!"#$%&'$()*+,-)$.

/$01-213)$4516,713$
Origin of Life: Synthesis of Prebiotic Chemicals
hormones were part of the earth(?)

Extraterrestrial Origins:
Meteorites with organic material PAH, amino and organic acids, ketones Alcohols, purines, C10

Terrestrial Origins:
Amino acids, sugars, purines from CH4, NH3, H2O, N2

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Figure 2-2: History of the evolution of the Earth and approximate times of the major stages in the evolution of life.

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Physiological Requirements: Internal Conditions -energy -raw materials Environmental Conditions: External conditions -Acceptable conditions-growth/metabolism -Poor conditions-quiescence When cells can coordinate these signals, they begin to live within a community either singly or communal and begin to establish a division of labour and therefore specialization of function
might have created multicell orgs through endocrine sys

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Theories of Prebiotic Synthesis: Energy Sources
There were, in theory, several sources of energy that could have been used for the prebiotic synthesis of organic chemicals

Sunlight

Geothermal vents

Lightning

Volcanic activity

Cosmic Radiation

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NH4

Figure 2-1: Examples of organic compounds present in the prebiotic Earth. Such compounds were used as substrates and building blocks for more complex molecules.

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Association of organic material on inorganic substrate Concentration of organic material
A

Evolution of Protocells

Partitioning of Hydrophilic and Hydrophobic material Formation of Hydrophobic membrane Separation of protocell From substrate
there was an E gradient

!"#$%&'$()*+,-)$./$01-213)$4516,713$

!"#$%&'$()*+,-)$./$01-213)$4516,713$
Protocells and First Signalling Networks
What is a protocell? Partitioning of biotic and non-biotic environments The need to signal Theoretical evolution and structure of a protocell Partitioning of hydrophilic and hydrophobic molecules Development of lipid membrane Ion gradients across membrane and energy production Signalling capacity Membrane diffusion Pores in membrane Protein channels Early adhesion proteins

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Protocells, as we examined had only a random and intermittent signaling capacity with little, if any, coordination between the excreted Molecule and the regulation of activity of another protocell. Once viable living cells developed, coordination between the secreted Molecule and the activity of the target cell was essential.

The expression of receptors on the target cell determines the specificity of hormone action

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First Cells and Signaling Ability
First true cell Metabolic and genetic replication ability Required Attributes Stable membrane Cytoskeleton adhesion signaling system endocytosis, exocytosis Reasonably accurate replication Evolution 2 billion years from first cell to first multicellular animal Elaboration of membranes act as a substrate for biosynthetic enzymes required for exocytosis and endocytosis Symbiosis with other organisms Transformation from anaerobic to aerobic formation of organelles

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Symbiotic cells increase of complexity

Figure 2-11: Increase in cellular complexity by symbiosis. The formation of organelles occurs by the ingestion or infection of other cells. Overtime, a symbiotic relationship is formed. Intracellular or intracrine signalling pathways are required to coordinate the actions of the new organelle with the actions of the rest of the cell.

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Selection of Types of Signaling Molecules
Amino acids among first amino acids synthesized acid/base ability form elongated chains Peptides and Proteins structural and signalling ability several types of structural organization can carry and transmit information Lipids pass through membrane early prebiotic synthesis Gases and ions present in prebiotic Earth Nucleic acids common as signal molecules but evolutionary origins are not clear Sugars impart a lot of E and are hydrophillic might not be great not common as signalling molecule sog mols because they dissolve

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A possible timeline for the evolution of cells and their signalling systems

Figure 2-15: Evolution of signalling systems in the Metazoa.

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Problems associated with the development Of multicellularity:
1.! Adhesion;cells need to be physically connected or attached 2. Nutrient, ion, water, oxygen transport to inner cells 3. Coordination of function: signalling systems 4. Procurement of nutrient sources

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Signaling Processes

Earliest Cells: Intracrine and Exocrine abilities More evolved systems Autocrine/Paracrine mechanisms Juxtacrine Mechanisms Membrane Mechanisms Simple and facilitated diffusion Channel and transporter mediated diffusion

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Basic signaling systems:
Growth and Differentiation feeding, growth maturation cell cycle differentiation cell death Sensory systems locomotion toward food source away from danger chemical (olfactory input) visual (input) geotaxis mechanical

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know how the signalling systems evolved
us

mollusks and insects

know the taxonomy

second mouth = opening

distinct organisms evolved from single celled orgs

no symetry bet ween sponges and placozoans

basal orgs = something that's close to the original

Figure 3-8: Phylogeny of the phylogenetically oldest lineages of metazoans.

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"8296):+$;31<3$=)+>?1>3$
the only signalling they have might be paracrine signalling. ionic transmission like gap junctions. simple signalling system doesn't require much sophistication to communicate and stuff

Trichoplax adherens Phylum Placozoa No endocrine or nervous system asymmetrical

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Signaling in primitive Extant Metazoans

Placozoa and Poriferans as models of early metazoans role of extracellular matrix Trichoplax adherens as a model asymmetrical, two tissue types autocrine paracrine signalling only

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Epithelial cells of the Poriferans (Sponges) are capable of transmitting signals

No true nervous system with axons and action potentials

works the same way as placozoans. so don't know if they're sister types or how they're related. they're negatively phototaxic, take the lar va are symetrical, their photoreceptors can stimulate the surrounding cells by gap junctions which can stimulate the flagella moving it away from light. it's capable of behavior despite the lack of NS

Figure 3-10 Photoreception in a sponge larvae. Cilia, associated with a ring of photoreceptive cells at the base of the larve act to propel the organism away from sunlight. Reprinted with permission from Leys and Degnan (2001) Biol. Bull.

!"#$%&'$()*+,-)$./$01-213)$4516,713$
have a reasonably elaborate NS, there's no transitional forms doe, don't know how it got to that point. but they go have a ner ve net, which is fundamentaly different from how we do it. looks like the next slide

Formation of the Nerve net with Neurons and neurosecretory cells

Placozoans and poriferans do not have nervous systems, however the Radiate phyla, Ctenophora and Cnidaria do. There are no known metazoans that display transitional forms of the nervous System. Phylogenetically older species of deuterostomes and protostomes also show The presence of a nerve net, We assume that the first functional neural/neurosecretory system was a nerve net.

!"#$%&'$()*+,-)$./$01-213)$4516,713$
blue = somatic cells, the other cells are neurosecretory cells, which are connected by axons and are in communication with each other, they get stimulate and interact with each other. can efficiently regulate the somatic cells

A model of a nerve net

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Immunohistochemical visualization of a Nerve net in Hydra

Figure 3-13 Nerve net in Hydra. A network of nerve cells is seen in A) hypostome B) peduncle and C) basal disc after visualization with an anti vasopressin antiserum. See text for additional details. The scale bars represent 100 m. Reprinted with permission from Morashita et al (2003) Elsevier Ltd.

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Signaling Systems in the Radiata
Growth and Differentiation pathways comparatively well developed
frizzled = GPCR and is a very early version of it.

Wnt/frizzled pathway: wnt (wingless) ligands: a family of polypeptide ligands frizzled receptor: a family of proteins consisting of membrane bound receptors with 7 alpha helices Similar to G-protein coupled receptors but without the G-protein component. Nuclear receptors: similar to steroid-thyroid hormone receptors although ligands are not well understood Serine/threonine and Tyrosine kinase based systems Sensory type pathways present G-protein coupled receptors are present but in low numbers and diversity

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Formation of the Nerve net with Neurons and neurosecretory cells
The first neuron/neurosecretory cells may have been derived from Early sensory cells These cells released a chemical signal via a robust depolarizing current resulting from the appropriate sensory input. Later these cells evolved into a new morphology with extended Process that could interact with structures associated with movement And feeding (ie. Cells with cilia). As movement became more complex, interneurons evolved and bridged The sensory cells with the locomotory cells

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Evolution of Signaling pathways
pre ner vous type of system.

Sensory input Sensory cell Ionic coupling

Locomotor cell with cilia

Nervous

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Evolution of Signaling pathways
Sensory input Sensory cell Ionic coupling Sensory input Sensory cell Chemical secretion and paracrine diffusion

Locomotor cell with cilia

Nervous

Neuroendocrine

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primitive NS

Evolution of Signaling pathways

primitive

primitive

Sensory input Sensory cell Ionic coupling

these t wo split and specialized

Sensory input Sensory cell Chemical secretion and paracrine diffusion

Sensory input Sensory cell Chemical secretion and paracrine diffusion Non-nervous (depolarizing) Secretory cell

Locomotor cell with cilia

Nervous

Neuroendocrine

Endocrine

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Genetic and Morphological Complexity and Signalling Systems: Key points
endocrine and NS developed complementary systems b/c NS = fast but can only reach few cells and endo = slow but reaches all cells .

Development of Nervous and Vascular Systems Integration of sensory systems and locomotor systems Formation of nervous system and integration with vascular system Triploblastic organism Formation of all known bilateral metazoans Increase in complexity of signalling systems

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Relationship among symmetry, endocrine system, and the nervous system
not the most efficent way of moving

How an organism moves has a bearing On where sensory organs are located. Once bilateral symmetry evolved, then Sense organs could be concentrated In the direction of movement (i.e. head)
more efficient, looks like a pnacake and move in t wo dimensions, sensory sys can be located on the edge, the NS linked all fo these things they have radial NS.

most efficient. they move along one direction. the sensory organs become concentrated on one secion. the amount of E to create a small but efficent it less than the first one. the amount of E utlized for movement is also less compared to the other ones.

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Relationship among symmetry, endocrine system, and the nervous system

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Symmetry allowed for structural similarity among individuals of a population A systemic signaling system could therefore develop in same way in all individuals. more effective. it also creates a behaviour which allows them to interact with each other even more and they
come together as a population

This set the foundation for co-ordination among sensory systems, feeding, and locomotory structures. This could not be consistently achieved in a non symmetrical animal.

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pangenomic theory that somehow the genomes weere similar (?) find more info

Precambrian explosion Arthropods Echinoderms Ecdysozoa Protostomes Molluscs Lophotrochozoa Acoela Radiata Placozoa Triploblastic animals

Chordates

Deuterostomes

Basal (phylogenetically older) Lineages of bilateral animals Porifera

Moving forward toward the Chordates

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Formation of the Chordate Nervous System Plan
Segmented nervous System with Ganglia and Ventral nerve chord Segmentation Encephalization Dorsal nerve chord
protestomes

bilateral animals. we see the beginning of a NS

Ventral nerve chord Radiates: Nerve ring Nerve net

@8A,-)$.B&$"#$%&'$(!$)!*+,!(,-#$&.!./.*,0!

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A few words about Endocrine systems
Any cell that evolved the capability for robust paracrine secretion had the potential to become and endocrine cell. These cells were present in all the basic tissue types of the earliest Metazoans, therefore, all tissues had the capability of becoming Endocrine organs. In fact, this is the case today: all tissues and organs have a variety of Substances that are secreted into the blood stream.

Therefore, all organs, by definition, may be considered endocrine organs.

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Nervous-Endocrine Interactions
Neurotransmitters Neuromodulators

Sensory Information

Nervous component Neuroendocrine component

Neurotransmitters Neuromodulators
mostly peptides

are small mobile molecules

Tissues
Hormones

Endocrine component

Releasing Factors Neurohormones they cover everything

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General Organization of the Vertebrate Brain

Nerve cord/spinal cord

shark brain

Hindbrain rhombencephalon Midbrain mesencephalon Forebrain prosencephalon

The brain is bilateral consisting of both left and right halves. One side is basically a mirror image of the other. This means also that the neuroendocrine cells are similarly divided.

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Human brain development
midbrain

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most is on diencephalon main component

Figure 4-13 Organization of the vertebrate nervous system A) Central Nervous System B) Spinal Cord and ganglia

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Ventricles of the Brain
The brain and nerve cord are hollow and consists of four ventricles filled with cerebral spinal fluid

fishes, sharks, amphibians

I II

III

IV

Forebrain

Midbrain

Hindbrain

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evolution, folded back on its self

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Cerebrospinal fluid CSF
Resembles ultrafiltrate of plasma There are no substances in CSF that are not found in plasma Proportions differ No diffusion barrier
protects brain from shock and stabilizing it and also clears out shit form there

I II

III

IV

There is a barrier between CSF and blood.

Brain-blood barier

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Blood-brain barrier

can expand and contract to allow openings

Figure 4-15 Basic components of the vertebrate blood brain barrier

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ones that are fat soluble they pass through BBB very easily which the water soluable can't

windows or holes

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Blood-brain barrier
Passive diffusion: water, lipophilic (steroids, ecosanoids) drugs (barbiturates) alcohol Transported: glucose, neutral amino acids used for neurotransmiter synthesis Peptides such as met-enkephalin, IL1, NPY, MCH, CRF use saturatable Transporters Peptides such as calcitonin, amylin, adrenomedulin enter via specific receptors In circumventricular organs

!"#$%&'$()*+,-)$./$01-213)$4516,713$
Pineal gland melatonin, rhythms Subfornical organ regulates fluids, osmoreceptor cells Organum vasculosum peptides, osmoregulation, fever Area postrema vomiting centre, osmoregulation Median eminence associated with anterior pituitary Neurohypophysis: neurosecretion AVP, oxytocin
Figure 4-16. Position of circumventricular organs in a human brain

!"#$%&'$()*+,-)$./$01-213)$4516,713$
1,(,-2%!3*-&4*&-,!$)!56*&6*2-/!1%2(7!

rostral front

III ventricle
Hypothalamus

caudal

Optic chiasm

Median eminence

The pituitary gland consists of two main lobes: Anterior Lobe:epithelial tissue Posterior Lobe: nervous tissue

Intermediate lobe (Pars intermedia)

Anterior lobe Anterior Hypophysis (Pars distalis)

Posterior lobe Posterior Hypophysis (Pars nervosa)

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not on the exam

Mammalian Brain Development

Neural plate Neural tube Ectoderm Neural crest

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not on the exam

Ontogeny of the Pituitary Gland in Amphibians

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Structure of Human Hypothalamic-Pituitary Portal System

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Hormones Produced in the Pituitary Gland
Anterior Pituitary Gland Cell Type: Gonadotropes: Follicle Stimulating Hormone (FSH) Luteinizing hormone (LH) Thyrotropes: Thyroxine stimulating hormone (TSH) Corticotropes: Adenocorticotropic Hormone (ACTH) Somatotropes: Growth hormone (GH) Lactotropes: Prolactin Somatolactotropes: Somatolactin (fishes) Posterior Pituitary Gland The posterior lobe consists of neurosecretory terminal originating from cells in the hypothalamus. In mammals, there are two main hormones that are released: Vasopressin and Oxytocin. In non-mammals variants of vasotocin are released.

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Chordate Evolution and Phylogeny

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Neuroendocrine secretion in a Tunicate

Figure 4-18. Neural Complex in a Tunicate. A number of hypothalamic and pituitary hormones and processing enzymes have been found in the neural complex suggesting a relationship with the chordate hypothalamo-pituitary system. See text for further details.CS, cloacal siphon; OS, oral siphon. Reprinted with permission from Kawamura et al, (2002) Elsevier Science (USA).

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Structure of Agnathan Pituitary Glands
The jawless fishes, or agnathans, are the most primitive known chordates. These fish which include the hagfishes (Myxini) and the lampreys (Cephalospidomorphi) are separate classes. The hagfishes lack a vertebrae and therefore are not classified As vertebrates. The Cephalospidomorphi do possess a vertebrae and are considered The phylogenetically oldest vertebrates. The Cephalospidomorphi likely evolved At least 50 million years after the Myxini. Both groups of fishes are highly degenerate. Despite this, there are common aspects to their structure of pituitary and hypothalamus That provide us with clues as to how this system evolved in the vertebrates
1 0 0

Hagfish Class: Myxini

2 0 0 3 0 0 4 0 0 5 0 0

6 0 0 Lamprey Class: Cephalospidomorphi

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Structure of Agnathan Pituitary Glands
both don't have cortical sys

For both the Myxini and the Cephalospidomorphi there appears to be a complete separation of the pars distalis from the rest of the hypothalamus. Diffusion of the hypothalamic releasing factors appears to be the main route of communication between the hypothalamus and pars distalis. However, a simple portal system may be present although this is unclear. In hagfishes, (Myxini), the pars nervosa is a relatively undifferentiated part of the hypothalamus where the axonal termini of neurosecretory cells impinge on a vascularized region at the hypothalamus wall. In lampreys (Cephalospidomorphi) the pars nervosa becomes more differentiated into a discrete lobular structure.

Agnathan Pituitary Structure

much closer association with distalisn and ner vosa

communicate through stemic bloods not efficent but it works

Myxini
Pars distalis Pars intermedia

Cephalospidomorphi
Pars nervosa

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Structure of Agnathan Pituitary Glands
Hagfish and lampreys may also utilize the systemic blood as a route for releasing factors to regulate the pars distalis Ectopic transplantation of the hagfish pars distalis in other regions of the body, for example do not appear to compromise the normal physiology of the these animals. The various cell types in the pars distalis are relatively intermingled.

!"#$%&'$()*+,-)$./$01-213)$4516,713$
Chondricthyan Pituitary Gland
The Class Chondrichthyes or cartilaginous fish are the phylogenetically oldest Group of gnathostomes or jawed fish. The cartilaginous fish include the Subclass Elasmobranchii (sharks, skates and rays) and the Subclass Holocephali (chimaeras). The cartilaginous fish evolved 50-100 million years After the lineage leading to the lampreys and about 400 million years before Present. The Elasmobranchs and Holocephali diverged early in Chondrichthyan evolution, but still possess a number of related features.

1 0 0 2 0 0 3 0 0 4 0 0 5 0 0 6 0 0

Elasmobranchii

Holocephali

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General Structure of Elasmobranch Pituitary Gland
The Elasmobranch pituitary consists of a fused pars nervosa and pars intermedia. In this situation where there is considerable Intermingling of pars nervosa and pars Intermedia, it is referred to as a pars Neurointermedia or Neurointermediate lobe. There is a well developed portal system that connects most of the pars distalis with the hypothalamus. However, the ventral lobe, has no vascular or nervous connection to the rest of the brain. Releasing factors are secreted into the systemic blood supply to communicate with the ventral lobe. Neurointermediate lobe

Portal system

Pars distalis Pars intermedia Pars nervosa

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Elasmobranch Pituitary Organization
The elasmobranch pars distalis consists of a second lobe ventral to the main Body. This structure is called the ventral lobe and is connected to the rest of The pituitary by a thin stalk of tissue. The ventral lobe does not appear to have Either a vascular or nervous connection to the rest of the brain.
Pars Nervosa Pars Distalis

Portal System

Ventral lobe

A view of the elasmobranch pituitary gland

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Pituitary and Diencephalon Structure in the Holocephali
The structure of the pituitary in holocephalans is similar to that of the Elasmobranchii. The main difference lies in the structure of the ventral lobe. In holocephalans, the ventral lobe is called a buccal lobe and is found in the roof of the mouth. It is separated from the brain by a layer of cartilage. The hypothalamus communicates with the buccal lobe by secreting releasing factors into the systemic blood.

Brain

Buccal lobe

Position of Pituitary Gland

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Pituitary Cell Organization in the Cartilaginous Fish

interdigitation

Although there are few studies to draw upon, the location of cells in the pituitary gland in the Cartilaginous fish appears to have some organization. For example, the gonadotropes are located primarily in the ventral lobe of elasmobranches (or the buccal lobe of holocephalons). Although there is some interdigitation of the pars nervosa with aspects of the pars distalis, there is currently no evidence that there is any direct neural link between the hypothalamus and pars distalis.However, given the organization of the neurosecretory cells of the hypothalamus and the presence of interdigitation in cartilaginous fish, there is the distinct potential that some form of rudimentary neural connection may exist. If so, one might also expect that some of the basic pituitary cell types are organized to a limited degree in clusters of related cells.

!"#$%&'$()*+,-)$./$01-213)$4516,713$
The Actinopterygian-Sarcopterygian Bifurcation And Pituitary Evolution
Around 350 million years ago, vertebrate evolution followed two main pathways. One group of fishes exploited a niche associated with the ocean and lake bottoms whereas another group exploited a pelagic niche. The bottom dwelling group adapted robust and limb-like 1 0 0 fins and developed into the lobe-finned 2 0 0 fishes (Sarcopterygii) and ultimately to land animals. The pelagic adapted 3 0 0 lineage developed into the ray-finned 4 0 0 Fishes (Actinopterygii). Pituitary evolution followed separate 0 0 morphological patterns in each group. 6
5 0 0
amphibians reptiles us at the bottom

teleosts

Lobe-finned fish lineage (Sarcopterygii)

Ray-finned fish lineage (Actinopterygii)

!"#$%&'$()*+,-)$./$01-213)$4516,713$
4516,713$1C$+D)$E8+,8+>-F$E1-+>6$"F:+)2$
Neurosecretory cells With projections to The median eminence And portal system

Early jawed vertebrate vIII oc

Neurosecretory cells With projections to the pars nervosa

Lobe-finned fishes/tetrapods vIII oc

Ray-finned fishes vIII oc

Jawed vertebrates began with a pituitary structure similar to chondrichtyan fishes where there was a portal system and a more or less distinct pars distalis and pars nervosa. In the lobe-finned fishes and tetrapods, the portal system became more developed, whereas in the ray-finned fishes the portal system was lost and a direct neural connection developed

Elaboration of portal system Direct neural connection

cells are intermingled here
cells are clumped by groups

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Amniotes
The earliest amniote tetrapods appears about 30 million years after the evolution of the first amphibians. This group includes the mammals, reptiles and birds, and refers to the amniotic egg present in all groups. The early amniotes began to radiate extensively in the Carboniferous and early Permian times, displacing the non-amniotes (early amphibians) from those habitats and niches. Evidence suggested that virtually all of the early amniotes were carnivorous. The extensive radiation of insects at this time is thought to have been a food source that was exploited by the early amniotes thus acting to stimulate this radiation of species. Physiological changes that accompanied this adaptation included changes in respiratory mechanisms, diuresis and excretion, reproduction and integration of sensory systems.
1 0 0

Phylogenetic Position of Amniotes

2 0 0 3 0 0 4 0 0 5 0 0 6 0 0

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Structure of Reptile Pituitary Glands
There are four subclasses of reptiles: Chelonia (tortoises, turtles and terrapins), Rhynchocephalia (tuataras), crocodilia (alligators, crocodiles, caimans, gharials) And squamata (lizards snakes).
chelonia

1 0 0 2 0 0
rhynchocephalia crocodilia

3 0 0 4 0 0 5 0 0

squamata

6 0 0
squamata

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Structure of Reptile Pituitary Glands
The general arrangement of the reptilian pituitary is more or less similar in most of the groups. Differences are most apparent within the pars nervosa morphology. The phylogenetically older reptile groups such as the chelonia (turtles) and rhynchocephalians (tuataras) tend to have a pars nervosa with some interdigitation of the pars intermedia. Note also in all reptiles is an expanded portal system and generally a mix of pituitary cell types instead of clusters of similar cells.
similar to mammals

Rhynchocephalia

Chelonia

Crocodilia

Squamata (lacertilia) (lizards)

Squamata (ophidia) (snakes)

!"#$%&'$()*+,-)$./$01-213)$4516,713$
Structure of Teleost Pituitary Gland
The Teleostei are a vast assemblage of fish that include about 25,000 species. The pituitary gland is unusual in that cells of the pars distalis are clustered in nuclei of similar cells. There is extensive interdigitation of the neural lobe with the rest of the pituitary, and direct neural connection between the hypothalamus And pars distalis
1 0 0 2 0 0 3 0 0 4 0 0 5 0 0 6 0 0

!"#$%&'$()*+,-)$./$01-213)$4516,713$
Anguilla anguilla (eel) as a model for the teleost pituitary gland
rostral
Neurosecretory cells of the hypothalamus

OC

The teleost pituitary gland morphology represents the most advanced structural features among the fishes. Two major changes have occurred: neurosecretory cells of the hypothalamus send their axons through the extensive interdigitations of the pars nervosa with the pars intermedia and pars distalis. Within the pars distalis cells have become grouped into discrete clusters. Thus, in this manner the releasing factor can stimulate the appropriate pituitary cell.

!"#$%&'$()*+,-)$./$01-213)$4516,713$

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