The Drosophila melanogaster life cycle
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1 The Drosophila melanogaster life cycle
2 Eclosion The first phenotype described in Drosophila as an endogenous rhythm (1954): Number of eclosed flies Hamblen et al., Gene3cs 1998 rhythm with a periodicity of about 24h (circa die); evident in synchronizing condi3ons (e.g. Light:Dark cycles) and in free running condi3ons (constant darkness and T).
3 A circadian output: the locomotor activity rhythm LD 12/12 1 giorno LD 12/12 2 giorno Day 1 ZT 12 ZT 24 ZT 0 ZT 12 12:12 LD Day 2 LD 12/12 2 giorno DD 3 giorno DD Day3 DD 3 giorno DD 4 giorno Day 4 DD 4 giorno DD 5 giorno Day 5 DD 5 giorno DD 6 giorno
4 Model of the circadian oscillator timekeeper input pathway output pathway daily changes in physiological and behavioral processes
5 GeneBc basis of circadian clock in Drosophila In 1967, PiOendrigh demonstrated the hereditability of the character free- running period ; : Konopka and Benzer isolated Drosophila strains with eclosion- rhythm abnormali3es. Wild- type Locus period (X chr.) Number of eclosed flies Arrhythmic 19h periodicity 29h periodicity per 01 per s per l
6 period mutants - locomotor ac3vity /0 12h per h per o - arrhythmic per s 19 h per l 29 h
7 PER DOMAINS NLS PASA/B Thr-Gly NLS 1218 aa per L (Val Asp) per s (Ser Asn) per o (stop) CLD Chang et al., Curr.Biol. 2003
8 period mrna and protein cycle delay mrna PROTEIN
9 3meless mutant 3m 01 (Sehgal et al., Science 1994; Vosshall et al., Science 1994) Wild- type 3m 01 eclosion locomotor acbvity
10 TIM NLS CLD ARM- 1 ARM- 2 (Vodovar et al., Curr. Biol. 2002) timeless mrna and protein cycle delay mrna PROTEIN ( Myers et al., Science 1995; Sehgal et al., 1995, ibid,; Gekakis et al.,1995 ibid)
11 The circadian clock at molecular level
12 Clock mechanism in Drosophila and Mammals (Wijnen and Young, 2006)
13 How many clocks? Drosophila per-luciferase reporter
14 Brain master clock in Drosophila and Mammals Drosophila master clock Mouse SCN
15 The brain circadian clock Helfrich- Förster, Microscopy Research and Technique2003
16 Rhythmic behaviour requires brain intercellular communicabon
17 The output signals in the circadian behavior Ozkaya and Rosato, Adv Genet 2012
18 The output signals in the circadian behavior Prepro- PDF PDF 102 aa
19 PDF is a key factor in the output signal Renn et al. Cell, 1999
20 Anatomical and physiological outputs from neurons expressing clock genes in the brain of Drosophila Hall, Adv. Gen. 2003
21 Clock ReseVng by environmental sbmuli 1) Light 2) Temperature 3) Social input
22 Light synchronization Hall,2000
23 Molecular light entrainment (from Wijnen and Young, 2006)
24 Temperature input T information: perceived from periphery by the chordotonal organ, stretch of receptors located internally to the cuticle at the joints between segments (Sehadova et al., Neuron 2009).
25 Role of social experience in the entrainement of the circadian clock (Levine et al., Science 2002; Krupp et al., Curr Biol 2008) The locomotor activity peaks of flies living together are more synchronized than those of isolated individuals. Vectors represent the mean phase of the 2 groups; The tail length indicates the coherence of the phase: a longer tail denotes a tighter distribution of the phase
26 The influence of the social experience on the phase depends on the composition of the groups: Wild-type hosts: wild-type flies + per 01 flies (visitors) in different ratios
27 => The olfactory system is involved in the perception of the social input Sensory mechanism on the basis of the social effect: Analysis of the phase of two groups of isolated flies: one group during the period of the experiment received fly air ; the second group during the period of the experiment received neutral air. Analysis of the phase of olfactory para sbl mutants reared alone (control) or in presence of per01 mutants (hosts): No change in the coherence of the phase was caused by the presence of per01 mutants
28 The dual oscillator Most animals display a bimodal ac3vity, showing one peak in the morning and one in the evening. In the seven3es, PiOendrigh and Daan proposed a model explaining this bimodal ac3vity paoern: There is not only one circadian oscillator, but two. One oscillator the so called morning oscillator (M) is responsible for the ac3vity in the morning. The other oscillator the evening oscillator (E) is inducing ac3vity in the evening.
29 The neuronal control of the morning and evening ac3vity peaks in Drosophila melanogaster Stoleru et al Nature 2004; Grima et al. Nature 2004
30 The GAL4/UAS binary system (Brand and Perrimon, 1998) X UAS gene X Reg. Seq. A Gal4 P- Element P- Element F1 Reg. Seq. A Gal4 GAL4 GAL4 UAS gene X X
31 In yeast, the GAL80 protein antagonizes GAL4 ac3vity by binding to the ac3va3on domain of GAL4, preven3ng interac3on between GAL4 and the transcrip3onal machinery. GAL4 is inactive Galactose GAL3 GAL4 is active
32 Reg. seq.a Gal4 Reg. seq.a UAS Reg. seq.b Gal4 Gene X Gal80 GAL4 GAL4 UAS Reg. seq.b Gene X Gal80 GAL4 Reg. seq.a UAS Gal4 GeneX GAL80 GAL4 GAL80 Reg. seq.b Gal80
33 The morning and evening anticipations of the locomotor activity rhythm in 12:12 LD conditions are determined by the endogenous clock. Grima et al. Nature 2004 Which are the clock neurons important in the regulation of the morning and evening activity in LD?
34 The UAS/GAL4 binary and UAS/GAL4/GAL80 ternary systems allowed to identify the MA and EA neurons Pdf-Gal4>Uas-GFP Pdf-Gal4>Uas-hid NO MA The evening activity is controlled by additional neurons different from LNvs cry-gal4>uas-gfp cry-gal4>uas-hid NO MA and EA
35 The UAS/GAL4 binary and UAS/GAL4/GAL80 ternary systems allowed to identify the MA and EA neurons cry-gal4; Pdf-Gal80; UAS-GFP The GFP is expressed only in CRY+ PDF- cells cry-gal4; Pdf-Gal80; UAS-hid Ablation of CRY+ PDF- cells (LNd and 2DN1) MA, no EA
36 MA requires s-lnv and l-lnv (PDF+, CRY+) EA requires LNds, DN1s and 5th s-lnv (PDF- CRY+)!!! DN 1a DN 2!!!!! LPN!! I-LN v s-ln v DN 1p DN 3!!! LN d 5 th S-LN v! CRY+ CRY- (Stoleru et al Nature 2004; Grima et al. Nature 2004)
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