In yeast, mitochondrial division and fusion are highly regulated during growth, mating and sporulation, yet the mechanisms controlling these activities are unknown. Using a novel screen, we isolated mutants in which mitochondria lose their normal structure, and instead form a large network of interconnected tubules. These mutants, which appear defective in mitochondrial division, all carried mutations in DNM1, a dynamin-related protein that localizes to mitochondria. We also isolated mutants containing numerous mitochondrial fragments. These mutants were defective in FZO1, a gene previously shown to be required for mitochondrial fusion. Surprisingly, we found that in dnm1 fzo1 double mutants, normal mitochondrial shape is restored. Induction of Dnm1p expression in dnm1 fzo1 cells caused rapid fragmentation of mitochondria. We propose that dnm1 mutants are defective in the mitochondrial division, an activity antagonistic to fusion. Our results thus suggest that mitochondrial shape is normally controlled by a balance between division and fusion which requires Dnm1p and Fzo1p, respectively.
Mitochondria undergo regulated fusion and division in many cell types ( Bereiter-Hahn and Voth 1994; Kawano et al. 1995), which appear to play key roles in establishing and maintaining mitochondrial shape ( Tyler 1992). In the yeast S. cerevisiae, mitochondria are elongated, tubule-shaped organelles that are very dynamic during growth, mating, and sporulation. Mitochondria constitutively divide and fuse during cell growth ( Nunnari et al. 1997), but change their number depending on growth conditions ( Stevens 1977). During mating, mitochondria fuse immediately after cell fusion, mixing their contents, including mitochodrial DNA (mtDNA) and matrix proteins ( Nunnari et al. 1997; Okamoto et al. 1998). When diploids sporulate, mitochondria are dramatically reorganized moving into the four spores and surrounding each haploid nucleus ( Miyakawa et al. 1984). The yeast homologue of fuzzy onions ( Hales and Fuller 1997), FZO1 was recently identified and shown to play an important role in mitochondrial fusion ( Hermann et al. 1998; Rapaport et al. 1998). However, the mechanisms that control mitochondrial division are unknown.
Strain YHS2, which expresses red-shifted GFP (F64L, T65C, and I167T) fused to the Cox4p presequence (residues 1–21) under the ADH1 promoter (ADH1-COX4-GFP), was constructed as follows. First, pHS1 was constructed by replacing wild-type GFP in pOK29, a HIS3-CEN plasmid which carries ADH1-COX4-GFP (Kerscher, O., unpublished information), with a NcoI-BamHI fragment carrying red-shifted GFP from pQBI25 (Quantum Biotechnologies). The EcoRV-BamHI fragment from pHS1 was inserted into pDH9, which carries 5′ and 3′ untranslated regions of MFA2 (a gift from S. Michaelis), forming pHS2. To integrate ADH1-COX4-GFP at chromosomal MFA2, a XhoI-SmaI fragment carrying 5′-MFA2-ADH1-COX4-GFP-3′-MFA2 from pHS2 was transformed into strain SM1235, which carries mfa2::URA3 ( Michaelis and Herskowitz 1988). Strain YHS2, which contains MATα mfa2::ADH1-COX4-GFP, was selected on 5-fluoro-orotic acid medium ( Adams et al. 1997). MATa mfa2::ADH1-COX4-GFP strain YHS1 was obtained by crossing YHS2 to SM1227 ( Michaelis and Herskowitz 1988). Strain 1002 (MATa, his3, trp1, ura3, mfa2::ADH1-COX4-GFP) was constructed by crossing YHS1 to BY4731 ( Brachmann et al. 1998).
YHS2 was mutagenized with 3% ethane methylsulfonate to ∼30% survival ( Adams et al. 1997). Mutagenized cells were suspended at ∼9 × 105 cells/ml on coverslips and observed using an inverted microscope and the HIQ GFP 41014 filter set (Chroma). Mutants were isolated using micropipettes (10 μm diameter; World Precision Instruments), transferred to a drop of SD on the same coverslip, and then to YPD plates. Micropipettes were handled by an Eppendorf micromanipulator 5171.
Crosses to wild-type strain 1002 showed that all class I, II, and III mutations were recessive and caused by a defect in a single gene. Complementation tests revealed that all eight recessive class IV mutants were defective in the same gene. Crosses between class IV mutants and TRP1 strain 194 (a gift from E. Schweizer) or dnm1Δ strain JSY1361 ( Otsuga et al. 1998) showed the class IV mutation was centromere linked, located on chromosome XII, and allelic to dnm1. DNM1-containing plasmid, pRU1-DNM1 ( Otsuga et al. 1998), rescued the mitochondrial phenotype of all recessive class IV mutants. All dominant and semidominant class IV mutations also segregated as alleles of dnm1.
Complete disruptions of the DNM1 and FZO1 were constructed by PCR-mediated gene replacement as described ( Lorenz et al. 1995) into strains BY4733 and BY4744 ( Brachmann et al. 1998). For dnm1Δ, we used HIS3 plasmid pRS303 ( Sikorski and Hieter 1989) and for fzo1Δ we used kanMX4 plasmid pRS400 ( Brachmann et al. 1998). MATa dnm1Δ fzo1Δ strain YHS27 and MATα dnm1Δ fzo1Δ strain YHS23 were constructed by crossing MATa dnm1Δ strain YHS19 to MATα fzo1Δ strain YHS22. Mitochondria in the disruption strains were visualized using pHS12, a CEN-LEU2 plasmid containing ADH1-COX4-GFP. pHS12 was created by inserting the XhoI-NotI fragment from pHS1 into pRS315 ( Sikorski and Hieter 1989).
The DNM1 gene with a NotI site immediately preceding its termination codon was PCR amplified from yeast genomic DNA and subcloned into pAA3, a CEN-LEU2 plasmid which contains the HA epitope with a NotI site at its NH2 terminus (Aiken, A., unpublished data), forming pDNM1-HA (pHS14). DNM1-GFP plasmid pHS20 was constructed as described above except that pAA1, a CEN-LEU2 plasmid which contains GFP with a NotI site at its NH2 terminus (Aiken, A., unpublished data), was used instead of pAA3. To form pHS15, DNM1-HA coding sequences were PCR amplified from pHS14 with 50 bp of flanking sequences homologous to the GAL1-URA3 promoter in pRS314GU ( Nigro et al. 1992). The DNM1-HA fragment and linearized pRS314GU were cotransformed into yeast and pGAL1-DNM1-HA (pHS15) was formed by homologous recombination ( Oldenburg et al. 1997). pGAL1-DNM1-GFP (pHS40) was constructed as described for pHS15 except that pHS20 was used instead of pHS14.
dnm1Δ fzo1Δ cells carrying pGAL1-DNM1-GFP were incubated in galactose media for 1–2 h, stained with MitoTracker Red CMXRos (Molecular Probes). 12 cells were examined by fluorescence microscopy and the mitochondrially associated Dnm1p-GFP dots (82 total) were assigned to one of two locations: (a) the end of a tubule (50 dots), or (b) the side of a tubule (32 dots). The end of a tubule was defined as when the center of a Dnm1p-GFP dot was located within 0.15 μm from the end. The average length of the mitochondrial tubules was estimated to be 2.7 ± 1.9 μm and the diameter ∼0.3 μm (n = 44). We calculate that the side of the tubule represents 89% of the mitochondrial surface area and the remainder (11%) represents the end.
We screened for yeast mutants defective in mitochondrial shape using a novel strategy in which mitochondria are visualized by the green fluorescent protein (GFP) and mutants were isolated by micromanipulation. GFP was fused to the presequence (residues 1–21) of mitochondrial cytochrome oxidase subunit IV (COX4; Pon and Schatz 1991). When expressed in yeast, COX4-GFP targets the mitochondrial matrix, and mitochondria were visible by fluorescence microscopy. We integrated the COX4-GFP gene at the nonessential MFA2 locus ( Michaelis and Herskowitz 1988), which made fluorescence intensity uniform among cells and enabled efficient screening. After mutagenesis, individual cells with abnormal mitochondrial shape were hand-isolated using micropipettes ( Fig. 1 A). This screening procedure allowed us to isolate individual mutant cells with interesting mitochondrial phenotypes from a large total population of cells.
Mutants defective in mitochondrial shape. (A) Screen to isolate mutants defective in mitochondrial morphology. (a) Mitochondria were visualized by COX4-GFP. (b–d) Individual mutants were isolated using a micropipette. The arrow indicates a potential mutant cell. (B) Wild-type YHS2 and a representative from each class of mutants were grown in YPD to log phase and then stained with 1 μg/ml 3,3′dihexyloxacarbocyanine (DiOC6) (Molecular Probes) to enhance mitochondrial fluorescence. Confocal images of cells are shown. (C) dnm1Δ cells contain an interconnected network of mitochondria. Wild-type (BY4733) and isogenic dnm1Δ cells expressing COX4-GFP (pHS12), were grown in YPGal to log phase. Fluorescence (COX4-GFP) and differential interference contrast (DIC) images are shown. Bars: (A) 10 μm; (B) 2 μm; (C) 3 μm.
Of ∼72,000 cells screened, we isolated 20 mutants, which were classified into four categories ( Fig. 1 B). Class I mutants (two isolates) contained one or two large, spherical mitochondria instead of the normal tubules seen in wild-type cells. Genetic crosses showed that both carried mdm10 mutants ( Sogo and Yaffe 1994). The single class II mutant contained one or two oblong mitochondria collapsed to one side of the cell and was found to be defective in SLM1. slm1 was previously identified as an mmm1 synthetic-lethal mutant ( Burgess et al. 1994; Burgess et al., manuscript in preparation). Class III mutants (three isolates) contained numerous mitochondrial fragments and were shown to carry fzo1 mutations. FZO1 encodes a GTPase anchored in the mitochondrial outer membrane that is required for mitochondrial fusion ( Hermann et al. 1998; Rapaport et al. 1998). Class IV mutants (14 isolates) exhibited a novel phenotype consisting of an interconnected network of mitochondrial tubules. In contrast to wild-type, which have 5–10 separate mitochondria per cell, class IV mutants appear to contain a single organelle. Because of their unique networked mitochondrial shape, these mutants were examined further.
Genetic crosses showed that our 14 class IV mutants comprised 8 recessive, 5 dominant and 1 semi-dominant mutations. Mapping studies showed that all 14 mutations were centromere linked (1.1 cM) and located on chromosome XII. We noted that DNM1 ( Gammie et al. 1995), a gene related to dynamin GTPase ( Obar et al. 1990), maps to chromosome XII near the centromere and is required for mitochondrial shape ( Otsuga et al. 1998). Using a dnm1Δ strain and a plasmid containing DNM1 (kindly provided by J. Shaw), we found that all 14 class IV mutants carried dnm1 alleles. These results were unexpected since mitochondrial shape in our mutants was strikingly different from previously seen in dnm1 mutants, where mitochondria collapse to one side of the cell and form a single tubule ( Otsuga et al. 1998).
A complete disruption of DNM1 coding sequences was constructed, and examined for mitochondrial shape ( Fig. 1 C). ∼90% of dnm1Δ cells showed a single highly branched mitochondrial network. ∼10% of dnm1Δ cells displayed a single mitochondrial tubule localized to one side of the cell, similar to that seen earlier ( Otsuga et al. 1998). The mitochondrial shape was not dramatically altered by growth conditions (not shown). Mitochondria in dnm1 mutants were efficiently segregated during cell division ( Fig. 1 B and 2). Small daughter buds often contained a single mitochondrial tubule without branches, while larger buds had small networks. Most mitochondria were continuous from mother cells to buds, and separate mitochondria were only seen after the two cells separated. Our results strongly suggest that dnm1 mutants are defective in mitochondrial division. We speculate that mitochondria in dnm1 mutants may be divided indirectly, perhaps by cytokinesis. The yeast cell division machinery is clearly robust enough for the job, since nuclei are efficiently severed by cytokinesis in S. pombe cut mutants ( Uzawa et al. 1990).
In yeast, mitochondria are very dynamic, fusing or dividing on average every two minutes ( Nunnari et al. 1997). Thus there appears to be an equilibrium between fusion and division. Supporting this idea, when FZO1, a gene required for mitochondrial fusion, is defective, mitochondria fragment due to continued fission of the organelle ( Hermann et al. 1998; Rapaport et al. 1998). We hypothesized that if mitochondrial division were blocked, cells would have fewer (larger) organelles. Our working model, based on the morphology of dnm1 mutants, is that Dnm1p is required for mitochondrial division. To test this hypothesis, we constructed double mutants containing both dnm1Δ and fzo1Δ by genetic crosses. We found that normal mitochondrial shape was restored ( Fig. 2). ∼85% of double mutants contained multiply-branched, tubular mitochondria very similar to those seen in wild-type cells (Table 1). This was in marked contrast to dnm1Δ mutants, which usually had a single organelle, and fzo1Δ mutants, with numerous mitochondrial fragments ( Fig. 2; Hermann et al. 1998; Rapaport et al. 1998). Mitochondria in dnm1Δ fzo1Δ cells were not always completely normal; the tubules tended to be longer and more curved than in wild-type cells, and occasionally formed bundles. Nonetheless, our observations suggest that excess mitochondrial division in fzo1Δ cells is suppressed by inactivating DNM1, and that excess mitochondrial fusion in dnm1Δ cells is rescued by fzo1Δ. We propose that division, which requires Dnm1p, and fusion, controlled by Fzo1p, have antagonistic effects on mitochondrial shape and number. Our results also suggest that mitochondrial tubule formation occurs by a mechanism independent of fusion and division.
dnm1Δ fzo1Δ cells have normal-shaped mitochondria. Wild-type (BY4733) and isogenic deletion strains (dnm1Δ, fzo1Δ, and dnm1Δ fzo1Δ) expressing COX4-GFP (pHS12), were grown in YPGal to log phase and examined by fluorescence microscopy. Bars, 2 μm.
Interestingly, mitochondrial shape and number in dnm1Δ fzo1Δ cells was dependent upon the order of gene disruption. When cells were first disrupted for FZO1 and subsequently for DNM1 (Table 1, fzo1Δ→dnm1Δ), ∼40% of cells carried mitochondrial fragments similar to those seen in fzo1Δ single mutants. In contrast, when cells were first disrupted for DNM1 and then for FZO1 (Table 1, dnm1Δ→fzo1Δ), ∼30% of cells displayed a mitochondrial network like that seen in dnm1Δ cells. Our results indicate that the mitochondrial networks found in dnm1Δ mutants persist in the absence of fusion activity, and fragments formed in the fzo1Δ mutant persist in the absence of fission activity. We also found tubular mitochondria in many of the double mutant cells formed by consecutive gene disruption (∼50% for fzo1Δ→dnm1Δ; ∼60% for dnm1Δ→ fzo1Δ). These results further indicate that mitochondrial tubules form in the absence of division and fusion. It is not clear why dnm1Δ fzo1Δ double mutants generated by crossing a dnm1Δ cell to a fzo1Δ cell contained mostly (>80%) tubular mitochondria and essentially no mitochondrial networks or fragments (Table 1). During germination and growth of a dnm1Δ fzo1Δ spore, it is possible that cells are simultaneously depleted of Dnm1p and Fzo1p, leading to the formation of tubules, but not networks or fragments.
To further test the role of Dnm1p in division, we induced Dnm1p expression in dnm1Δ fzo1Δ cells and observed its effect on mitochondria. Dnm1p was fused to the HA epitope (Dnm1p-HA) ( Field et al. 1988) and expressed under the galactose-inducible GAL1 promoter ( Nigro et al. 1992). Our pGAL1-DNM1-HA rescued the dnm1Δ phenotype on galactose medium (not shown). When dnm1Δ fzo1Δ cells containing pGAL1-DNM1-HA were grown in the absence of galactose, no Dnm1p-HA was detected ( Fig. 3 B) and ∼70% of cells displayed the tubular mitochondria typical of dnm1Δ fzo1Δ mutants ( Fig. 3 A). Upon transfer to galactose medium, Dnm1p-HA levels gradually increased, while the level of hexokinase, a control protein, remained constant ( Fig. 3 B). Concomitant with the accumulation of Dnm1p-HA, mitochondrial shape changed dramatically ( Fig. 3 A). The number of cells with tubular mitochondria decreased, and those with fragmented mitochondria increased. By 5 h, ∼65% of the cells contained completely fragmented mitochondria. At intermediate times (2 h) after inducing Dnm1p-HA, cells contained partially fragmented tubules, and many mitochondrial tubules were adjacent to small fragments. Our results clearly show that the division of mitochondria in dnm1Δ fzo1Δ cells coincides with the expression of Dnm1p.
Expression of Dnm1p causes fragmentation of mitochondria in dnm1Δ fzo1Δ cells. dnm1Δ fzo1Δ cells carrying pCOX4-GFP (pHS12) and pGAL1-DNM1-HA (pHS15) were pregrown in raffinose medium, centrifuged and resuspended to an OD600 of 0.2 in galactose medium (SGS) to induce Dnm1p-HA expression. Cells were examined for mitochondrial shape (n = 100) or used to prepare total protein at the indicated timepoints. (A) Mitochondrial shape was classified into the following four groups: tubules (○), partially fragmented tubules (□), fragments (▴), and other (⋄). (B) Total protein was extracted as described ( Yaffe and Schatz 1984) and subjected to Western blot analysis using antibodies to the HA epitope ( Field et al. 1988), or hexokinase (Davis, A., unpublished data) followed by chemiluminescence (Pierce). Relative amounts of Dnm1p-HA (+) were quantitated and plotted in A.
Further supporting a role for Dnm1p in mitochondrial fission, we found that the Dnm1 protein was preferentially localized to sites of mitochondrial division. We constructed a fusion between Dnm1p and the green fluorescent protein (GFP). Consistent with previous results ( Otsuga et al. 1998), we found that much of Dnm1p-GFP was associated with mitochondria in punctate structures ( Fig. 4 A). In cells that were constitutively expressing Dnm1p-GFP, it was difficult to determine the precise location of Dnm1p because of the complex morphology of the mitochondria and the large number of Dnm1p-GFP dots. To simplify our analyses, we induced the expression of Dnm1p-GFP in the dnm1Δ fzo1Δ mutant and examined cells at early times after induction. We found a tight correlation between the appearance of Dnm1p-GFP and fragmentation of mitochondria. Two representative cells are shown in Fig. 4B and Fig. C; both cells contained two Dnm1p-GFP dots, one of which was located at the end of a tubule, the other appeared to be reside near a constricted region of the mitochondrion. After analysis of additional cells, we found that Dnm1p-GFP was localized to ends of mitochondrial fragments much more frequently (>60%) than predicted if Dnm1p-GFP was randomly distributed on mitochondria (∼11%). These results suggest that Dnm1p acts at the site of mitochondrial fission. We note that Dnm1p-GFP is not exclusively found at the ends of mitochondria. We surmise that Dnm1p on the sides of the tubules may mark future sites of division, or represent Dnm1p-containing complexes that have diffused away from the end of the tubule. More definitive experiments (e.g., time-lapse videomicroscopy) to determine the role of Dnm1p in fission are in progress.
A Dnm1p-GFP preferentially localizes to site of mitochondrial division. (A) dnm1Δ strain YHS19 was transformed with pHS20 expressing Dnm1p-GFP. Cells were grown in SGal medium, labeled with 0.1 μM MitoTracker Red CMXRos (Molecular Probes) and then examined under the fluorescence microscope. Merged images taken in the red (Mitotracker) and green (GFP) channels are shown. (B and C) dnm1Δ fzo1Δ diploid cells carrying pGAL1-DNM1-GFP were pregrown in YPglycerol/ethanol medium, centrifuged and grown in galactose medium (SGS) for 1–2 h. Cells were then stained with MitoTracker and examined for mitochondrial localization of Dnm1p-GFP. Two representative cells are shown. Bar, 3 μm.
The relatively normal mitochondria seen in dnm1Δ fzo1Δ mutants could be explained by a restoration of fusion activity; for example, if Dnm1p were an inhibitor of mitochondrial fusion. To test this possibility, we monitored mitochondrial fusion during mating ( Nunnari et al. 1997; Okamoto et al. 1998). Mitochondria were visualized in one parent, by the galactose-induced expression of a matrix-targeted GFP (CS1-GFP) on plasmid pCLbGFP ( Okamoto et al. 1998). MATa cells containing pCLbGFP were pregrown in galactose medium to induce CS1-GFP expression, then transferred to glucose to inhibit further synthesis. MATa cells were then mixed with MATα cells, which did not carry pCLbGFP, and allowed to mate on glucose medium. Mitochondria were visualized in zygotes using MitoTracker. If mitochondrial fusion occurred, GFP and MitoTracker fluorescence would completely overlap, because the matrix-localized CS1-GFP from the MATa mitochondria diffused into the mitochondrial matrix of the MATα cell. If no fusion occurred, GFP-labeled mitochondria would be seen in only one half of the zygote.
Zygotes formed by two wild-type cells, or two dnm1Δ mutants, exhibited efficient mitochondrial fusion, with GFP fluorescence and MitoTracker overlapping in all mitochondria ( Fig. 4 A). In contrast, fusion was defective in fzo1Δ mutants, consistent with previous observations ( Hermann et al. 1998). Mitochondrial fragments tended to aggregate in fzo1Δ cells and individual fragments were difficult to distinguish. Nonetheless, in matings between two fzo1Δ cells, MitoTracker showed clusters of fragmented mitochondria in the zygote and diploid bud, but we detected GFP fluorescence in only half of the mitochondrial clusters. Like fzo1Δ mutants, dnm1Δ fzo1Δ double mutants failed to fuse their mitochondria. Although mitochondria in dnm1Δ fzo1Δ/dnm1Δ fzo1Δ diploid cells had normal shape, only half of the organelles contained GFP. Our results indicate that dnm1Δ fzo1Δ cells are defective in mitochondrial fusion.
To eliminate the possibility that low, basal levels of fusion occur in dnm1Δ fzo1Δ cells, we used a more sensitive fusion assay using the matrix markers, CS1-GFP and mitochondrial DNA (mtDNA; Okamoto et al. 1998). 4′,6-diamidino-2-phenylindole (DAPI) stained mtDNA was a more stable probe compared with MitoTracker, allowing us to examine cells for longer times following the initial mating event. MATa cells, which lacked mtDNA and carried pCLbGFP were mated to MATα cells, which contained mtDNA, but not the plasmid ( Fig. 5 B). The DAPI and GFP fluorescence overlapped in all the mitochondrial tubules in wild-type zygotes (52 zygotes examined). When 500 dnm1Δ fzo1Δ zygotes were examined, we found no overlap between DAPI and GFP. The fusion activity in dnm1Δ fzo1Δ mutants is therefore at least 500-fold less than that in wild-type cells. Even after the zygotes were allowed to grow and divide, we found no fusion in the mutant cells ( Fig. 5 B). When 100 dnm1Δ fzo1Δ/dnm1Δ fzo1Δ diploid cells were examined 24 h after mating, none contained an overlap between GFP and DAPI, whereas a complete GFP and DAPI overlap was seen in 43 wild-type diploids. dnm1Δ fzo1Δ cells clearly lack significant mitochondrial fusion activity. Our results above also suggest that dnm1Δ fzo1Δ cells lack fission activity. We therefore propose that in cells lacking Fzo1p and Dnm1p, mitochondrial tubule formation occurs by a mechanism independent of fusion and division, such as growth from the ends of preexisting organelles.
dnm1Δ fzo1Δ cells are defective in mitochondrial fusion. (A) MATa cells that express CS1-GFP from the GAL1 promoter of pCLbGFP were grown in galactose-containing medium (SGS). MATα cells lacking the plasmid were grown in glucose medium (YPD). MATa and α cells carrying the indicated genotypes were mated on YPD for 3.5 h, and then stained with 0.1 μM MitoTracker. The distribution of MitoTracker and GFP in representative zygotes containing a medial bud (asterisks) are shown. Arrows indicate three clusters of mitochondria lacking CS1-GFP in fzo1Δ zygotes. (B) Wild-type and dnm1Δ fzo1Δ MATa cells were treated with ethidium bromide to induce loss of mtDNA ( Fox et al. 1991), transformed with pCLbGFP, and then grown in galactose-containing medium (SGS). Wild-type and dnm1Δ fzo1Δ MATα cells containing mtDNA, but lacking the plasmid, were grown in YPD. Cells were mated on YPD and stained with 0.5 μg/ml 4′,6-diamidino-2-phenylindole (DAPI; Molecular Probes). A representative zygote formed by mating between wild-type cells or between dnm1Δ fzo1Δ mutants is shown. Fluorescence (GFP or DAPI), differential interference contrast (DIC) and merged images are shown. Mating mixtures were also incubated for an additional 24 h at 30°C on YPD, and representative diploids are shown. N indicates nuclear DNA staining. Bar, 3 μm.
Dynamin has been proposed to work as a mechanochemical enzyme that 'pinches off' plasma membrane invaginations, forming intracellular vesicles ( Takei et al. 1995). Supporting this idea, dynamin self-assembles into spiral-like structures ( Hinshaw and Schmid 1995) which can sever artificial membranes in vitro ( Sweitzer and Hinshaw 1998). It is also possible that dynamin plays a regulatory, instead of an enzymatic, function in membrane scission ( Sever et al. 1999). Future studies are clearly needed to determine the precise mechanism that Dnm1p plays in mitochondrial division.
We especially thank J. Shaw for generous gifts of strains and plasmids. We also thank J. Holder, O. Kerscher, S. Michaelis, J. Boeke, A. Aiken, A. Davis, R. Butow, K. Okamoto, D. Murphy, R. Bustos, C. Machamer, and T. Kai for reagents, equipment and technical advice. We thank C. Machamer, K. Wilson, and the members of the Jensen lab for valuable comments on the manuscript.
This work was supported by a PHS grant (R01-GM54021) to R.E. Jensen and a JSPS fellowship to H. Sesaki.
Abbreviations used in this paper: GFP, green fluorescent protein; mtDNA, mitochondrial DNA.
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It is often cited how the Milwaukee Junction and Belt Line Railroad area east of Detroit's New Center has been thought of as the "cradle of the automotive industry," but it seems that Commonwealth Street just west of New Center used to be something of an automotive innovation hot spot as well. A string of vacant industrial buildings huddled together on Commonwealth's 5700-5900 block are all that remain to attest to this brief chapter in Motor City history.
With several spurs peeling off of the main railroad trunk lines that cut right through this area, it appears destined to have been an industrial haven by the turn of the century. Through-rail access to the other automobile manufacturing sectors of the city such as Milwaukee Junction seem to have made this area ideal as the hub of a light manufacturing and supply industry, and for smaller carmakers who relied on large components (such as motors) from other suppliers. It also stood on the edge of the popping New Center area that was rising in the 1920s as the inner sanctum of General Motors, which was soon to be crowned by the towering Fisher Building.
Some of the supply-related or brassworking companies clustered in this vicinity included Detroit Lubricator Co., Penberthy Injector Co., Detroit Auto Specialty Co., Schroeder Paint & Glass, Monarch Foundry, Acme Brass Foundry, Crescent Brass & Pin, the Detroit Brass Works, and the Westinghouse Electric & Mfg. Co. Meanwhile, the Warren Motor Co. (a short-lived carmaker), and Scripps Motor Co. (a marine engine manufacturer) were also located in this corridor. You may recall my older post exploring the nearby Caille Motor Co., another marine engine maker and pioneer of coin-operated machines. Furthermore, Burroughs Corp. was also close by; one could think of this area of the city as a sort of extension (or predecessor, rather) of what is now being branded as "Tech Town" in the northern Cass Corridor.
Detroit1701.org explains that it was because of the abundance of these sorts of "parts makers" and the skilled workforce they employed here that Detroit was destined to become the center of the auto industry; we had already been producing these sorts of mechanical accouterments for the carriage, bicycle, railroad car, and ship building industries since the mid-1800s. But the story of the Commonwealth corridor (or at least this block of it) is really the story of a nearly forgotten inventor from Detroit's golden era, by the name of Joseph LaVigne.
Mr. Joseph P. LaVigne (also seen spelled "Lavigne;" I'm not sure which is correct) is not found in the usual social registers of notable Detroiters that I checked, but he was in the newspapers a fair bit, and held no less than 385 patents when he died. He lived with his wife and daughter at 521 Lincoln Ave. (renumbered to 5465 Lincoln after 1921), which stood about three blocks southeast of here. LaVigne was once quoted as saying "Don't call me an inventor. I am a designer; nothing more."
His LaVigne Manufacturing Co. used to stand here on Commonwealth where these buildings are now, and made brass items such as radiator valves, unions, elbows, oilers, and other such fittings used on early gasoline engines. Its success was heralded by the Detroit Free Press as an example of the boom in brassworking firms in Detroit (you may recall my older post exploring the Roberts Brass Works, which also boomed during the same period).
By the way, these buildings you're seeing here were actually erected later by the Commonwealth Brass Corp., which took over LaVigne's company sometime around WWI.
Organized in early 1906 to handle his automobile-related inventions, the LaVigne Mfg. Co. started with just his patents—some of which were contested in a lawsuit by his former employer, Detroit Lubricator Co. (which incidentally was located one block over, on Trumbull; they also made brass items such as valves and oilers). Mr. LaVigne seems to have been in court rather often based on a lot of the search engine hits I was seeing, but let us suffice it to say that it was fairly common back then for employers to try to *appropriate* the innovations of their employees. Sadly, it is often still the case today.
LaVigne was a stockholder in the Universal Wheel Co., and served as an officer of the National Union of Cyclecar Makers. He also ran the Liberty Mechanical School out of the rear of his home—over the profits of which he was sued again by a partner. Almost all of LaVigne's patents were in use however, which is pretty impressive.
Among the more lucrative of his patents: a quick-opening hot water valve, a carburetor, and a steering gear (for the manufacture of which he formed another company, the LaVigne Gear Co. of Corliss, Wisconsin). He also designed several embroidery attachments for sewing machines that were in common use, and in 1906 he was working on a way to reduce the price of cars by eliminating the need for a transmission. Had LaVigne been successful in this, it might've given him the chance to compete with Henry Ford's Model-T for the title of most affordable car, right before it exploded in popularity. Almost all the machinery in his factory on Commonwealth was "automatic," and designed by LaVigne himself, the Free Press said.
Griswold Motor Co. was sold to the C.H. Blomstrom Mfg. Co. the same year, mainly to acquire the rights to his idea for a vertical-crankshaft friction drive setup, which Blomstrom marketed as a safer alternative to crankstarting. LaVigne designed its vertical-crankshaft engine, which was pull-started by a lever from the driver's seat to eliminate the danger of injury from the crank's dreaded "kickback." The "gyroscope" engine, as it was called, still never took off. Incidentally, the compendious website coachbuilt.com explains that the Griswold Motor Co. was not to be confused with the separate Griswold Body Co., which also moved to a location somewhere on Commonwealth St., in 1909.
The April 23, 1908 Detroit Free Press carried a story, "Blazes Away at Cat, But Pinks Neighbor," describing the time Mr. LaVigne was accidentally shot while reading the paper on his back porch by an unhinged neighbor aiming at a troublesome cat. Mr. LaVigne recovered from the minor injury, and returned to the automobile industry in 1913 with the LaVigne Cyclecar Co., and the LaVigne Motor Co. He had built a promising prototype in a small shop behind his house (his wife was apparently just as accommodating of his boyish tinkerings as Clara Ford was to Henry).
The LaVigne Cyclecar was a light, air-cooled "underslung" car that was built in his plant here on Commonwealth. There was a roadster, a coupe, and a touring, ranging from $425 to $500—decidedly on the cheaper side of things. It also had a unique electrical setup where the headlights were integral to the front fenders, which turned with the front axle. The 1914 New York Auto Show yielded a promising contract for 5,000 units, but the "cyclecar" craze soon died down, so LaVigne marketed it as a "light car." Unfortunately the market for light cars collapsed as well, signaling the end of this venture.
There is a somewhat famous photograph of his seven-year-old daughter Olive LaVigne seated in the "Le Petite" car her father designed, and she created something of a sensation in those days by being seen piloting it around town (undoubtedly so as to advertise the ease with which her father's invention could be operated).
Olive later drove a 10-ton truck, and according to local historian Richard Bak she bragged that by 1955 she had driven over 500,000 miles in her "automobubbling" career. It is probable that she was even more famous than her father, and was heralded as the "first female driver," although I am not sure that is technically true. She later became a notable employee of the Detroit Recreation Department in the 1930s and 1940s.
Traveler Motor Car Co. was another separate venture of Mr. LaVigne's in 1913, which was initially also building cars in the same plant as the LaVigne Cyclecars on Commonwealth, although Traveler was financially backed from different sources. They manufactured two full size five-passenger touring cars and a roadster (marketing from $1,100 to $2,500), but despite being a full-size car, Traveler Motors also failed in 1914 and LaVigne was forced out of the auto industry again.
A blurb in the Detroit Free Press from May, 1921 reported that LaVigne and salesman E.J. Kleist had been held up by robbers on the Dixie Highway near the suburb of Trenton, while stopped at a train crossing. The "four thugs" got away with $1,500 in cash and diamonds. What LaVigne was doing down on the Dixie Highway carrying that much money (and diamonds!) during the Prohibition Era I don't know, but we can all certainly have fun speculating.
A Sanborn map from 1950 shows Commonwealth Brass Corp. then owned these buildings on the west side of the street where LaVigne Mfg. used to stand, from 5787 to 5947 Commonwealth. Commonwealth Brass Corp. actually did similar work as LaVigne, producing brass items for radiators, valves, etc. for steam and pneumatic applications; perhaps the corporation originally purchased LaVigne Mfg. and consolidated it with other smaller local firms. I suspect they took over LaVigne's foundry sometime around WWI.
In 1918 a young man named Milton Donahey went to work at Commonwealth Brass, where he advanced to a foreman position. Commonwealth Brass was a cutting edge firm that innovated the process of forging brass instead of using cast brass for their fittings, which was the industry standard at the time. There, Donahey produced the second brass forging die ever made in the United States.
Commonwealth Brass was part of UAW Local 157, which included tool & die workers from plants such as Fisher Body #21 and #23, McLaughlin Co., Ternstedt Mfg., etc. In 1950 Donahey decided to start his own company, United Brass Manufacturers, which still operates today, located out in the suburb of Romulus.
I personally think these buildings were erected sometime between 1930 and 1950. This is the one labelled "press shop" on the Sanborn maps.
An article by Max Homfeld at gasenginemagazine.com says Mr. Kermath was part owner of Eclipse Manufacturing Co. in 1907, a small machine shop where he built engines on contract for the Detroit Auto Marine Co. He operated in the Boydell Building in Greektown, alongside the Dodge Brothers—who at the time were still building engines for Henry Ford before they went into business for themselves (I went to a pretty cool loft party in the Boydell many years ago).
"The Detroit auto fever must have infected Kermath," Homfeld writes, "for he built an experimental automobile in 1907," but his destiny was in engines for boats. In 1910 he formed the Kermath Manufacturing Company, and soon the first Kermath marine engine was introduced, a four-cycle four-cylinder rated at 12 horsepower.
Kermath Mfg. produced many different models of reliable engines over the next few decades, from single cylinders to V-12s, which were commonly used by such prominent local boat builders as Garwood, Chris Craft, and Matthews as well as many others. Kermath also often converted conventional automotive engines for marine use, such as the Lincoln Zephyr V-12.
Counting the Scripps Motor Co., and the Caille Motor Co., that makes at least three marine engine manufacturers that were based in this area of the city (despite being located nowhere near a body of water). Some of this building's additions shown on the map were built as late as 1950. Kermath Mfg. stopped production in 1958.
Don't be fooled by the corrugated metal siding on the building; that's just cladding that has been added to cover up the ugly old brick...yuck—brick looks so old, right gang? Metal is modern! And modern is good!
It appears that the most recent tenant of 5900 Commonwealth was Commonwealth Industries, Inc., a company that did heat treating and other metalworking according to the journal Air Travel News, of April, 1929. The book Corporate Power and Urban Crisis in Detroit seems to indicate that Commonwealth Industries, Inc. was later absorbed by the Masco Corporation, which is today one of the giant conglomerates of the Detroit automotive supply industry.
How all this junk got here is perhaps another question, but there sure was some interesting stuff in here to wander around and look at. Yes, I habitually hang out in junkyards.
According to Sanborn maps, the Talbot Coal Co. (started in 1906 by Cash W. Talbot) originally occupied the footprint that is now filled in by this junkyard.
The M.E. Griswold Co. was formed in the 1890s making bicycles with stamped sheet metal, which was a new method at the time. However, according to the compendious website coachbuilt.com, Griswold soon found greater profits in stamping automobile parts, and moved to Detroit in 1909, to Commonwealth Street, in a building that had previously been the King Stock Yards Hotel (what the hell kind of a name is that for a hotel?). It was destroyed by fire in 1911 but was rebuilt with the large brick structure we see today. From 1912 to 1930 Griswold stamped bodies for many early Detroit car and truck makers such as Columbia Motors, King, Chrysler, Jewett, Paige, Willys-Knight.
When Graham took over Paige-Detroit, they cancelled all outside contracts, and Griswold lost a large chunk of their business. Their "swan song" however, as coachbuilt.com put it, was in 1928 when they built a one-off body for the Auburn Automobile Co., which was famous for their iconic boat-tail speedster. Griswold's "Cabin Speedster" concept body was advertised in Auburn's catalogue that year as "...tomorrow's automobile design," blending the concepts of "racing car and airplane," designed by famous aviator Wade Morton.
Griswold's Auburn speedster was a sensation at the New York and Los Angeles auto shows, but unfortunately the concept car was lost in a fire. Griswold did some production bodies for Willys-Knight in 1929, but after that it was pretty much a wrap.
I wrote before about another similar street on the city's west side that was dedicated to the automotive supply industry, in a post entitled, "The Epworth Corridor."
Other buildings I have explored in the area of Commonwealth are Redeemer Episcopal Church, and the National Biscuit Co.
Our visit was cut short when we noticed a police car of some sort parked in the gravel lot between the buildings, two officers outside casually poking around. I waited until the officers had their backs turned before I made a swift and silent emergence from the doorway my partner and I were watching them from, so that they would not actually see us exiting a building.
I greeted the officers with perfect nonchalance, and they naturally asked what we were up to. I explained that I was showing my visiting New York friend around the city and photographing graffiti, while noting that the officers were still quite at ease, and apparently not about to do anything to us. They must have stopped when they saw my ramshackle pickup parked, thinking it was a stolen vehicle, or that I was a scrapper? Thankful that we didn't *fit the description,* we were allowed to continue on with our day unmolested.
I would definitely stay at the King Stock Yards Hotel.


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By the Shores of the Lake, by Herbert Kastle (epub/Kindle/pdf)
Zelda's Tavern was at the crossroads, and most of the local boys, farmers' sons and youngsters from the tiny town of Liggon's Falls, congregated there for a cold beer about eight o'clock each night. Sometimes they brought dates, but that was a rarity since the area was notoriously short of eligible females. That didn't include the summer vacationists from New York City, of course... And that's where the trouble started.
Traces of Brillhart: A William Deacon Mystery, by Herbert Brean (epub/Kindle/pdf)
Brill Brillhart, the songwriter, was known as the biggest heel to hit New York in four decades. So t
Traces of Merrilee: A William Deacon Mystery, by Herbert Brean (epub/Kindle/pdf)
William Deacon, magazine journalist and sometimes detective, suddenly heard himself say, "Okay, if y
They're Chasing Us!, by Herbert Kastle (epub/Kindle/pdf)
The Lake Monster, by Stephen Wasylyk (epub/Kindle/pdf)
Malcolm Sage, Detective, by Herbert Jenkins (epub/Kindle/pdf)
"Malcolm Sage had been a hot-shot intelligence agent for Britain's Division Z during the Great War,
"An astonishing book: don't miss it." --BOSTON GLOBE In the treacherous maze of post-war Berl
Mrs. Pigafetta Swims Well, by Reginald Bretnor (epub/Kindle)
Mr. Coastguard, this is what has happened to Pietro Pugliese, who is captain of the fishing boat Il
When he saw the school teacher in the window, he had to have her. Not only would the robot be a grea
Over the course of the late summer and early autumn, six horrible deaths had occurred. Five men and
The Almost Perfect Murder, by Hulbert Footner (epub/Kindle)
A collection of crime fiction short stories featuring Madame Rosika Storey and her resourceful assis
The Modern Wizard in His Magic Attic, by Cottrel Hoe [Jenny Baxter #7]
#7 in the Jennie Baxter series has Jennie investigating a new explosive in Vienna.

redpajama_set_name: RedPajamaCommonCrawl


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Meet Roxy (9 yo boxer) & Buddy (10 yo corgi/beagle) are a bonded pair of pups that are in search of a new home to spend our golden years. The family they lived with their entire lives are moving and their circumstances prevented them from taking them. They are both house trained, very sweet, well mannered and know basic commands. They are very confused being displaced. Looking for a new home that will accept both of them, love and pamper them throughout their golden years. Please lets find them a great home!


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To protect individual items such as ornaments or drinking vessels we have wine glass and bottle boxes depending on size.


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text:
Q: How to add cert and chain .pem files provided by a website to the list of ubuntu CA? irrelevant but possliby helpful background:
I'm trying to build openocd for using picotools to program raspberry pi pico and while running ./bootstrap I get the error "SSL connection timeout" when it's trying to connect to https://git.savannah.nongnu.org/git/git2cl.git/ from the terminal. But then i have no problem opening the same url in firefox. I read somewhere that it's mostly related to the proxy settings for the browser and the system not being the same but i double checked and my browser is using system proxy settings.
the question is:
I researched a bit and i guess the problem is that my ubuntu doesn't use the same CA certificate authority as the browser. I opened the url in my browser, clicked that lock icon next to the address bar,  digged a little bit and found two .pem files PEM(cert) and PEM(chain) to download. Should i just add the two files using this guide? It doesn't seem quite right to me... Shouldn't I somehow extract the chain file into multiple .pem files first? If so how do i do it?

redpajama_set_name: RedPajamaStackExchange


Just bought a 2001 Boxster in early January.
Once I received it I changed the oil and filter and saw the engine had more oil (1/2 gallon more). I was expecting the issue could be resolved after changing and filling just the necessary oil but still leaking…. Any ideas?
Then a couple of weeks after getting the car, the engine light turned on (error P0430). I was finally able to pass the emission test using Cataclean.
Also noted the clutch is "high and short". I hope this is not a serious issue. When shifting Clutch does not slip. Any suggestions?
I also noted some suspension noises in the rear suspension also. What can I do to define what it is?
Even with all that, at home we absolutely love the car and want to enjoy it!
Thanks for adding me to this forum. I really enjoy your stories, cars, comments!


text:
wdvalgardsonkaffihus.com Blog
general articles by W. D. Valgardson but also many about Iceland and Icelandic North American society and history
Tag Archives: trade
Iceland: War With England
Posted on May 2, 2013 by william
Have you heard of the Napoleonic wars? Did you know that because of them, England and Denmark were at war? Did you know that Danish ports were blockaded by the English and that the trade ships that were supposed to supply Iceland with food and trade goods and take Icelandic goods in return were not able to leave port?
Did you know that the people of Icelandic were brought to the brink of starvation? That friends of Iceland in high places in England intervened and obtained a declaration from the English king to lift the blockade on Iceland?
That's okay. Neither did I. Not, that is, until I began this arcane, strange research into life in Iceland in the late 1700s and the 1800s. I mean, most of us don't know Canadian history, never mind what was going on in Europe in the early 1800s.
However, The Iceland Journal of Henry Holland has numerous references to the Napoleonic War and its consequences for Iceland. They are just tidbits, footnotes, brief mentions but by the time I'd finished re-reading Holland's journal, I had a sense of Iceland being affected by something I've never heard anyone talk about.
In a footnote on page 114, Holland says" Bjarni (Sivertsen) may already have been engaged in negotiating the restoration (completed 1812) of funds pirated from Iceland during the 1808 Gilpin raid. Bjarni was no stranger to Britain. He was one of the Icelandic merchants whose ships were detained under war regulations at Leith 1807-1809; Sir Joseph Banks helped secure their release and offered financial support."
Later, he says about the School House, "Behind the building which forms the present schoolhouse, a new range of buildings has been begun upon, with a view to an enlargement of the Institution. The war with Denmark, among its other consequent calamaties, has been the means of entirely suspending this scheme—from the want which it creates in Iceland of all the materials for building."
Later in his trip he visits Mr. Jacobæus, whom, he says, "is one of the most considerable merchants in Iceland; and had much commercial connection with Denmark; though this trade has of course suffered greatly from the war between England and Denmark."
He visits the Leira printing office, the only printing office in Iceland, and says, "It has latterly been much injured by the winter floods, & it is now in projection to construct a new building on the same spot – the execution of which is only delayed form the scarcity of timber in Iceland during the period of war."
They visit a Mr. Gudmundson, a merchant, "who has commercial connections at Copenhagen, at Reikjaviik & at Havenefiord….We found from conversation with him that the war between England & Denmark had been greatly injurious to his trade—Three years have now elapsed since any vessel has come from Copenhagen to Buderstad; though previously to this time, it was usual for one or two vessels to enter the port annually – This privation of the accustomed inter course is severely felt by the inhabitants of the interior, who are greatly in want of corn, timber, iron &c. The warehouses at Buderstad, and other ports in this part of Iceland, are filled with the commodities of the country—fish oil, fox skins &c, which it is impossible at the present time profitably to dispose."
When they visit Stappen, he says the merchant of the place is a Mr. Hialtalin. However, he is not there because "At the commencement of the war, he was taken prisoner, & carried into England –Thence he contrived to get into Norway, where he was about 1 1/2½ years ago. Since that time no intelligence what ever of him has reached Iceland. His wife, Madame Hialtalin, with a family of six children, continues to reside at Stappen, where she carries on the business as well as lies in her power."
When he visits Mr. Clausen, another Danish trader, he discovers that "He collects for exportation, fish, oil, tallow, fox-skins, & the various woolen manufactures of the island–& sells to the inhabitants, both in a retail & wholesale way, different articles of foreign produce or manufacture, procured from the continent of Europe. The war between England & Denmark has been greatly detrimental to this trade. Beside the intercourse with Denmark much profit was formerly derived from the exportation of fish to France, Spain & the ports of the Mediterranean Sea; a branch of commerce which is now entirely suspended. The intercourse with England has not yet acquired a sufficiently settled footing to relieve these evils—Mr. Clausen's warehouses are crowded with goods for which a market is wanting—He reckons that he has lying by him, (either under cover, or collected into large heaps upon the shore) many hundred thousand fish, salted or dried—Of the woollen goods, manufactured in Iceland, his stock is proportionally large. He has about 50,000 pair of mittens, or woolen gloves, and almost an equal quantity of stocking of different qualities of fineness."
"Previously to the war, Stikkes-holm was a place of considerable trade, three or four vessels from Norway or Denmark generally coming to the port every year. The only vessel which arrived last year was one from Norway, & none has appeared here in the present summer."
"Mr. H. had been present at the late unfortunate attack upon Copenhagen—his house & much of his property had been destroyed in the bombardment of the place. He shewed us an umbrella, broken by a shot whilst he was sleeping under it, in a tent."
"Both Mr. Hialtalin (this is the brother of the missing Mr. Hialtalin) & Mr. Benedictsen spoke much of the distress produced in Iceland by the war between England & Denmark; & seemed to consider the English Order in Council as likely to afford only a very partial relief."
Reykavik has an annual trade fair, "The Handel", which Holland says is better this year, 1810, than the previous year "however, by no means equal to previous years."
When they arrive at Hyindarmule, they attempt to buy some items of Iceland dress from the women. "We were unable, however, to effect any thing of this kind—Not that the people were unwilling to sell – they were all pleased but the impossibility, during the present period of war, of replacing any article with which they now parted."
"We reached Eyarback at 5 in the afternoon…Previously to the war, three ships usually came there every year."
"The merchant proprietor at Eyarback is a Mr. Lambasson—This gentleman being detained in Norway by the war, the business is conducted at present by his wife, & by an agent, Mr. Peterson. We were received by them with great hospitality, & remained at the house a few hours to refresh ourselves."
It's interesting that, time and again, where they stay is with Danish merchants. Their countries are at war. The English bombardment of Copenhagen resulted in 2,000 civilians being killed and 30% of the buildings being destroyed. Danish ports were effectively blockaded and the Danish navy neutralized.
However, except for one instance, Holland and his companions never felt any hostility or disapproval.
This was not a two way war between France and England but a complex web of alliances and re-alliances with Denmark having to not only fight England in order to avoid being invaded by France but by Sweden. Germany and Russia, at various times, were part of the mix.
When I get time, I'll type up the letter of the English king who writes up the letter for the British Order in Council that is mentioned.
Posted in Uncategorized	| Tagged Denmark, England, France, Holland, Iceland, Napoleonic War, Order In Council, The Journal of Henry Holland 1810, trade	| 8 Replies
Was Afi A Smuggler
Posted on April 22, 2013 by william
Many of us, I say us, because it was certainly true of me, had no idea of what life was like for Icelanders in the 1700s and 1800s even though it was social and economic conditions during these times that led to our ancestors leaving Iceland the latter part of the 1800s. What I heard, repeatedly, was that our ancestors left Iceland because of the volcanic eruptions. I've heard this repeated time and again, as if it were gospel, both in private conversation and in public speeches.
While it is true that the effects of volcanic eruptions contributed to people making the decision to leave for North America, another hardship piled upon previous hardships, there was much more to it than that.
Uno von Troil, writing his Letters on Iceland, based on his visit in 1772, gives a summary of what trade was like for the Icelanders. Some of the conditions he mentions in brief (unfortunately, he does not elaborate) give us a glimpse into life for our lang (x6 or so) ammas and afis.
Von Troil points out that over the centuries there had been many changes in the way that trade was conducted with the Icelanders. What is obviously the major problem is that the trade was not under the control of the Icelandic people. The days of Viking ships were long past. The Icelanders were in no position to carry their goods to various markets.
He points out that until around 1400, the Norwegians were virtually the only people trading with the Icelanders. What fish the Icelanders didn't eat or export in their own ships (the Icelanders still had ships with which to export goods), the Norwegians bought.
The English took over this trade, he says, until the Reformation. Although the mention of the English is just in this one line, the relationship of the English to Iceland waxes and wanes but never stops. It is long and, sometimes, troubled relationship.
During the Reformation, trade with Iceland was given to the Germans, particularly traders from Hamburg. Christian IV prohibited the Hanseatic League towns from trading with Iceland in 1602 and gave the right to trade to Copenhagen, Malmo and some other cities that were controlled by Denmark.
The Iceland company had the trade rights from 1620 to 1662.
In von Troil's notes, I found a most interesting comment about the Iceland Company and the Turkish raids. We all know about the pirate raids that resulted as a large number of Icelanders being taken and sold as slaves in Algeria.
According to von Troil, the king was upset with the Iceland Company because not only had the company, in return for their exclusive license, promised to provide the goods necessary for the Icelanders but to protect Iceland from marauders.
Von Troil doesn't explain the situation in detail but he says that the people who owned shares in the company were paid for their stock holdings but those people who had 1000 rix dollar shares were only given 500 and those who had 200 rix dollar shares got nothing. The company had paid the king for every trading port plus two rix dollars to the governor. It also contributed to the king's "magazines" on the Westman Islands.
After the Iceland company was done away with, the trade of every port was auctioned off to the highest bidder every six years. However, he says, since 1734, a trading company has been given exclusive trading rights to the Icelandic ports for a yearly sum of 6000 rix dollars.
In return for this payment, the company is supposed to send 24 to 30 ships a year. They bring (or are supposed to bring) grain, bread, wine, iron and wood, etc. There are 22 trading ports. There they get fish, meat, butter, blubber, skins , wool and woolen goods.
He says that he doesn't know if the trading company actually makes much money from the trade. The reason for that, he says is that the Dutch, illegally, bring much better goods to trade with the Icelanders. The Icelanders, illegally, sell their goods to the Dutch. This is regarded as smuggling and a number of Dutch ships have been seized. The legitimate trading company knows that smuggling and illegal trading is going on because the Icelanders bring such a small amount of goods to trade with them.
Von Troil mentions that at Reykjavik there is a woolen "manufactory, where twenty or thirty workmen are employed" but he does not explain what it is that is manufactured. Is it wool being readied for market or is it finished products? Surprisingly, he says that there are only a few looms here and there. Later travellers make reports of looms but do say that they are small and primitive compared to the looms in Europe.
The state of the Icelandic economy can be judged by von Troil comments on money. He says there is so little actual money that in the entire country can't amount to more than a few thousand rix dollars.
The Icelanders keep their accounts not in money but according to yards of wadmal and number of fish. Forty eight two pound fish are worth one rix dollar. Twenty-four ells (about two to two and a half feet) are worth one rix dollar.
He says that you can buy a horse for 150 fish. So that would be about three rix dollars. You can buy a farm for 6000 ells. Think of that as 1000 yards of woven cloth.
So, there you have it, in the lines and between the lines. The Danish company that was supposed to protect the Icelanders in return for their trading license, didn't, and Icelanders were killed and taken into slavery. Profit before responsibility. Not by the king but by the trading company.
The goods supplied by the later trading companies was often of poor quality. This did not improve with time. Later reports in the 1800s make note of the fact that some ships, to increase profits, did not even bring poor goods to trade, but nothing except cheap Danish brandy.
Your ancestors, if they could, in the face of harsh penalties, traded smuggled goods because they were of better quality.
Your ancestors lived in an economy where there was no banking system, hardly any actual cash in silver or gold, and this would not change until the English and Scottish traders started coming and were paying cash for sheep and horses. That was still long in the future. When Anthony Trollope goes to Iceland on the yacht, The Mastiffe, owned by John Burns, in 1878, he comments on the fact that there is still no bank in Iceland. More than a hundred years have passed since von Troil's visit in 1772. Trollope says "One deficiency in Reykjavik which the most surprised me was the want of a bank. There is no such thing as a commercial bank in Iceland."
One must, of course, be wary of historical documents, particularly personal papers, for much is included that is hearsay. For example, von Troil says of the Icelanders kidnapped by the Turkish pirates that most were returned to Iceland. We know that is not true. Still, his observations are interesting and informative and give us some sense of what life was like for those great (x6) ammas and afis of ours, and a sense of the social and economic conditions that led, eventually, to our people going to the harbours and getting onto English ships to begin their journey to Amerika.
Posted in Uncategorized	| Tagged 1772, 1878, Danish, Hanseatic, Letters On Iceland, Mastiffe, Norwegian, smuggling, trade, trading monopoly, Trollope, Turkish raids, Uno von Troil	| Leave a reply
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Once you have booked the course and secured your place, you will be sent joining instructions within 24 hours.
You should firstly watch the introductory video, this will give you an overview of the course and additional information about preparing for the exam.
Before starting the course, it's best to familiarise yourself with the online portal and the additional references available.
You then need to work through the 13 modules of self-study.
Once you have completed the online course and revision, you will need to attend the 2-day exam preparation workshop and exam.
The first day and a half of this workshop will be spent preparing you for the exam. Your facilitator will help you to understand what the examiners are looking for in questions and the best way of structuring answers.
On the afternoon of the 2nd day, you will sit the exam which is a three-hour, essay-based assessment.
Notification of your results will be provided within 10 weeks' time by the APM.
We are confident that our experienced facilitators will get you through the exam, first time. However, if you don't happen to quite make it over the pass line, our Guaranteed Pass promise means we will continue to support you and pay for you to resit the exam and workshop.
Our blended package costs £1,495 + VAT and includes all of the above- no hidden fees!


The Child Care Program has been operated by the MLK Center since 1970. This program is State Certified to accommodate 105 children. In 1992, the Day Care Unit began providing services for at-risk children though direct referral from the Erie County Office of Children and Youth (OCY) under a "Family Support Center Project" in response to the growing incidence of child neglect and abuse in Erie County.
An expansion of the Child Care Program took place in May of 1995 to accommodate infants six(6) weeks of age through toddlers aged 18 months in response to the need within the community for qualified care for the younger child.
An additional building, owned by Bayfront NATO, Inc. and located across the street from the main facility, was extensively renovated for this Day Care expansion.
Infant/Toddler Day Care starting from birth to 24 months. Great emphasis is placed on early child development stages including cognitive, physical and social stimulating activities. Age appropriate deeding program is included, Similac Advanced formula provided.
Pre-School Care starts from 3 years of age. Curriculum includes standard structured learning of letters, numbers, colors, shapes, calendar activities, and library time. Creative play includes free choice of dramatic play, blocks, puzzles, table top activities, science and math centers. Nutritious state approved meals and snacks are included.
Pre-Kindergarten Readiness starts from 4 years of age. Prepares children for entry into school age environment, building on the Pre-School curriculum. Pertinent personal information is a focus to have each child be able to state their name, parent's name and home address. Emphasis is placed on teaching the role of public safety personnel, fire, and police.
Before-School Care for children aged 5 up to 13 years of age. Includes a state approved nutritious meal and interactive learning and/or sport activities. Children enrolled in the Before-School Care are transported to school if they are present prior to the bus departing the Center.
After-School Care for children aged 5 up to 14 years of age. Homework assistance is offered and children have the opportunity to participate in the Youth Department's program for additional programs.
The Summer Care Program is provided for children aged 5 through 13 years while their parents are working, attending school, or in training classes while children are out of school for summer vacation.
We have open enrollment year-round for Bayfront Childcare.
Bayfront Childcare has an income-based payment program and accepts CCIS.
To enroll your child in Childcare, please contact Darlean Henderson, Daycare Supervisor at 814-459-2761 x237 or fill out the form below.


Insane Clown Posse fans marched on Washington DC at the weekend in protest against the FBI's designation of them as a criminal "gang", showing that not all clowns are scary and out-numbering a nearby pro-Trump rally in the process.
The Juggalos – the fond name given to fans by the shock-rap group – were labelled a "loosely-organized hybrid gang" by the FBI in its 2011 National Gang Threat Assessment. The report argued that "many Juggalos subsets exhibit gang-like behaviour and engage in criminal activity and violence". This designation, Juggalos argue, has made it harder for many to find work, housing and access to other social services.
But despite their gang reputation, the Juggalos march on Washington was largely a peaceful and successful one, according to reports, with the 1,500 Juggalos in attendance outnumbering the couple of hundred pro-Trump supporters rallying behind their President elsewhere in the National Mall on Saturday (September 16). The Juggalos say they chose the particular setting due to it being the meeting place in an iconic scene from Forrest Gump.
The Juggalos were out in force with humorous signs ("Clown Lives Matter", "Grab America by the Posse" and "the FBI listens to Nickelback" among them), a sense of unity and even brought their kids along for the occasion. There was free pizza and soft drinks and, at one point, a Chinese tour group appeared to get lost amongst the rally.
"We're a family, a large one. I'm here to march for my people," one fan told the Associated Press.
See a selection of tweets, videos and photos from the Juggalo march below.


The project sought to reduce the sexual transmission of HIV in MARPs through the use of evidence based interventions and promising practices for Most At-Risk Populations. The goal was to increase the organizational capacities of MARP groups to ensure comprehensive response to the HIV epidemic and guarantee protection of human rights through promotion of public policies.
In particular, CAI contributed to the promotion of behavior change among MARPs regarding HIV prevention, stigma and discrimination, and gender equity, through the implementation of enhanced outreach education and social marketing of appropriate educational material.
CAI partnered with the Ministry of Health, NGOs and civil society to increase access to and uptake of voluntary counseling and testing services for MARPs by establishing a system for community-based voluntary counseling and testing, training of government health care providers, and ensuring quality of services provided to target population. Healthcare providers play a significant role in the provision of HIV prevention services to their clients. In order to ensure that MARPs received culturally competent services, CAI provided specialized trainings for healthcare providers; and this improved their competency to provide services to MARPs by providing a safe and respectful environment.
CAI contributed to the development of leadership and sustainability through the creation of a Human Rights coalition to advocate for MARPs rights. Key activities included developing agreements with private sector on workplace issues, establishing guidelines for health care providers on services that are friendly to the targeted groups, and training most at risk individuals to register complaints with local authorities.
Please view the project's final report by clicking here. The report is an interactive brochure, please click on the bottom right corner of it to turn page.


A half-Russian, half-Korean rocket likely exploded a few minutes after liftoff Thursday, dealing a second blow the South Korea's $400 million program to develop its own satellite launcher.
The Korea Space Launch Vehicle, or KSLV 1, blasted off from the Naro Space Center at 0801 GMT (4:01 a.m. EDT), or at 5:01 p.m. local time. The rocket flew through a nearly transparent deck of clouds about a minute after liftoff.
Live video from the space center showed no signs of any problems during the early moments of the launch, but officials with the Korea Aerospace Research Institute say they lost contact with the rocket 137 seconds into the flight.
At that point, the booster should have been flying between 40 and 50 miles in altitude on the power of a Russian first stage built by Khrunichev, the contractor in charge of the workhorse Proton launcher.
Some Korean news reports say the rocket appears to have exploded and crashed as controllers lost communications with the booster.
Nearly 375,000 pounds of thrust from the Russian-made first stage RD-151 engine was supposed to propel the 108-foot-tall rocket more than 100 miles above the East China Sea in less than four minutes. The KSLV's nose cone was expected to separate 215 seconds after liftoff.
Khrunichev built the kerosene-burning first stage and lended expertise in construction of the Naro launch pad and control center under a 2004 contract signed with KARI.
After pulling free of the first stage, the Korean second stage was programmed to coast for nearly three minutes before firing its solid motor for a 58-second burn to inject the flight's 219-pound payload into orbit.
The launch was carrying the STSAT satellite with instruments to observe Earth and a laser reflector to help engineers track the craft's orbit. The rocket was targeting an orbit ranging in altitude from 186 miles to 932 miles, according to KARI.
The rocket is nicknamed the Naro 1 after its launch site.
South Korea has invested more than $400 million into the Naro rocket project.
The flight was delayed one day after the launch pad's fire suppresion system unexpectedly activated during a countdown Wednesday.
It was the second KSLV launch from the new space center, which was built 300 miles south of Seoul in the South Jeolla province near the southwestern tip of the Korean peninsula.
The Naro's first flight in August 2009 did not achieve orbit because half of its payload shroud clung to the rocket when it was supposed to fall away. The launcher tugged the extra mass into space, keeping the booster from reaching the planned orbit.
Read our story on the KSLV's first flight.
Engineers resolved the issue and started preparations for a second launch at Naro this spring.
If the flight was successful, it would have put South Korea on a short list of eight countries, plus Europe, with their own space launchers.
The flight was also a key test of the Russian first stage, which is similar to the rocket that will power Angara boosters off launch pads beginning around 2012.
Russia is building the Angara rocket family to replace the Proton and smaller vehicles in the country's arsenal. Designers plan to cluster between one and five universal first stages to customize the Angara rocket for a range of payloads.
Khrunichev is the Angara's prime contractor.
South Korea says it wants to build a larger rocket to haul up to 3,300 pounds to a sun-synchronous orbit popular with Earth observation satellites. Officials want to use indigenous technology to achieve the goal by 2020.
North Korea's state news agency had no official response to Wednesday's launch. Before last year's launch attempt, North Korea criticized the muted international response to the South's foray into the space arena.
An apparent satellite launch attempt by North Korea in April 2009 drew reprimands from the international community, which said the launch was just a disguised long-range missile test.
Tensions are high on the Korean peninsula after the sinking of a South Korean navy ship that killed 46 sailors in March. A South Korean investigation concluded the North torpedoed the vessel.


FDN's Foreclosure Defense Seminar which was scheduled for Friday, February 25, 2011 is being rescheduled as Mr. Barnes has been called to a court hearing in a foreclosure defense case in Indiana for that date.
The new date will be a Friday in March. We will update as soon as the date is confirmed.


The Human Resources (HR) department is typically the area where International employee benefits are introduced and implemented, such as long-term and short-term International Medical Insurance or Business Travel Accident (BTA).
Some international medical policies will need underlying domestic health insurance in place, others do not.
If your company only has a few employees traveling working and living abroad, an individual and family international medical plan will most likely be advised. For larger numbers of global employees the company can implement an international group medical plan for employees on short-term and long-term global assignments.
An international medical plan should always include; hospital services, outpatient, emergency evacuation, rx, Dr visits along with lab and X-Ray.
Some multinational companies will choose to go with a Business Travel Accident (BTA) policy. This can work well for medical coverage as a result of an accident. The policies will also include emergency evacuation along with some reimbursement features for luggage and lost expenses from cancelled flights/hotels etc.
The BTA premiums will typically be lower than a full international medical plan, because most do not include "illness" coverage, and there is typically a maximum benefit per trip.
Domestic Health Insurance provides very little and often NO coverage when outside the U.S. Worse yet, if an emergency strikes and you need a medical evacuation or need to be flown home for care, international medical provides this coverage while your domestic group health plan does not. Emergency medical evacuation and repatriation services can cost upwards of $100,000 depending on your location, the distance to care, and if a nurse or medical staff is necessary during the flight.
Some international medical insurance companies have concierge-level service for it's members. Global services are available 24/7/365 to assist members with anything from arranging direct pay, coordinating an emergency evacuation or getting the proper follow up care. For Human Resource administrators, the right policy is readily available to help you assist your employees while on their foreign work assignment.
The One-Call solution along with the mobile app, which is available with certain companies, allows the employee to receive or send vital information or connect directly with the company for assistance. This communication is a much needed asset for the employee and their family while living abroad.
If help is needed during a holiday, over the weekend or late at night, can they call their domestic carrier? Is HR open or available to assist? Global assistance is one of the many assets International medical companies provide.
If you have a unique situation with an employee, independent contractor or third country national, feel free to contact me anytime. I'm happy to assist with your international medical, group or individual needs.