you've outsmarted someone, but trust me here.
If you know of some other apps that serve this purpose, feel free to share them in the comments. Offer up your own tips for keeping your photos safe in such a situation, whether with a smartphone or a DSLR.
For more on photographer and photojournalist intimidation by civilians and police, check out the Pixiq blog.
And remember: Photography is not a crime.


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Cognitive Dissonance and the Problem of Evil
This article is a summary of a portion of an interview with Social Psychologist Carol Tavris on Point of Inquiry, the Podcast of the Center for Skeptical Inquiry. She and Social Psychologist Elliot Aronson are the authors of a book on Cognitive Dissonance called Mistakes Were Made (But Not By Me): Why We Justify Foolish Beliefs and how it affects us in everyday life. It covers the manifestation of Cognitive Dissonance in prosecuting attorneys, law enforcement officials, politicians, smokers etc. In the interview she was asked if Cognitive Dissonance is manifested in religious belief and this article summarizes her response.
In the beginning of the interview she talks about characteristics of Cognitive Dissonance and how it manifests itself in Attorneys that have discovered they have wrongly prosecuted someone, law enforcement officials that are trained to believe the person being questioned is just as good as guilty thereby justifying whatever means necessary to elicit a confession, and politicians that support policy that is shown over time to be wrong but will not change their position. She uses the resulting situation of the Iraq War and the position of the Bush Administration as an example of a manifestation of Cognitive Dissonance.
Q: There are religious people that don't demand proof for their beliefs, is this a way of relieving their cognitive dissonance?
A: The more important a particular belief is to us the more strongly we will ignore or reject evidence suggesting we are wrong. Religion is central to what gives many people meaning and purpose in life. This type of belief will be defended at all costs. Examples of dis-confirming evidence creating Cognitive Dissonance are Evolution, the Holocaust and disasters.
Most religious people are not threatened by evolution. They find a way to fit it into their beliefs, but some cannot fit it into their beliefs and they will go to great lengths to try to refute the dis-confirming evidence.
How do Jews deal with the Holocaust? The Jews believe they are the chosen people, and god is looking after them. How could a good loving god have permitted genocide? Students of Cognitive Dissonance Theory would predict that people would become more religious and their faith would be strengthened. What most people do is not lose their faith in God but reduce the dissonance by saying God is responsible for the Good in the world, human beings are responsible for the Evil or God is testing faith. The Christian response to the question of how Jesus could permit enormous suffering to happen is to believe that it is to test faith. Anything that is not consonant with a belief in God is reinterpreted to make it consonant. For example after a terrible disaster the survivors will say something like "god was looking after me" but discounting the fact that God was not looking out for other people that died.
Another interesting interview related to cognitive dissonance is from the radio show "All in the Mind". They interviewed Phillip Zambardo, the lead researcher involved with the Stanford Prison Experiment. The experiment had to be canceled because it got out of control. The participants started self-justifying doing terrible things to each other and it had to be stopped. He was the expert witness for the defendants in the Abu Ghraib trial, explaining how situational factors can make good people do bad things using cognitive dissonance to self-justify their actions. He talks about it in his book The Lucifer Effect.
It made me think about slavery, the crusades, Old Testament atrocities and Craigs defense of killing pregnant mothers with a sword. (thanks Steven Carr!)
Point of Inquiry podcast with Carol Tavris interview.
Science Friday podcast interview with Elliot Aronson
Mistakes Were Made (But Not By Me): Why We Justify Foolish Beliefs
Wikipedia on Cognitive Dissonance
All in the Mind
Stanford Prison Experiment
The Lucifer Effect


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Emerson CF4801VNB Premium Select 70" Ceiling Fan in Venetian Bronze
- Accommodated many blade sets (not included) with swivel sockets
- Motor is protected for life against failure and breakage
- Personalized to your style
- Reviews
- Description
- Specifications
- Benefits
Does not include blade set. Must be purchased separately.
Emerson CF4801VNB Premium Select 70" Ceiling Fan in Venetian Bronze gives you the freedom of comfort and efficient cooling.
Flexible and Versatile
This ceiling fan installs almost anywhere for targeted cooling and ventilation. Outfit this unit with 52", 58" or 70" blades (except B61/B62/B63 models) and utilize wall or a remote for total control.
Custom Appeal
Express yourself through your appliance. With an ample range of blades from which to choose, customize your unit to reflect your taste and style.
Dependable Performance
On the hottest days you don't want to be left down and out without a superior way to cool and refresh. This unit offers dependable performance to keep you comfortable.
Limited Lifetime Warranty
Your investment is protected for life to ensure the highest level of reliability and sustained performance for as long as you own your unit.
May be subject to restocking fee.
Guaranteed Satisfaction.
- Up To 30 days to return your purchase
- Money back guaranteed
- Free return shipping
- No restocking fees
- Model: CF4801VNB
- Product Type: 70" Ceiling Fan in Venetian Bronze
- Product Color: Venetian Bronze
- Application: To cool and circulate air.
- Warranty: Limited lifetime
- Type: Ceiling Fan
- Max Air Flow: 7264 CFM
- Diameter: 52 inches
- Voltage: 120 volts, 120 volts
- Oscillation: No
- Watts: 100 watts, 100 watts
- Usage: Indoor
- Certified for Commercial Use: No
- Energy Star Certified: No
- Fan Blades Finish: Does not include fan blades - must be purchased separately
- Motor Finish: Metal
- Room Size: Large Room
- Ceiling Fan Style: Transitional/Casual
- Number of Paddles: 0
- Includes Light: Yes
- Control Type: Pull chain
- Mount Type: Standard
- Includes: Downrod , Mounting Hardware
- Remote Control Compatible: Yes
- Downrod Length: 4.5 inches
- Must Be Purchased Separately: Fan Blades , Remote Control
- Light Kit Compatible: Yes
- Optional Accessories: Remote Control , Light Kit , Fan Blades
- Accommodated many blade sets (not included) with swivel sockets
- Compatible with 52", 58" and 70" blade sets
- Modular design fits in a variety of areas
- Motor is protected for life against failure and breakage
- Personalized to your style
- Reduced energy consumption doesn't compromise power


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Juelz Santana (Feat. Un Kasa & Richmond Rabb) - Aggy [Skull Gang]
*World Premiere*
The official video for Skull Gang's "Aggy" featuring Juelz Santana, Un Kasa and Richmond Rabb, from their mixtape Juelz Santana Presents Skull Gang Takeover (The New Movement To Move Wit) in stores now. Directed by Scenario.
Please click the "Report" button below if the video on this page is not working properly.
Comments will only be displayed by verified emails. No Spamming or Racism.


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GRAND ISLAND, Neb.—No injuries have been reported after a small plane ran into a terminal at the Central Nebraska Regional Airport in Grand Island.
The Grant Island Independent reports ( http://bit.ly/XOEHyT) that fire crews were dispatched to the scene just before 3 p.m. Wednesday because fuel was leaking from the single-engine aircraft.
Airport Executive Director Mike Olson says the pilot had started the engine and was preparing to taxi onto a runway when he lost power to the brakes and nose wheel control. The plane rolled into the front of the Trego Dugan terminal, breaking a window and denting the plane's nose.
Records say the plane had landed in Grand Island on its way from Chicago to the Rocky Mountain Metropolitan Airport near Broomfield, Colo.
Information from: The Grand Island Independent, http://www.theindependent.com


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Celebrating illustration, design, cartoon and comic art of the mid-20th century.
Friday, September 26, 2008
The Bu$ine$$ of Illu$tration: Comic Books
"This field uses much the same type of talent employed by the syndicates," begins Rodewald, "but mostly on a free-lance instead of a salary basis."
"It is perhaps the lowest-paid field in the whole art business (twenty-five dollars for a whole page of four to six line drawings is not uncommon), and it is worth considering only as a refuge of desperation. A young beginner may glean a certain amount of valuable experience from this work; otherwise only an artist of phenominal speed or one who works on a royalty basis can hope to earn more than the barest subsistence."
Its mind-boggling to think that an illustrator of the calibre of Don Heck, who drew the page above in 1959, would be willing to pour this much quality into work that paid only $25 per page. Jim Amash has interviewed over a hundred comic book professionals from the early days of the business for Alter Ego magazine and says, "almost all of them said they had wanted to get out of the business - if they could have. Some, like Jerry Robinson, tried showing their samples to editors in more lucrative markets like book and magazine publishing. But there was a stigma attached to the comics industry, and as soon as editors saw the samples were comic pages, they didn't care how good they were, they wanted nothing to do with you."
Some artists like Jack Kamen left the comic book business for exactly that reason. Kamen, who quit the comics the same year Fred Rodewald's article was published, found he could do similar style line drawings for advertising clients at $40 to $50 per drawing. In an interview in The Comics Journal #240, Kamen says he could walk into a New York ad agency, "and walk out with $1,500 worth of work. I'd say, 'What do you got for me?' 'Well, we've got 30 or 40 drawings over hear at $40-$50 bucks a piece.' 'I'll take it.' "
Ten years later, page rates in the comics business had only gone up about ten or fifteen dollars, and even then, the business was so unstable that they would sometimes drop down to as low as twenty dollars. Long-time industry veteran, John Romita Sr., described in a 2001 interview how the comics business of the late 50's/early 60's was "a roller-coaster."
"I remember going from $20 dollars a page, pencil and ink, to $40 a page, pencil and ink. And then, the next two years, every time I took a job in, I got a cut and I ended up at $20 a page again. We were flying without a parachute. I used to be afraid of getting a mortgage."
In spite of those stressful conditions, the business was able to attract some remarkable talents. John Buscema (above), undisputably one of the finest draftsmen the comics have ever seen, was working then at Fredman~Chaite studios as a storyboard artist.
In an interview published in Alter Ego #15, Buscema says, "Advertising's a brutal life, believe me. For example, I would work a whole week and get ready to go home on a Friday night, and I'd get a call - the owner of the studio would get a call from an agency, that there was a campaign that had to be in by Monday. You know, when you talk about a campaign, you're talking about dozens of artists working on one product. It could be cars, it could be cigarettes, it could be anything, and you'd have to stay in the city. I'd have to call my wife and tell her I wasn't going to be home for the weekend."
"So when I got the call from Marvel, [then Marvel editor-in-chief] Stan [Lee] said, 'I don't know if you'll agree to work, We'll make you happy, financially, and we have plenty of work' I never thought twice and I returned to comics."
By then it was the mid-60's and page rates for top talent had climbed to around $60, according to Jim Amash, who was quoted that number by one of the industry giants, Jack Kirby.
John Buscema drew on his advertising art acumen in his approach to his comics work. Reflecting Rodewald's advice that one must be incredibly fast to make a living in comics, Buscema once told a group of young professionals during a 'chalk-talk' tutorial, "Are you guys familiar with the one line? ... You put in three or four lines, I put in one. I did that in half, a third, a quarter of the time.... We picked that up in advertising. This is a very simplified way - its supposed to be very, very sketchy. But carried it over into comics. ... You guys want to struggle along with the pages, you want to fill in a black background... what's the point of filling in a whole black area when you can just scribble it in? ... Your problem is to tell a story the best possible way, the fastest possible way. ... Do your job. If you earn your own pay, that's it. You're in this for the money!"
I asked several industry professionals for a rough estimate of page rates typical of mainstream comics today. One artist, who has been out of the business for some time now, told me he was recently offered an assignment at $300 a page, pencils and inks. He told me, "That's about what the rate was in l980."
"When I was working in comics," he continued, "it was a decent living. With royalties you could expect to do fairly well on a monthly book. Just about everything else I've worked on as an artist has paid better- animation, storyboards, advertising...probably even signpainting if I had tried that."
Other artists confirmed this number, although noting that there are some working for less. However, said one pro with more than a decade of experience, "As long as you can be consistent and get things done (mostly) on time, it's more than possible [to make a living in comics]. There's also additional income from royalties (if you're on a book that sells well enough), trade paperbacks, and original art sales. I also do commission work through my art dealer. I'm not rich, by any measure, but I manage to pay all my bills on time and I haven't had to skip any meals yet."
"It's definitely not as lucrative as mainstream illustration work, but it's a hell of a lot more fun!"
And echoing that sentiment are these sage words from one long-time professional comic artist:
"I'm occasionally surprised how much, and how little, some artists are paid per page, but you can do well either way if you apply yourself to the work. If you get into a field or have a specific career based on the potential of making money you've lost already. Having a passion for what you do will always bring a monetary reward, some will become wealthy but most will have a very enjoyable life doing what they love to do, and that ain't bad, considering."
10 comments:
Nice article. I always like hearing these sorts of things in context to the time period.ReplyDelete
So what is $25 a page equivalent to in far as today's page rates?
I had the sheer joy of filming John Buscema for the Frazetta documentary. At the end of the interview he did a sketch for a book we were going to give to Frank at the end of filming.
Watching him start with a wrist, and sticking true to his "one-line" rule Conan emerged out of nowhere and we all gasped at the ease and proficiency of such seasoned skills.
He was sooo damn good.
- Harley5:35 PM
Yikes! never wanted to do comic books (especially the non-cartoony kind), and I'm too slow and deliberate anyways. Waaaaaaaay too slow.ReplyDelete
This is a sad story, but at least these fellas seem to enjoy their work.
Thanks again for another great presentation. These business posts have been fascinating.
Shane, I'm dying of envy that you had this amazing experience - you lucky bastard. How great that such a film exists... is it online somewhere? I'm out of the loop on this stuff, I'm afraid...ReplyDelete
I hope you don't mean people working in the mainstream for major publishers are getting paid $25 a page, because if that's so, we're going to have to stage an intervention.
harley; thanks for your feedback - its good to know these posts were appreciated. Let's hope younger readers give what was presented this week some thoughtful consideration. Whatever career path they choose, these posts were intended to provide some historical context and some perspective on current opportunities.
Thanks for this post. I'm an art student studying illustration and I'm interested not only in comics, but also in animation, character design, and storyboards. How much of this article still applies to today? For me, as long as I have a job that pays me to draw (pays me enough to get my bills done and eat) I'm pretty happy. Keep in mind that I'm super naive, though. :DReplyDelete
There are a lot more comic companies out there now, so it's good for someone starting out, but there are smaller pay rates for everyone.ReplyDelete
Seasoned pros who made a lot of money in the 90s can't get much comic work these days. Now the best paying work is in Advertising! Storyboards can be a easy job for someone used to doing comics.
I agree with Leif,about ShaneReplyDelete
"I'm dying of envy that you had this amazing experience"
Have you seen this.. John Buscema Drawing Capt. America and ink...
Fantastic article. Too many comics history stories concentrate on art styles, genres and characters ... without talking about the money. Thanks, thanks, thanks!ReplyDelete
$300 a page, pencils and inks. He told me, "That's about what the rate was in l980."ReplyDelete
Thats $216,000 a year in 1980. I would say thats pretty damn good.
Even King Kirby got $57,600 in 1966, pretty good for the time.
Yes theyn should've got a piece of the action but they were far from paupers. At least for the top guys.


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Info-Tech M-300C Discontinued!
Please check
Please check
|Discontinued RTTY & Facsimile Equipment
The Info-Tech M-300C tri-mode microprocessor keyboard generates International Morse Code, Baudot and ASCII. There is TTL output for connection to the M-600A to monitor outgoing transmissions. The AFSK output is Mark at 2125 Hz and Space at 2295 (170 Hz shift) or 2975 (850 Hz shift). There is RTTY FSK isolated output for the FSK input to a transmitter or may also be used to key a printer loop circuit. Supported Baudot speeds are 60, 66, 75, 100 and 132 WPM. ASCII at 110 or 300 baud. There is a built-in RY and Quick Brown Fox generator. There is a 700 character running buffer and 11 message memories of 120 characters each.
Requires 115 VAC 50/60 Hz 15W. 14.25 x 4 x 12 inches 11 lbs. With Owner's Manual.
Copyright 2005-2018, Universal Radio, Inc.


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TSP Innovations necesitaba un nuevo diseño de logotipo y lanzó un concurso de diseño en 99designs.
Un ganador ha sido elegido entre 363 diseños de 100 diseñadores freelance.
yakstand.com is an exciting new blog concept using the content of magazines to spark engaging conversation and interaction with it's members. Many of us have read a magazine and been encouraged by an informative how-to article, sold by a great product review (as I was with 99designs in Fast Company) or received a call to action by a heart warming article. At yakstand.com you can further build on those emotions and interact with a community of members who can relate and offer further information on the topic, expand your knowledge past the information received from the article, give a different viewpoint on the subject matter or offer their personal experience with a product or service.
Our goal is to bring an extension to the content of magazines and provide our community with a site full of relevant discussion, magazine news and targeted product offerings.
Avid magazine readers of all ages who would like to take their experience to a new level.
Let me give you a brief background on the name YAKSTAND. Since we are a magazine/newspaper discussion blog we decided to merge a couple words into the name. First is YAK, which is to talk, gab or chatter. Second is STAND, from newsstand.
We are interested in logos that embody the traditional Web 2.0 style including a full size logo as well as the square icon (ex. Facebook, Twitter, etc.).
We have uploaded a sample idea that we came up with. By no means is that the only design we want. Use it as inspiration and BE CREATIVE! We would like to see options with the Yak (animal) incorporated and/or magazines/newsstands.
Be SIMPLE and MEMORABLE. Web 2.0 logos are not complicated or detailed. BE ORIGINAL. We will not accept logos that are too similar to other logos used by other companies.
Please integrate the name yakstand.com in the logo either UNDER the main graphic or to THE RIGHT SIDE. We like the .com to be smaller font and vertical after the name. Please do not use script or overly curvy fonts.
Our tagline is Regurgitate What You Read so integrate that as well.
For colors use blue, green, orange or red (or any subsequent shade). Again, BE CREATIVE and use what works well for your design.
We will need .AI and layered .PSD files as well as the font names used.
Descubre cómo funciona un concurso de diseño u obtén más información sobre nuestros servicios de diseño de logo.


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Mitt Romney has singled out the Drudge Report as one of his favorite websites. But the candidate's public embrace of the right-wing publication is more than an effort to win over conservative readers: Romney actually believes in some of the debunked conspiracy theories extremist groups peddle.
During a town hall in Ohio on Wednesday, Romney responded to question about the United Nations by declaring that the international body will undermine Americans' Second Amendment rights and dictate how families should raise their children:
ROMNEY: Turning to the United Nations to tell us how to raise our kids, or whether we can have the Second Amendment rights that our Constitution gave us, I mean, that is the wrong way to go, right? Do not cede sovereignty. I'm happy to talk there. I'm not willing to give American sovereignty in any way, shape or form to the United Nations or any other body. We are a free nation. We fought for freedom and independence. We are going to keep freedom and independence.
Watch it:
The "guns" reference concerns Arms Trade Treaty, a U.N. initiative to prevent weapons from getting into the hands of terrorists and genocidaires. Both Drudge and the NRA have insisted that it poses a threat to American gun ownership. However, as ThinkProgress has documented, the treaty can't and won't: there are no provisions being negotiated in the treaty that affect domestic gun ownership, the State Department has publicly committed to rejecting any treaty that does, and Constitutional protections for gun ownership would trump a U.N. treaty according the Supreme Court even if a treaty infringing on the Second Amendment somehow made it through Congress.
Romney's "Telling us how raise our kids" conspiracy refers to the notion peddled by pundit Dick Morris. He claims that the U.N. is coming for our children through the Convention on the Rights of the Child, a treaty expressing standards for the acceptable treatment of children that every country — except Somalia and the United States — has ratified. The Convention "can only be implemented through domestic legislation enacted by Congress or state legislatures, in a manner and time-frame determined by our own legislative process." Thus, the U.N. can't force the United States to pass laws interpreting Treaty provisions in any particular fashion and "contains no controlling language or mandates" for the signatory nations.


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- 1 1/2 pounds large yellow or red beets, washed and trimmed
- 1/2 cup rice wine vinegar
- 1/2 cup granulated sugar
- 1 tablespoon extra-virgin olive oil
- 1/2 recipe Herbed Goat Cheese, recipe follows, cut into 8 round slices
- 1 to 1 1/2 cups mixed baby lettuces, washed and dried
- 1/4 cup Spago House Dressing, recipe follows
- 1/4 cup Citrus Hazelnut Vinaigrette, recipe follows
- 1 ounce toasted hazelnuts, recipe follows, coarsely chopped
Preheat the oven to 400 degrees F.
Place the beets in a small roasting pan and pour in enough cold water to reach about 1/4 of the way up the sides of the beets. Cover the pan with foil, and roast the beets until they are tender, 2 hours to 2 hours and 15 minutes. To check for doneness, gently insert a bamboo skewer into a beet. The skewer should slide through easily. Remove the beets from the pan, allow to cool, and then peel.
Cut each beet into 1/4-inch-thick round slices. Cut each slice with a 3-inch round cookie cutter. (You will need 10 rounds). Cut trimmings into 1/4-inch dice and reserve 1/2 cup for garnish.
In a saute pan, bring vinegar and sugar to a boil. Lower to a simmer and poach beets 1 minute on each side. With a slotted spatula, remove and place on a baking tray lined with parchment paper. Cover and refrigerate until needed.
When ready to assemble, heat the olive oil in a small saute pan. Arrange the slices of Herbed Goat Cheese in the pan and warm slightly, turning them with a small spatula just to warm both sides. This has to be done quickly, or the cheese will melt.
To assemble the napoleons, place 1 of the beet rounds on a firm, flat surface and begin to layer. Top with a slice of goat cheese, then another beet round, a second slice of cheese, another beet round, a third slice of cheese, and a beet round. (Continue until you have 5 layers of beets and 4 layers of cheese.) Carefully cut through layers, dividing into 3 wedges. Repeat with the remaining beets and cheese.
To serve, arrange 3 of the wedges, pointed ends facing out, in a circle in the center of each plate. Toss the baby lettuces with the Spago House Dressing and mound half of the lettuces on top of each arranged napoleon. Drizzle 1/2 of the Citrus Hazelnut Vinaigrette around each mound. Sprinkle toasted nuts and reserved diced beets on top of drizzled vinaigrette. Serve immediately.
Herbed Goat Cheese:
- 2 teaspoons chopped fresh flat-leaf parsley leaves
- 2 teaspoons chopped fresh chives
- 1 teaspoon chopped fresh thyme leaves
- 1/2 teaspoon freshly ground black pepper
- 1 (7 or 8-ounce) log goat cheese
Combine the parsley, chives, thyme, and pepper and place on a flat surface. Roll the log of cheese in the mixture, coating all sides and retaining the shape of the log. Wrap in plastic wrap and refrigerate until needed.
Yield: 1 (7 or 8-ounce) log
Recipe courtesy Wolfgang Puck, "Pizza, Pasta & More!," Random House, 2000
Spago House Salad Dressing:
- 2 large shallots, minced (1 heaping tablespoon)
- 1 tablespoon Dijon mustard
- 2 tablespoons zinfandel vinegar
- 2 tablespoons sherry wine vinegar
- 1/2 cup olive oil
- 1/2 cup vegetable oil
- Salt
- Freshly ground white pepper
In a small bowl, whisk together the shallots and the mustard. Whisk in the vinegars and then the olive and vegetable oil. Season with salt and pepper, to taste. Transfer to a covered container and refrigerate until needed.
Yield: 1 1/4 cups
Recipe courtesy Wolfgang Puck, "Adventures in the Kitchen", Random House, 1991
Citrus Hazelnut Vinaigrette:
- 1 1/2 cups fresh orange juice
- 1 shallot, peeled and minced
- 1 teaspoon minced fresh thyme leaves
- 2 tablespoons balsamic vinegar
- 1/2 teaspoon orange zest
- 1/3 cup hazelnut oil
- 1/3 cup extra-virgin olive oil
- Salt and freshly ground black pepper
In a medium saucepan, bring orange juice to a boil. Lower to a simmer and reduce until only 1/3 cup remains. Cool to room temperature.
In a medium bowl, combine orange juice, shallot, thyme, vinegar, and orange zest.
Slowly whisk in both oils until thick and emulsified. Season with salt and pepper. Refrigerate until needed.
Yield: 1 1/3 cups
Recipe courtesy Wolfgang Puck, "Pizza, Pasta & More!," Random House, 2000
Toasted Hazelnuts:
- 1 cup hazelnuts
Preheat the oven to 350 degrees F. Arrange the hazelnuts on a medium baking tray and toast for 10 to 12 minutes, turning after 5 minutes. (This can be done in a toaster oven). Remove to a clean towel and rub to remove as much of the skin as possible. Chop coarsely and use when needed.
Yield: 1 cup
Recipe courtesy Wolfgang Puck, 2000
My much similar version below:


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In the fall of 1967 University campuses across the country became important centers of protest against the Vietnam War. This associated press article appearing in the Greenfield Recorder refers to demonstrations in California, Wisconsin and Washington. When the principal supplier of Napalm to the United States military, the Dow Chemical Company, sought future employees at the University of Wisconsin at Madison, students occupied a building in protest of military and corporate recruiting on campus. The University of Wisconsin chancellor ordered local police to clear demonstrators from the building. This led to violent encounters with student protesters and mass arrests. Sixty-five people were injured during the confrontation. Similar incidents occurred at the Berkley campus of the University of California. The author points out that these protests drew forth strong responses on both sides. The National Committee for Responsible Patriotism claimed to have "received hundreds of inquiries as a result of the anti-war demonstrations." It was the general belief of these callers that the anti-war protesters showed "disrespect for law and order."
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"Violence Seen Probable in D.C. Demonstration" article from The Greenfield Recorder newspaper
"UM Students Protest Move Into Cambodia" article in The Greenfield Recorder newspaper
"War Protest Keeps New England College Campuses in Ferment" article in The Greenfield Recorder newspaper
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HESPERIA >> An early morning head-on crash killed a woman and critically injured four other people after one of the cars apparently crossed over into oncoming traffic, sheriff's officials said in a news release Saturday.
Maria Zavala, 44, of Hesperia was pronounced dead at the scene, coroner's officials said.
The crash was reported in the 10700 block of Balsam Avenue near Pendleton Street around 3:30 a.m., authorities said.
Sheriff's officials said two men in a 2007 BMW sedan were traveling at an undetermined rate of speed when they crossed over into oncoming lanes, striking a 2011 Toyota Camry driven by Zavala. Two passengers were in her car, both teenage girls.
Firefighters extricated two victims who were trapped in the wreckage.
Weather conditions prevented the use of air ambulances, so the injured were taken by ground to Loma Linda University Medical Center and Arrowhead Regional Medical Center in Colton.
Authorities had not determined if alcohol or drugs played a role in the crash.
Anyone with information or who may have witnessed the collision is asked to call sheriff's Deputy Charles Foster at 760-947-1500.
Staff Writer Doug Saunders contributed to this report.


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Feb. 17, 2014
Florida State 75, UNC 81 2013-14 Season Statistics Coaches' Quotes Photo Gallery
TALLAHASSEE, Fla. (Seminoles.com) — Ian Miller, Montay Brandon and Aaron Thomas combined to score 56 points but it wasn't enough in Florida State's 81-75 loss to North Carolina at the Donald L. Tucker Center Monday night.
The triumph was the Tar Heels' (18-7, 8-4 Atlantic Coast Conference) seventh in a row while FSU fell to 15-11 and 6-8 in the mighty ACC.
The Seminoles were within two scores with 21 seconds to play but couldn't capitalize on some missed UNC free throws by converting on the offensive end. The Tar Heels finished the game by scoring 75 points in the final 30 minutes of action after going into the locker room at halftime down 36-33.
When it came down to it, UNC's depth coupled with FSU's quick turnaround following Saturday' win at Wake Forest proved to be the difference.
"We'll work through it," FSU coach Leonard Hamliton said afterwards, citing his team's lack of depth at this point in the season as the "hand we've been dealt."
Miller has been FSU's ace off the bench all season but Hamilton elected to start the talented senior at point guard. The Charlotte, N.C. native responded by scoring 22 points on 7-of-16 shooting.
He was aided in his offensive effort by back-court mates Thomas and Brandon. Thomas finished with 16 points while Brandon poured in a career high 18. Okaro White was limited in his minutes as he battled with foul trouble and finished with four points.
"We are not quite as deep as we have been in the past," Hamilton said. "For us to lose our most experienced senior player, it probably was a little more challenging than it would have been in other circumstances."
Defensively, the Seminoles couldn't defend the paint against Kennedy Meeks, who dropped 23 points on the hosts and grabbed seven rebounds. Marcus Paige had just four first-half points but erupted for 16 in the second frame to finish with 20 from his point-guard spot.
Brice Johnson (14) and J.P. Tokoto (10) also scarred in double figures, which helped counteract the fact that UNC standout James Michael McAdoo was held scoreless before fouling out in the second half. The Tar Heels shot 51.6 percent from the field for the game while FSU finished 43.1 percent in the same category.
North Carolina — one of the nation's top rebounding teams — grabbed 42 boards to the `Noles' 33 but FSU had just eight turnovers. Carolina had six blocks and scored 50 of its 81 points in the paint.
FSU built a 21-6 lead to start the game on Thomas' reverse lay-up before a North Carolina timeout at the 11:13 mark of the first half.
But UNC chipped away in a foul-plagued opening frame and cut the deficit to a one-score game with less than 30 seconds before the break. Gilchrist's nifty pass to a cutting Brandon was then fouled by a Carolina 3-pointer as time expired for a 36-33 FSU advantage at halftime.
Despite eight early points from Miller in the second half, Carolina tied the score at 53-53 and then built a 70-62 lead on Paige's 3-pointer with 6:03 left to play. Brandon's lay-up and free throw then made it a five-point contest before White and Thomas combined for four free throws to cut the score to 70-69.
A J.P. Tokoto dunk and two more points-in-the-paint from Meeks gave the `Heels a 74-69 but the visitors couldn't convert on two separate one-and-one opportunities from the charity stripe to extend the advantage and Brandon's put-back made it a 74-71 UNC lead with 30 second left.
Paige then hit a pair of free throws and the Seminoles couldn't cut it any closer than a 76-72 score down the stretch as UNC hit five of its six final free throws.
"We got beat by a team that had a little more firepower than we had," Hamilton said.
A tough task now awaits the `Noles on the road as they travel to play at Pitt Feb. 23. Tip-off from the Peterson Events Center is scheduled for 7 p.m. on ESPNU.


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Federal Inmate Indicted On Contraband Charge
ERIE, Pa. - An inmate at the McKean Federal Correctional Institution in Bradford, Pennsylvania, has been indicted by a federal grand jury in Erie on a charge of possession of contraband in prison, United States Attorney David J. Hickton announced today.
The one-count indictment named Pedro Kline, 25, as the sole defendant.
According to the indictment presented to the court, from on or about May 11, 2013 to on or about May 13, 2013, Kline was in possession of contraband, namely, marijuana.
The law provides for a maximum total sentence of five years in prison, a fine of $250,000, or both. Under the Federal Sentencing Guidelines, the actual sentence imposed would be based upon the seriousness of the offense and the prior criminal history, if any, of the defendant.
Assistant United States Attorney Christian A. Trabold is prosecuting this case on behalf of the government.
Officers of the McKean Federal Correctional Institution conducted the investigation leading to the indictment in this case.
An indictment is an accusation. A defendant is presumed innocent unless and until proven guilty.