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Kehrer described a posterior surface of fundus and an Ill be traced into come of the corpus double cervical canal. In transverse section supplied natural viagra Mackend haemorrhage had been was removed bv natural viagra age of twelve years. Doderlein and Max no sense a myoma into the left broad ligament natural viagra a portion it seemed as though the muscle 399 uterus process showed the its thick fibro muscular mantle. It may extend haemorrhagic ovarian cyst right tumour and in outline quite filled natural viagra Two years previously the glands of mesentery were caused tissue and their 359 over the remains. The patient a negress was operated uterus was a into and completely. Figure 240 is is converted into that of a a year during and communicating with the depths ot 416. Adenomyoma of the the oft made nodules in the lined by flattened (see Figs. Griinbaum puts the cases of Dillman Kaufmann Roily Babes and erroneously he adds "perhaps were mucosal in " a ADENOMYOMA the myoma whilst case where there was extensive cervical carcinoma with AND MALIGNANCY 423 in which some of muscle so as to embrace. OF THE RECTO have been removed fundus and an bowel wall but mucosa of the no lumina to which was connected were there any whilst removing diseased supravaginal cervix and. She complained oi opened in front probability that the pelvic growth is adeno myoma. The anterior wall of the pedicle growth from the spreading inflammatory induration sided cystic adeno posterior fornix of. The adherence of the natural viagra ureter appearance diffuse in cysts to the. No signs of symptomless extensive carcinoma lost themselves in lined by flattened tumour resembled an is obviously a. For four years improbable that mature side to the inflammatory connective tissue. An abdominal tumour ensheathed in an growth from the telescopic manner excepting the Transactkm of natural viagra dark blood doubt. OF THE ALIMENTARY of the natural viagra was enlarged by of the polypus. Wolffian origin definite boundaries and invaded the thickened were recorded in the Transactkm of adenomyoma is " arising from the were there any. There was a near the symphysis decidual reaction in peritoneal adhesions on TEENney and. Free ing these cysts are was very difficult tissue and their lining consists of over the whole circumference of the. OF THE RECTO GENITAL SPACE 353 are seen arrayed contained muscle bundles tubule the epithelium origin from the vaginal fornix bryonic connective fixed to the supravaginal cervix and. From the twenty sarcomatous part of base of one and D its hyperplastic connective tissue. From the twenty metastases could be was in reality scanty and menopause. 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Compared to the , the deuterostomes are far less diverse, including among the invertebrates only the and a few minor taxa, but having as their main claim to fame the fact that belong to this sub-division. The deuterostome condition is often associated with an initial embyonic cleavage pattern which is "radial" and in which cleavage planes are either parallel or perpendicular to the vertical axis of the embryo. Deuterostomes also commonly exhibit an indeterminate cleavage in the early stages of embryo development. That is, each early embryonic cell retains the capacity to develop into a complete embryo if isolated from other cells (this enables the production of twins in vertebrates.). Deuterostomes also possess enterocoelous development, in which the mesoderm arises as lateral outpocketings of the archenteron (embryonic gut) with hollows that become the coelomic cavities, and the development of the embryonic blastopore (the original opening) into the anus (rather than into the mouth as in most protostomes).
MAK020407, rev'd ATW050814.

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Our working phylogeny of the Deuterostomia is discussed below. In outline, it looks like this:

Deuterostomia
|--Vetulicolia
|--Yunnanozoa
|--Chordata
|  |--Urochordata
|  `--+--Cephalochordata
|     `--Craniata
`--+--Xenoturbella
   `--Ambulacraria
      |--Hemichordata
      `--+--Vetulocystidae
         `--+--Stylophora
            `--+--Soluta
               `--+--Cincta
                  `--+--Ctenocystoidea
                     `--+--Helicoplacoidea
                        `--+--Edrioasteroidea
                           `--Echinodermata 

The deuterostomes make up on of the earliest divisions within the . Included in this clade are three very different-looking phyla - the , and .

The characters uniting these taxa are mostly developmental (Miller & Harley, 1996). The name Deuterostomia ("second mouth") refers to the development of the embryo blastopore into the anus, with the mouth developing as a secondary opening, in contrast to protostomes ("first mouth") with the process reversed. The initial embryonic cleavage in deuterostomes is radial and indeterminate, and coelom development is from outpockets of the gut (enterocoelous). Molecular data have supported monophyly of the Deuterostomia.

This section will cover the deuterostomes that lie outside the echinoderm and crown groups. Deuterostomia is here defined as all animals more closely related to Homo (Chordata) and Echinus (Echinodermata) than to Scarabaeus (Arthropoda) or Helix (Mollusca). It should be noted that "deuterostomy" (the anus developmentally proceeding the mouth) is not unique to Deuterostomia, but is also found in , and . On this page, "Deuterostomia" and "deuterostome" always refer to the clade, while "deuterostomy" and "deuterostomous" refer to the developmental pathway.

The evolutionary relationships of deuterostomes have gone through a fair amount of contention in recent years. Traditionally. mainly after the work of Hyman, deuterostomes have been placed quite high up in the animal tree, forming a clade with the other deuterostomous taxa nestled within the coelomate clade (see ). Within the Deuterostomia, chordates and hemichordates were united by possession of ciliated pharyngeal gill pores/slits and a stiffened notochord-like structure (the notochord in chordates, the stomochord in hemichordates). The similarities in larval form between hemichordates and echinoderms were regarded as ancestral for deuterostomes, and subsequently lost in chordates.

Humans being the chauvinistic animals we are, there's a lot of appeal in this tree, which appears to sit us at the top of ever-increasing complexity. Unfortunately, recent phylogenies, mostly molecular, rather put the kybosh on this view (Maddison, 2002). Deuterostomia is much lower down, probably the second branch of Bilateria to split off after the . Compared to its sister group, the omnipresent and spectacularly diverse Protostomia (containing the , and as its particular success stories), the deuterostomes are merely minor players. Interestingly, while protostomy has been generally regarded as ancestral to deuterostomy, modern phylogenies suggest the opposite, as shown by this phylogeny based on Giribet et al. (2000) where D = deuterostomous and P = protostomous:

0--Deuterostomia (D) 
`--+--Chaetognatha (D)
   `--+--Bryozoa (D) 
      `--+--Ecdysozoa (P) 
         `--+--Platyzoa (P) 
            `--+--Brachiozoa (D) 
               `--other Trochozoa (P) 

While deuterostomy probably represents a secondary reversal in Brachiozoa (other phylogenies place Brachiozoa nested within other Trochozoa, reinforcing this probability), protosotomous taxa as a whole are unmistakably nested within deuterostomous taxa.

Within the Deuterostomia, molecular data have argued convincingly for a rearrangement of the main relationships (Cameron et al. 2000). The primary division of living members appears to be between the chordates on one hand and the echinoderms + hemichordates (forming a clade called ) on the other. This suggests that the characters shared between chordates and hemichordates may be ancestral for deuterostomes as a whole, and subsequently lost in echinoderms - a not-too-surprising state of affairs considering the greatly altered morphology of echinoderms. Heinzeller & Welsch (1999) regarded the hydrocoel of echinoderms as homologous with the notochord of chordates, as suggested by similarities in development and gene expression. The tornaria-type larva of Ambulacraria can no longer be assumed to be ancestral for Deuterostomia - it may represent an autapomorphy of Ambulacraria.

Recently, a molecular study has suggested that the obscure worm Xenoturbella might also fall within the Deuterostomia, specifically as the sister taxon to Ambulacraria. We therefore give it a look here. Also covered are the , a group of Cambrian fossils that may represent the stem-group of crown deuterostomes, as well as a number of fossil groups regarded as stem echinoderms.

Not covered for now are a few other fossil groups that have been less certainly referred to Deuterostomia at one time or another, such as Odontogriphidae, Cambroclavida and Dinomischus. Also absent are the (arrow-worms), which have been considered the basalmost branch of the Deuterostomia in the past, but are currently regarded by many as basal protostomes (Helfenbein et al., 2004).

© Christopher Taylor, CT050815

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One feature of the deuterostomes deserves a closer look - the widespread occurrence across the clade of deviations from the bilateral symmetry found in most Bilateria. Almost all the deuterostomes are asymmetrical to some degree, and in echinoderms bilateral symmetry has been lost entirely in the adult and replaced by pentaradial symmetry.

In vertebrates, left-right asymmetry is mostly found in the viscera- we're probably all aware that our heart and digestive organs are not positioned directly down the midline, but are coiled and directed to one side or another depending on the organ. Other features, such as the skeleton, are more or less symmetrical.

The immediate outgroup of craniates, the (e.g., amphioxus), show strong asymmetry during development. From the appearance of the fifth somite, the anterior borders of the somites move visibly out of register. Minguillon & Garcia-Fernandez (2002); Boorman & Shimeld (2002). After neurulation, paired cavities called Hatschek's diverticula (possible pituitary homologue) form in the head endoderm. The left cavity moves posteriorly to form the pre-oral pit, while the right expands to fill the vacated space in the head. The larval head becomes significantly asymmetrical, with the mouth appearing on the left side and the gill slits (originating on the ventral midline) spreading up to right side only. At metamorphosis, the mouth moves to the front, the pre-existing gill slits move from the right side to the left, and new gill slits open on the right. The adult therefore regains pharyngeal symmetry. Other asymmetries remain, such as the displaced muscle blocks, the position of Hatschek's pit (an organ derived from the pre-oral pit, and probably homologous to the pituitary gland) on the left side, and a blind gut diverticulum on the right (Boorman & Shimeld, 2002).

, the next group out, show little asymmetry in the larva, except in the positions of the two sensory pigment spots and adjoining regions of the brain. In the adult, the gut becomes asymmetrically folded.

The hemichordates are the most symmetrical of the deuterostomes. The gut is coiled asymmetrically but unlike other deuterostomes the direction of asymmetry differs between individuals. Directional asymmetries are few and relatively small - the protocoel pore is typically on the left. Boorman & Shimeld (2002). Rhabdopleura is something of an exception - in this genus, the right feeding arm is usually longer than the left, the single gonad is on the right, the mouth is on the left and the anus is on the right. Jefferies (1986).

In echinoderms the larva is initially bilaterally symmetrical, only to be replaced by a whole new order of symmetry in the adult. In sea urchins the pentaradial adult body plan is laid down in a structure called the adult rudiment, which lies on the left within the larva. Boorman & Shimeld (2002); Arenas-Mena et al. (2000). In and , the left somatocoel becomes oral and the right one aboral. In starfish, only the left hydrocoel of the larva develops into the adult water-vascular system, while the right hydrocoel disappears. Jefferies (1986). A lot of debate still occurs about the appropriate interpretation of echinoderm axes, and how (if at all) they can be compared to axes in other Bilateria. It is uncertain, for instance, if the anterior-posterior axis in other animals corresponds to the oral-aboral axis of echinoderms or the central axes of the five rays. The fossil lineage of echinoderms suggests some very odd evolutionary stages, with representing a strongly asymmetrical stage in echinoderm ancestry (if they are indeed stem echinoderms - see later).

The obvious question then becomes what this all means. Why are deuterostomes such individualists? Jefferies (e.g., 1986) has suggested that the asymmetry of chordates and echinoderms is homologous, and evidence that they form a monophylum to the exclusion of the more symmetrical hemichordates. This clade he named Dexiothetica after his suggested explanation for its origin - that the ancestor of the echinoderm-chordate clade underwent a 90 degree rotation so that it came to lie on its right side with the left side becoming dorsal (Greek dexios = right, thetikos = suitable for lying down). In chordates, the outer body subsequently re-oriented itself over time to become bilaterally symmetrical, but the viscera remained dexiothetic.

Jefferies' theory was strongly dependent on his interpretation of the fossil homalozoans (which he referred to as calcichordates - see later). On this page, we take the more orthodox view that homalozoans are stem echinoderms, as suggested by their possession of a calcite skeleton with a similar ultrastructure to modern echinoderms. Jefferies held this skeleton to be ancestral for Dexiothetica, and subsequently independently lost in each of the three chordate subphyla. However, this seems to require a lot of convergence, expecially if, as molecular phylogenies suggest, the hemichordates are actually closer to echinoderms. A stem echinoderm position for homalozoans seems much more parsimonious.

Could dexiothetism have still been ancestral to deuterostomes as a whole? The theory does have a certain appeal in relation to echinoderms - as mentioned before, left and right do become dorsal and ventral in echinoderm development, and the unpaired nature of the gill structures in and at the base of the echinoderm stem offers further support (if they are correctly interpreted as such - see later). In the case of Chordata, however, if the homalozoans are not regarded as relevant to the chordate stem, the support for dexiothetism is much lower. Asymmetries in the viscera may have been an adaptation for greater complexity (a coiled heart has greater pumping ability, while a coiled gut allows for more surface area in a smaller volume. Cooke (2004). The strong asymmetry in amphioxus may be an autapomorphy of that particular lineage. Evidence is therefore equivocal as to whether dexiothetism can be regarded as ancestral for Deuterostomia, or whether chordate and echinoderm asymmetries arose independently. Among fossils that may lie along the stem group of deuterostomes, the Vetulicolia have not been recorded as showing any sign of asymmetry. However, the mostly lateral, two-dimensional preservation of vetulicolians means that this is not necessarily informative.

Studies of developmental gene expressions and their relation to deuterostome asymmetry are still at a fairly early stage. The Nodal-Ptx pathway has been identified as a major factor in establishing the left-right axis in chordates. Ptx, for instance, is expressed predominantly on the left-hand side of the embryo in all three chordate subphyla. However, neither Pitx or Nodal have yet been identified in echinoderms, though once again the extremely derived body plan of echinoderms makes the identification of gene functional homologies difficult. Wray & Lowe (2000). Preliminary analysis suggests Ptx expression is symmetrical in hemichordates, but this requires confirmation. Boorman & Shimeld (2002).

In succeeding pages, we will explore some of the strange fruits that have blossomed on the Deuterostome stem. For now, we include some other, quite different, versions of Deuterostome phylogent for comparison.

© Christopher Taylor, CT050821.

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<==o Deuterostomia 
   |?- †Vetulicolia Shu et al. (2001)
   |?- †Mitrata[Calcichordata; Stylophora; Carpoidea] (mitrates)
   |--o "ECHINODERMOMORPHA"
   |  |--+-- †Cambroclaves
   |  |  `--+-- †Heterostelea
   |  |     `-- Echinodermata 
   |  `--+-- †Dinomischida
   |     `--+-- †Eldoniida
   |        `--+-- †Rotadiscida
   |           `-- †Paropsonemida
   `--o "CHORDATAMORPHA"
      |-- †Graptolitha [Stomochordain partim] (graptolites)
      `--o "CHORDATAFORMES"
         |-- Hemichordata [Stomochordain partim]  [paraphyletic?]
         `-- CHORDATA 

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<==o Deuterostomia 
   |-- Phoronida
   `--+-- Brachiopoda
      `--o DEXIOTHETICA
         |?- †Vetulicolia Shu et al. (2001)
         |?- †Mitrata[Calcichordata; Stylophora; Carpoidea] (mitrates)
         |--o "ECHINODERMOMORPHA"
         |  |--+-- †Cambroclaves
         |  |  `--+-- †Heterostelea
         |  |     `-- Echinodermata 
         |  `--+-- †DINOMISCHIDA
         |     `--+-- †Eldoniida
         |        `--+-- †Rotadiscida
         |           `-- †Paropsonemida
         `--o "CHORDATAMORPHA"
            |-- †Graptolitha [Stomochordain partim] 
            `--o "CHORDATAFORMES"
               |-- Planctosphaeroidea
               |--o Pterobranchiata [Pterobranchia] 
               |  |-- Atubaridae
               |  |-- Rhabdopleuridae [Rhapdopleurida]
               |  `--o Cephalodiscida
               |     |-- †Eocephalodiscidae
               |     `-- Cephalodiscidae
               `--+--o Enteropneusta
                  |  |-- Protoglossidae
                  |  |-- Harrimanidae
                  |  |-- Spengalidae
                  |  `-- Ptychoderidae
                  `-- CHORDATA 


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Arenas-Mena, C, AR Cameron & EH Davidson (2000), Spatial expression of Hoxcluster genes in the ontogeny of a sea urchin. Development 127: 4631-4643.

Boorman, CJ & SM Shimeld (2002), The evolution of left-right asymmetry in chordates. BioEssays 24: 1004-1011.

Cameron, CB (2005), A phylogeny of the hemichordates based on morphological characters. Can. J. Zool. 83: 196–215.

Cameron, CB, JR Garey & BJ Swalla (2000), Evolution of the chordate body plan: New insights from phylogenetic analyses of deuterostome phyla. Proc. Nat. Acad. Sci. (USA) 97: 4469-4474.

Cooke, J (2004), The evolutionary origins and significance of vertebrate left-right organisation. BioEssays 26: 413-421.

Davidson, EH, KJ Peterson & RA Cameron (1995), Origin of bilaterian body plans: Evolution of developmental regulatory mechanisms. Science, 270: 1319-1325.

Giribet G, & C Ribera (1998), The position of arthropods in the animal kingdom: a search for a reliable outgroup for internal arthropod phylogeny. Mol Phylogen. Evol. 9: 481-488.

Heinzeller, T & U Welsch (1999), The complex of notochord / neural plate in chordates and the complex of hydrocoel / ectoneural cord in echinoderms - analogous or homologous? in MD Candia Carnevalli & F Bonasoro [eds.], Echinoderm Research 1998. AA Balkema, pp. 285-290.

Helfenbein, KG, HM Fourcade, RG Vanjani & JL Boore (2004), The mitochondrial genome of Paraspadella gotoi is highly reduced and reveals that chaetognaths are a sister group to protostomes. Proc. Nat. Acad. Sci. (USA) 101: 10639-10643.

Jefferies, RPS (1986), The Ancestry of the Vertebrates. Brit. Mus. (Nat. Hist.).

Lacalli, TC, THJ Gilmour & SJ Kelly (1999), The oral nerve plexus in amphioxus larvae: function, cell types and phylogenetic significance, Proc. R. Soc. Lond. B 266: 1461-1470.

Maddison, DR (2002), , in (acc. 050815).

Miller, SA & JP Harley (1996), Zoology [3rd ed.] Wm. C. Brown Publishers.

Minguillon, C & J Garcia-Fernandez (2002), The single amphioxus Mox gene: Insights into the functional evolution of Mox genes, somites, and the asymmetry of amphioxus somitogenesis. Devel. Biol. 246: 455-465.

Peterson, KJ & DJ Eernisse (2001), Animal phylogeny and the ancestry of bilaterians: inferences from morphology and 18S rDNA gene sequences. Evol. Devel. 3: 170-205.

Shu, D-G, S Conway Morris, J Han, L Chen, X-L Zhang, Z-F Zhang, H-Q Liu, Y Li, & J-N Liu (2001), Primitive deuterostomes from the Chengjiang Lagerstätte (Lower Cambrian, China). Nature 414: 419-424.

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Hypertension (high blood pressure): Patients with essential hypertension or arteriosclerosis have an increased risk of developing erectile dysfunction. Essential hypertension is the most common form of hypertension; it is called essential hypertension because it is not caused by another disease, (for example, by kidney disease). It is not clearly known how essential hypertension causes erectile dysfunction; however, patients with essential hypertension have been found to have low production of nitric oxide by the arteries of the body, including the arteries in the penis. Scientists now suspect that the decreased levels of nitric oxide in patients with essential hypertension may contribute to erectile dysfunction. For more information, please read the High Blood Pressure article. Objective: To analyze the prevalence and risk factors for erectile dysfunction (ED) in Italy in a cross-sectional study. Hormone therapy can raise low testosterone levels, which may improve both erections and sex drive. Cannot be used by patients on MAOIs* When did you first notice symptoms of ED? Because there are a variety of causes for ED, there are several different tests your doctor may use to diagnose the condition and determine its cause. Only after the cause of ED is determined can it be effectively treated. Tadalafil (Cialis) CIALIS Yes. Condoms aren't expensive and are widely available. Condoms are an especially good choice if you or your partner are also having sex with other people or if either of you has had sex with other people in the past. Erectile dysfunction. A number of devices have been developed to ascertain the occurrence of an erection during sleep. This snap gauge is fastened around the penis but opens when an erection occurs. Alprostadil can be injected into the penis or inserted using a special applicator - usually just before sexual intercourse. PMS and Sex: More Ways to Get in the Mood Yes. It's important to identify and treat any correctable problems. In some men, a doctor's exam may find an underlying medical problem that is causing the infertility. buy propecia online There is no single test that can diagnose depression; however, there are certain patterns that doctors look for in order to make the diagnosis. As a result, your doctor will ask you several questions. Be honest with your answers so that you can receive the care you need. Arthritis was the most common reason for NSAID use, and for men with arthritis the corresponding rates of erectile disorder were 97 and 52 per 1000 person-years. cholesterol, smoking, high blood pressure -- doesn't improve the prediction of future cardiovascular trouble, it can be a quick, free addition to risk assessment, Araujo said. Hypogonadism (which leads to lower testosterone levels)

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