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September 2002
Note: Because some of the information in this article may be outdated, it has been archived.
1. Introducing LUCA
LUCA is short for Last Universal Common Ancestor, and it is from this organism that every living cell on the planet has descended. LUCA does not represent the earliest stage in the evolution of life — it is widely accepted that before the evolution of proteins and DNA (which are common to all cellular life) there was a period where RNA carried out the roles now performed by proteins and DNA [Jeffares & Poole 2000]. There are a lot of uncertainties when we go this far back in evolutionary time and perhaps all we can be certain of is that, at a point in Earth's history (probably over 3 billion years ago), cells emerged which stored recipes for making both proteins and RNA on a third molecule, DNA.
Nevertheless, studying LUCA is not science fiction. In the same way as humans and chimpanzees shared a common history until less than 10 million years ago, all modern lifeforms shared a common history back as far as the split that gave rise to the three 'domains' of life we now know of as archaea, bacteria and eukaryotes; that is, back as far as LUCA. That there are three domains was first established by Carl Woese and colleagues, who found that the group called prokaryotes was actually two groups, the archaea and bacteria [Woese & Fox 1977; Woese et al. 1990]. Amazingly, this work has largely stood the test of time, and though it is argued that there has been extensive gene swapping between these two groups [Pennisi 1998, 1999; Doolittle 1999; Eisen 2000], recent analyses using complete genomes supports Woese and colleagues' decision to split prokaryotes into archaea and bacteria [Snel et al. 1999; Sicheritz-Pontén & Andersson 2001; Brown et al. 2001].
Woese and colleagues' discovery rested on an even more astounding one — all life stores its genetic information on DNA, using a common code which we call the genetic code. The information is stored as packets, called genes — recipes for making RNA, and proteins [see Appendix A: Making Protein]. The languages of DNA and RNA are so similar they may as well be called dialects, but both are markedly different from the language of protein. For RNA and DNA, the information-carrying part of both molecules is made up of four bases (analogous to letters in an alphabet) read in a linear fashion, as with written human languages. In RNA, the four bases are A, G, C and U. In DNA, A, G and C are also used, while T is used instead of U. Establishing the evolutionary basis for this change from U to T is not a trivial exercise, and is an interesting problem in itself [Poole et al. 2001]; but in terms of the actual language, the difference is as minor as the variant spelling of English words, e.g., civilisation and civilization.
The unearthing of the genetic code, and the subsequent demonstration that it is common to all life (a gene from a human can be read by the translation machinery of a bacterium) is without a doubt a key piece of evidence in establishing that there was a LUCA. But what else can we find out about LUCA? Knowing that the genetic code had arisen tells us there was probably a LUCA, but gives us very little information on the nature of LUCA.
In a nutshell, the study of LUCA broadly revolves around two questions:
- What features are common to all cellular life?
- What sets the three domains — archaea, bacteria and eukaryotes — apart from one another?
At first sight, building a list of LUCA features might seem a fairly straightforward process, especially now that advances in technology allow all the genes possessed by an organism to be identified by sequencing its genome. A sensible approach would perhaps be to compare all the genes from representative genomes of archaea, bacteria and eukaryotes. Those genes that are common to all three domains were in the LUCA and those that aren't must have been added later. Unfortunately, it's not that straightforward, for two main reasons:
- Some genes appear to have moved from organism to organism like genetic gypsies, confounding our ability to distinguish between features that are universal and date back to LUCA, and features that are universal because of genes moving about.
- Some genes which were found in LUCA may no longer be universal. That means it may be impossible to distinguish some LUCA features from genes that arose later, say in the evolution of eukaryotes.
Ask any two researchers to give an overview of what they think the LUCA was like, and you will no doubt get different answers. With such a tricky scientific endeavour as this — working out what an organism that lived billions of years ago was like — this is hardly surprising. Some of my own views on one aspect of the LUCA question are to be found in an earlier piece posted on Actionbioscience.org which I co-wrote with Dan Jeffares [Jeffares & Poole 2000], but what follows is a broader overview of the fast-growing field of 'LUCA biology.'
2. The minimal genome project
One hands-on approach to trying to uncover the biology of the LUCA has been to look for genes that are universal — that is, genes that all life forms possess. Once a list of these genes has been made, they also lead to another possibility: perhaps this list encapsulates the essence of cellular life — the minimum number of genes required to make a cell. In 1996, with the sequences of the first two bacterial genomes (Mycoplasma genitalium & Haemophilus influenzae) in hand, Arcady Mushegian & Eugene Koonin [Mushegian & Koonin 1996] tried exactly this. The most striking features of their minimal genome were:
- A mere 256 genes
- No biosynthetic machinery for making the building blocks of DNA
From this they tentatively concluded that LUCA stored its genetic information in RNA, not DNA, and made suggestions on how to further reduce the number of genes in their minimal genome. The work heralded the arrival of comparative genome studies, and there is no doubt that a good number of the genes in their 256-strong list do date back to the LUCA. However, the work was squarely criticised because of the omission of DNA [Becerra et al. 1997]. Both these bacteria are human parasites and it seems most likely that they did away with parts of the machinery for making their own DNA because they can steal from the host (i.e., the organism infected with these pathogens). Indeed, why put in the effort to make your own when it's there for the taking?
Regardless of whether or not DNA was a part of the LUCA (I think it was [Poole et al. 2000], but there are plenty of researchers that beg to differ [e.g., Leipe et al. 1999]), this omission highlighted a wider problem with the minimal genome. Namely, the genomes you begin with probably affect the final set of genes. This is a problem for the following reasons:
How many genomes must be compared before we are confident we aren't leaving anything out?
'Lifestyle' can affect the final list (in Mushegian & Koonin's work, the minimal gene set may in fact be a generic set required for parasitism in humans, and has little in common with what was required for a free-living cell to go about its business billions of years ago)
Gene losses: if a gene was in the LUCA, but now remains in only one of the three domains, this method would consistently leave it off the list of LUCA genes.
Finally, if genes can move from organism to organism (so-called horizontal or lateral gene transfer), certain genes may have done such a good job of spreading that they sometimes appear to date back to the time of LUCA, whereas in actual fact, they arose more recently.
Despite its limitations, the minimal genome concept is probably the best attempt to put money where mouth is and come up with a hard list of genes that may have been a feature of the LUCA. It is also the only sensible framework we currently have. That said, if gene transfer is extreme, genes will have moved about so often that this and related methods are rendered futile [Doolittle 1999].
Koonin has recently published an updated minimal gene set, using 21 complete genomes [Koonin 2000]. Surprisingly, of the 256 genes in the original set, only 81 remain, and this list is clearly insufficient to describe either the minimum number of genes required for a cell to function, or the genetic makeup of LUCA.
While working out which genes were part of LUCA is no easy task, the various attempts [Mushegian & Koonin 1996; Kyrpides et al. 1999; Hutchison et al. 1999] are a good starting point, and have served to highlight important problems that must be dealt with in the field of 'LUCA biology."
3. LUCA genomics
As the minimal genome work demonstrates, the major issues are:
- How much does reliance on universal features underestimate the genetic makeup of LUCA?
- How much gene swapping has gone on during the evolution of life from LUCA to the present?
The magic number of universal features is likely to shift about, and there have been plenty of criticisms of all the attempts to reconstruct the LUCA. Nevertheless, universal features are important because they describe a lower limit from which to build upon, and importantly, all these attempts converge on agreement insofar as concluding that LUCA was quite complex. Some of the gaps will be relatively simple to fill, but others may be close to impossible.
LUCA biologists are aware that universal features may underestimate the complexity of LUCA to some extent, but another concern is emerging that could cause even more headaches — the spectre of 'horizontal gene transfer' (also called lateral gene transfer):
- If there is lots of gene swapping between organisms, the tree of life becomes more like a web, and it may not be possible to disentangle the branches.
- If genes are extremely nomadic, truly universal features cannot be distinguished from genes that have successfully spread themselves by gene transfer.
The problem of gene transfer was made apparent in a landmark study of two bacterial genomes, Escherichia coli and Salmonella. Jeffery Lawrence & Howard Ochman [Lawrence & Ochman 1998] concluded that, since diverging from a shared ancestor 100 million years ago, at least 10% of the E. coli genome has been acquired in somewhere in excess of 200 horizontal gene transfer events.
In an equally insightful commentary, William Martin [Martin 1999] has discussed the implications of this work for our ability to reconstruct phylogenetic trees:
- The further back in time an evolutionary divergence, the greater the likelihood that any given gene in a genome has been transferred.
- Indeed, it may be the case that all bacterial genes have been subject to horizontal gene transfer at some point in their evolutionary history.
- This could undermine the utility of phylogenetic tree reconstruction for deep divergences.
Currently, there is a lot of debate over whether gene transfer is so rampant that evolutionary trees cannot be built, or whether the levels of gene transfer are negligible. Both extremes are currently championed in the literature, and ironically, when it comes to the LUCA, Carl Woese's work is central to both — many of those that view Woese's three domains as correct have been arguing for little or insignificant levels of transfer, whereas Woese has recently suggested that very early in evolution, gene transfer between organisms was more important than inheritance from generation to generation [Woese 1998].
While it sounds like Woese is being inconsistent, his more recent claim is limited to the earliest periods in the evolution of life, and arose from concerns that LUCA was beginning to appear totipotent, a crazy notion that would have LUCA as the ultimate source for all life's diversity:
- A fertilised egg is totipotent — from a single cell it will develop into all the different cells and tissues that make up an adult human being.
- If genes move between organisms, LUCA might mistakenly appear totipotent because many features would be incorrectly counted as universal.
Extrapolating Lawrence & Ochman's result back billions of years may not be realistic. But what if horizontal transfer was the default state? This is the idea Woese has developed. His argument is that genes were so free to exchange that there were no distinct lineages — genes moved more through horizontal transfer than by vertical inheritance. As the genetic system becomes more accurate and as the complexity increases, more genes become interdependent, and transfer gives way to vertical inheritance. Woese argues that translation (and therefore the genetic code) was the first thing to be fixed or crystallised, with other cellular functions following later. From this horizontal transfer dominated system, the three domains (archaea, bacteria and eukaryotes) each emerged independently as lineages.
This is certainly food for thought, but there are several issues:
Gathering evidence to support it is not exactly easy since there's no real way to establish that horizontal transfer was the initial state.
Koonin's shrinking minimal gene set (Koonin 2000), shows that the minimal genome approach is not creating a totipotent LUCA. Instead, the number of genes ascribable to LUCA is becoming smaller as more genomes are added.
Another problem has to do with the switch from horizontal transfer to vertical inheritance. How many genes would have been able to partake in global transfers before becoming crystallised and therefore unable to transfer? Would it really have been complex enough for the ancestors of the three domains of life to have emerged independently as distinct lineages?
Gene transfer is going to be a hotly debated topic for a while, and will continue to confound the reconstruction of the LUCA. The problem is a complex one:
Horizontal gene transfer has been demonstrated — e.g., the spread of antibiotic resistance.
Limitations of the methods for building evolutionary trees can give false evidence for gene transfer.
Methods that don't make use of evolutionary information are being used to examine genetic relationships and, in many cases, the data that have been used to argue for horizontal gene transfer are weak.
There is little consensus on the reliability of methods for detecting horizontal gene transfer.
What data are required to demonstrate ancient horizontal gene transfer events?
If natural selection is considered, most horizontal gene transfers will probably result in the gene being lost — by analogy, the organism needs a new gene like a fish needs a bicycle! For instance, antibiotic resistance genes won't spread and be maintained by selection unless the organisms with the genes are being assaulted with the antibiotic.
This last point has been too often ignored, and there has been little attempt to establish patterns (e.g., are all genes equally nomadic?). So how should LUCA biologists deal with horizontal gene transfer?
If we accept that there is or has been massive unbridled horizontal gene transfer between the three domains [e.g., Doolittle 1999], we must conclude that all our tools for looking into the evolutionary past are invalidated, which means we might as well give up on the question of the LUCA. We know that there are demonstrated cases of horizontal gene transfer, but this extreme position is like throwing the baby out with the bathwater.
If we take as our starting point the opposite extreme, that the effect of horizontal gene transfer has been negligible, we are in a much better position — we still have our tools in place, and any suggestions of horizontal gene transfer will need to be backed up with good evidence.
There is no doubt a middle ground can be found, but amidst the furore over horizontal gene transfer, a number of researchers, making use of whole genome sequences, have reported results suggesting gene transfers have minimal effect on the ability to recover evolutionary trees [e.g., Snel et al. 1999; Sicheritz-Pontén & Andersson 2001]. These results suggest that it is possible to reconstruct the tree of life, and moreover, conclude that the 3 domain structure of the tree, as first reported by Woese and his colleagues, is supported by whole genomes.
In a timely article, Chuck Kurland has firmly criticised the eagerness of many to attribute horizontal gene transfer [Kurland 2000]. One particularly interesting aspect of his exposition is that he suggests a number of non-scientific factors that have contributed to the hype around horizontal gene transfer, and is as much a comment on how science currently operates as it is about gene transfer.
The root of the tree of life is hard to pin down [Pennisi 1999], and unbridled horizontal transfers early in the evolution of life can't easily be distinguished from the limits of the sensitivity of our phylogenetic tools — that researchers have failed to reach a consensus on the shape of the tree does not mean that there must therefore have been horizontal transfer.
Indeed, there is another issue here — the reliability of the methods used for building evolutionary trees. Many researchers are very confident of the reliability of these methods, yet it is well known that these are based on mathematical algorithms which are convenient, but which do not necessarily accurately model real biological sequence evolution. These methods are likely to be robust for recent evolutionary events, and are definitely the most robust of the methods for detecting gene transfers. The problem is that near the root of the tree of life, they may be just too inaccurate to be useful for scrutinizing the very earliest events in evolution. In a worst-case scenario, the situation might in fact be a bit like timing the 100M sprint at the Olympics with a sundial!
David Penny, Bennet McComish and their coworkers have recently tried to address this question by investigating how far back in time the standard models used in evolutionary tree building can go before they start to go wrong. Their overall conclusion is that the models used do seem to do a little better than might be expected from theory, but that the models still do poorly for very early evolutionary events. Penny and colleagues also criticise the recent trend in reporting conflicting trees as evidence for horizontal gene transfer — given how hard it seems to accurately reconstruct the tree of life, it is hard to say whether conflicting answers are evidence for gene transfer, or just reflect the limitations of the methods for building the trees. Their testing of the models suggests that it is just not reasonable to say that there is horizontal gene transfer just because two trees made with two different genes don't come back with the same relationships between organisms. They make the following comment, which sums up the problem very succinctly:
"… there are major difficulties between data sets for ancient divergences. It is difficult to see why researchers are so confident in their results when the relatively recent divergences within mammals, birds, or flowering plants are only now being resolved."
This work of Penny et al. [2001] and the picture coming from evolutionary trees of whole genomes [Snel et al. 1999; Sicheritz-Pontén & Andersson 2001] seems to bolster Kurland's provocative assertion that horizontal transfer is 'an ideology that is begging for deconstruction.'
Nevertheless, horizontal gene transfer does occur to some extent — Lawrence & Ochman's 1998 paper is but one of many demonstrating this. Moreover, many of the technologies biologists use for inserting genes are simply human exploitation of what has been described as natural genetic engineering. The following naturally occurring mechanisms of 'genetic engineering' are routinely used in molecular biology laboratories:
- Plasmids: small, usually circular, pieces of DNA that often carry genes that enable them to move from one bacterium to another.
- Viruses: many will naturally insert themselves into the DNA of the organism they are infecting, and can be engineered to carry extra pieces of DNA.
- Natural or assisted DNA uptake by bacterial cells.
- Restriction endonucleases: molecular scissors that allow precise 'cutting' of DNA.
We also know it is possible to identify ancient gene transfers that may have occurred as far back as 2 billion years ago. Biologists can readily identify genes in the eukaryote repertoire that have come in via the mitochondrion, a compartment in the eukaryote cell which is bacterial in origin. Indeed, the handful of genes remaining in this compartment have been shown to be bacterial in origin, as have some that have since taken up residence in the eukaryote nucleus [Lang et al. 1999].
Returning for a moment to the biology of nomadic genes, the consensus emerging from studies of bacteria is that we should indeed start thinking of bacterial (and perhaps archaeal) genomes as being an ever changing collection of genes, but only to a degree [Hacker & Carniel 2001]:
- Genes which are central to the running of any cell — these are often referred to as housekeeping genes — make up the 'core' genome.
- Genes which come and go make up the flexible genome.
The flexible genome might be a window into the nomadic gene pool of bacteria — Lan & Reeves [2000] point out that closely related strains of bacteria differ in the genes they carry by as much as 20%, and this requires we reevaluate how we categorise species of bacteria. The hope is that the flexible genome can tell us how a particular bug is currently making its living. Losing genes isn't in itself particularly surprising for organisms that are in competition to reproduce as fast as they can [Jeffares & Poole 2000] — genes that aren't being used aren't kept. 'Use it or lose it' is the maxim of natural selection, and there are plenty of examples wherever you look in biology (our appendix and tail bone both appear to have headed in that direction for instance).
If this picture of core and flexible genomes is correct, it is good news for LUCA research because many universal features can in theory be recovered. This goes too for the ancient horizontal transfers seen with the mitochondrion. We should be optimistic that some patterns of horizontal gene transfer can be analyzed, though we still need to exercise care when looking so far back in time.
4. Fusion
When it comes to horizontal gene transfer, the hype about archaea and bacteria is arguably a case of squabbling over crumbs when compared to what seems to have happened in the early evolution of the eukaryotic cell. The now popular idea that eukaryotes emerged from a massive fusion (the ultimate gene transfer) event between a bacterium and an archaeon is also raising problems for LUCA biology.
While fusion is all about the origin of the eukaryotes, it is also about LUCA:
- Fusion scenarios challenge Woese's division of the living world into three domains.
- Rather than a tree with three branches all tracing back to the LUCA, fusion has two lineages, with the eukaryotes emerging by fusion.
- Fusion is in conflict with the emerging picture of the direct link between eukaryote biology and the RNA world.
A number of researchers have argued that the genes in the average eukaryote look to be a mixture of bacterial-like and archaeal-like. That is to say, at the genetic level, eukaryotes look to be some sort of genetic fusion between archaea and bacteria [Ribeiro & Golding 1998; Rivera et al. 1998; Horiike et al. 2001].
In understanding this, it has been helpful to divide genes into two categories: informational or operational [Rivera et al. 1998]. Informational genes are those which are involved with the copying, storing and regulation of genetic information, while operational genes are the recipes for making proteins for synthesis and breakdown of molecules in the cell, and are largely involved in energy metabolism.
Consistent with earlier research [see Gupta & Golding 1996] Rivera and colleagues found that there was rhyme and reason to the mixture of bacterial and archaeal genes in eukaryotes:
- For informational genes — archaea and eukaryotes share more in common
- For operational genes — bacteria and eukaryotes share more in common
Mark Ridley [2000] has suggested a good analogy for what many think has happened — a business merger. Instead of doubling up and having two departments for every aspect of the new company (Eukaryote Inc.), only one of each was kept, with the result being that the informational department came from Archaea Inc. and the 'operational department' from Bacteria Inc.
With the ongoing debate on how much horizontal gene transfer there is between organisms, the most exciting contribution to this picture looks not at the genes, but at gene networks. Taking a page from the study of complex networks such as the Internet, Eörs Szathmáry and colleagues [Podani et al. 2001] have recently shown that, while eukaryotic operational genes appear bacterial in origin, the structure of the metabolic network that these genes make up is in fact much much more like what is observed in archaea. In keeping with the business merger analogy, this is perhaps equivalent to keeping the management structures of Archaea Inc. in place.
This is an exciting picture, and there is no question that modern-day eukaryotes are the product of some sort of fusion [Ribeiro & Golding 1998; Horiike et al. 2001]. However, the tricky thing is working out what it all means for the origin of the eukaryotic cell. These are some of the outstanding issues:
- Why has such a merger apparently only happened once?
- No one has ever observed modern bacteria and archaea fusing.
- Why is it we don't see 'anti-eukaryotes' (that is, organisms which have the operational genes of archaea and the informational genes of bacteria)?
- A number of features found exclusively in eukaryotes are tricky to explain by a fusion event.
Indeed, there are a number of ways of explaining the fusion data, and consequently, there are quite a few different opinions on how the eukaryotes came to be [Minkel 2001].
If eukaryotes are the result of a fusion between a bacterium and an archaeon, then the 3 domain picture that Carl Woese's work supports would be wrong. Fusion would imply that everything in eukaryote biology is either a recent innovation specific to this domain, or an offshoot of the biology of archaea and bacteria. In other words, if you want to know about LUCA, archaea and bacteria are the only two domains worth looking at. This is an assumption that is often made, regardless of fusion, and a point against which some researchers, myself included, have argued [see Jeffares & Poole 2000; also Forterre & Philippe 1999; Poole et al. 1999].
To make sense of the motivation behind the many emerging fusion scenarios for the origin of the eukaryote cell and how these might impact on LUCA biology, it helps to concentrate on the big picture, rather than wading through the details of the various scenarios. Laura Katz [1998] has written a good overview of the various fusion scenarios, though several new scenarios have been published since then [e.g., Margulis, et al., 2000; Horiike et al. 2001; Bell 2001; Hartman & Fedorov 2002]. Fusion theories have developed out of the endosymbiotic theory for the origin of the mitochondrion:
- The endosymbiotic theory was first formulated by Mereschkowsky at the beginning of the 20th century, but reintroduced and updated by Lynn Margulis in the 1970s [Martin et al. 2001].
- This theory argues that the mitochondrion, sometimes called the powerhouse of the cell, was originally a bacterial cell that took up residence in the ancestor of modern eukaryotes.
- Both structural and genetic similarities have shown without a shadow of a doubt that the endosymbiotic theory is correct — the DNA in the mitochondrion is more closely related to bacteria than to the DNA stored in the eukaryotic cell nucleus.
- It is now widely accepted that this event happened once only.
Despite much agreement, there is ongoing debate surrounding the endosymbiotic theory:
- How was this partnership founded (e.g., oxygen-based or hydrogen-based metabolism)?
- Was the host that ultimately engulfed the bacterium a eukaryote or an archaeon?
The first question opens up a whole can of worms (which we'll avoid here), and is a current source of intense debate [Andersson & Kurland 1999; Rotte et al. 2000]. The second question is what has the major impact on LUCA biology, but these two questions have been unnecessarily muddled. The bottom line is that the genomic & gene network data supporting fusion between an archaeon and a bacterium can as easily be made to fit a fusion between a eukaryote and a bacterium.
The state of the field is as follows:
- Everyone agrees that the mitochondrion evolved from a bacterial ancestor (though there is current debate as to what the bacterial ancestor was like, and how it interacted with its host).
- There is disagreement as to whether the host was a eukaryote, or an archaeon.
- Archaea-Bacteria fusion hypotheses require all genes found only in eukaryotes to have arisen post-LUCA, post-fusion — that is, they are indirectly descended from LUCA.
- This comes into conflict with the picture of LUCA from RNA [Jeffares & Poole 2000], and Woese's tree of life [Woese et al. 1990] — both require that eukaryotes were directly descended from LUCA.
So how do we distinguish between an archaeal and a eukaryotic host? The key is in two parts — one is historical and the other requires careful thought about how archaea and eukaryotes might be related:
- The historical aspect centres around understanding the shift in thinking from the original picture of an ancient eukaryote playing host to the now largely agreed-upon picture of an archaeon playing host. This shift largely revolves around the changing branches in the eukaryote evolutionary tree [Dacks 2002].
- The relationship between archaea and eukaryotes cuts to the heart of how researchers view the evolution of cells.
Archaezoa - missing links lost
So why is it that fusion hypotheses have become so popular? Indeed, this goes against the classical interpretation, most thoroughly espoused by Tom Cavalier-Smith [1987], who identified a disparate group of eukaryotes that appeared to him to be missing links — the so-called Archaezoa, which look like eukaryotes but lack mitochondria. His hypothesis, that the Archaezoa evolved before the introduction of mitochondria into the eukaryote lineage, held sway for many years, though has recently been dropped in favour of fusion:
- There is growing evidence that all eukaryotes once harboured mitochondria
- Thus, the Archaezoa have probably all lost their mitochondria, rather than never having had them [Embley & Hirt 1998].
- For instance, one group of the Archaezoa called the microsporidia are now widely accepted to have been incorrectly placed very deep on the eukaryotic tree. Indeed, probably most of the Archaezoa, if not all, are incorrecly placed on the tree. Rather than being missing links leading back to the origin of eukaryotes, they probably arose more recently [see Dacks & Doolittle 2001; Keeling 1998; Dacks 2002].
- If the Archaezoa aren't a series of missing links, the origin of eukaryotes may have been concurrent with the endosymbiosis that gave rise to mitochondria.
The conclusion from the above is that all eukaryotes probably had a mitochondrion, and without the Archaezoa, the only ancestor of eukaryotes is archaea. Voilà! We have fusion.
A case of throwing the baby out with the bathwater?
The important point to keep in mind about the picture for fusion is that it is a partial one, based largely on gene data. There are a large number of differences in the general structure of eukaryotic and prokaryotic (archaeal & bacterial) cells that aren't explained by fusion [Poole & Penny 2001]. However, the major inconsistency is that the picture provided from trees is not the same for the relationship between archaea and eukaryotes, and that of bacteria and eukaryotes [Poole & Penny 2002, submitted]:
Margulis' hypothesis is evidenced from trees. There is now overwhelming agreement that the mitochondria branch is within the bacterial tree, specifically within a subgroup called the alpha-proteobacteria [Lang et al. 1999], and a bacterial origin is also observed for chloroplasts (where photosynthesis takes place in plants and other photosynthetic eukaryotes).
Comparisons of relevant genes from eukaryotes and archaea should give this picture also, yet the evidence points to archaea and eukaryotes being very distinct domains.
This has strong parallels to the way the Archaezoa case is being treated — if there are no modern groups of archaea that appear to have split from the trunk of the tree of life before the appearance of eukaryotes, should we accept fusion? Stronger evidence was certainly required in testing the origin of the mitochondrion!
Another issue has to do with missing links. If the disappearance of the missing links (the Archaezoa) is used to suggest fusion, it is surely just as reasonable to argue against fusion on exactly the same grounds — there are no intermediates between eukaryotes with mitochondria and the archaea. For example, we don't see examples of archaea with mitochondria in them, or archaea with nucleus-like structures.
With perhaps a couple of billion years separating the divergence of archaea and eukaryotes, it would be incorrect to require that the archaeon in the fusion must have been just like modern archaea. This cuts right to the heart of the problem — there is no inherent requirement that evolution leaves behind a series of intermediates for us to use to piece together the different evolutionary trajectories of archaea and eukaryotes. As with Chinese whispers, the end point may be very different from the starting phrase, but with evolution, all we have to look at is a number of different endpoints, from which we can only guess at the starting phrase!
While the specifics of the Archaezoan hypothesis are most probably wrong, it should not be thrown out completely. Explanations of eukaryote origins by fusion or via Margulis' original scenario each suffers from the disappearance of intermediate forms, but this is expected. As Stephen Jay Gould had often said, evolution results in bushes, not ladders.
A number of researchers [Forterre & Philippe 1999; Andersson & Kurland 1999; Penny & Poole 1999] maintain that the data for fusion can be reconciled with Lynn Margulis' endosymbiotic theory and Carl Woese's three-domain tree. Indeed, David Penny and I have argued that fusion does the worst job of explaining the available data [Penny & Poole 1999; Poole & Penny 2001]. For instance, fusion doesn't fit with the hypothesis that some eukaryote features, which have since been lost in archaea and bacteria, actually date back to the LUCA (see Jeffares & Poole 2000).
What it very tentatively implies is that archaea and eukaryotes may have shared a more recent ancestor than either shares with bacteria, as is often shown in textbooks, but this too is not certain, since the relationships between these three groups is also a point of controversy [see Forterre & Philippe 1999; Pennisi 1999]!
5. Conclusions
We are now entering a very exciting period in uncovering the history of the LUCA — the field has been given a major boost from a broader range of ideas being applied to the problem:
Acknowledging the technical challenges with building the tree of life is an important step in the right direction. So is the idea of using the RNA world period in the origin of life for establishing aspects of the nature of the LUCA (see Jeffares & Poole 2000).
Despite the deluge of genome data available, it is hard to say whether we will ever actually manage to get a complete list of genes which LUCA possessed. We may pick out certain characteristics, but each needs to be evaluated with extreme care. The minimal genome study of Mushegian & Koonin [1996] demonstrates this and, in its failure, it stands as a strong caveat. Most researchers have toyed with this idea, and many were probably disappointed (then later relieved) that Mushegian & Koonin beat them to it!
Horizontal gene transfer is likely to be a factor in confounding such efforts, but it is better to err on the side of caution with respect to how pervasive this is in the history of life. The emerging picture from genome research suggests that not all genes transfer equally easily, and that there may be an ecological underpinning to the nature of gene transfer.
The fusion hypothesis has important consequences for the LUCA — if correct, the LUCA must have been like bacteria and/or archaea, because those unique features of the biology of eukaryotes had not yet evolved.
The three domain tree that emerged from Woese's original work permits features of all three domains to trace back to the LUCA while the fusion hypothesis, in its strictest form, does not. Unless an argument for loss of a feature in all modern archaea can be made, it is diametrically opposed to the nature of the LUCA as suggested from RNA world fossils [Jeffares & Poole 2000].
Currently, many major assumptions are being questioned:
- Were there three domains or two, with the third arising by fusion?
- Was LUCA prokaryote-like or eukaryote-like or even a mixture?
- Is the genetic code the only one possible?
- Was early evolution more reliant on horizontal gene transfer than inheritance?
- Was there one or more LUCAs?
Each of these questions could easily fill a book, and it has become impossible to cover every aspect of LUCA biology in one article. To the casual observer, the field of LUCA biology looks to be in disarray, with everyone having their own pet theory. This can be exciting, frustrating, and, at times, bordering on the absurd, but above all it is a sign of healthy debate! Many views and varied approaches to the problem means some exciting answers to some fundamental questions about life's origins are just around the corner…
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Appendix A: Making Protein
To make a protein, the information stored in DNA language must be translated into the language of proteins, where, instead of a four-base alphabet, the alphabet is made of 20 amino acids. Translation is predominantly an RNA-affair, with a working RNA copy of the DNA gene (called messenger RNA, or mRNA) being read by the ribosome, the protein synthesis factory found in all cellular life forms. The actual translating from DNA/RNA language into protein language involves an RNA called transfer RNA, or tRNA for short.
tRNA is the key to understanding the nature of the genetic code. In making an RNA, each of the bases A,G,C,U is a word in itself, but in order to go from 4 bases to the 20 amino acids that make up proteins, the bases are grouped into 'words' three bases long (called codons), making a dictionary of 64 possible words. In this way, all 20 amino acids can be described in three-letter or triplet code, with room to spare. 61 combinations are used (the remaining three are read as 'STOP MAKING PROTEIN'), and an important feature of the code is redundancy — for instance, there are 6 codons that code for the amino acid leucine. The consequence of such redundancy is that the translation process is less susceptible to errors than a one-to-one code.
Translation requires that each tRNA is charged at one end with an amino acid. At the other end, an anticodon matches up with one of the codons that match that amino acid. Translation involves docking charged tRNAs at the appropriate point on the mRNA as the ribosome moves along, reading the triplets off. When the 'correct' tRNA is in place, the ribosome joins the amino acid carried by that tRNA to the growing chain of amino acids, which together make a protein.


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India is witnessing an unprecedented economic boom and this is visible through online shopping and E-commerce. Nearly 50% of the population in urban areas now shop online and this number is increasing day by day. It is a win-win for both buyers and sellers as consumers save time and efforts by shopping online and sellers have access far and wide. More and more merchants are realizing the enormous potential of the medium of Internet and are gearing up to offer products and services.
"When everything is available at our doorstep, why going and shopping outside?" say many youngsters. India, being a country of young, this mind-set of youngsters point towards the huge potential online business has here. Cashing on this new craze amongst the youngsters, the internet has become a shopping mall in itself and online shopping has become a breeze in the recent times. The outlet stores, which used to be jam packed during festive season, now bear a deserted look as the best deals and offers are available online. You can also save your money, especially when you couple it with online savings codes, deals of the day, and free shipping offers.
When you order something online, you often come across a text box where you need to write a promo code for some extra discount in case you have one. Websites like Couponhaat keep offering these online coupons and promotions on a regular basis. But often, you find that you don't have one and then you feel bad about not getting that extra discount. There's no need to worry as several websites are there that searches consumer emails, forums, blogs, merchant sites, and newspapers for any coupon codes from online retailers. So, the next time you shop online, go to these websites and get a coupon code from there to help you save a little bit of extra money the next time you shop online simply by typing in a code. For instance, if you want to shop on Jabong, look out for some great offers with Jabong coupons.
Thousands of online coupons and deals are available online on a regular basis. The same product can have different deals on different sites. This sometimes confuses the consumer as he is not sure if he got the best deal or not. To help buyers with this, many online sites are there, which list the best deals available online across the stores. By clicking on the deal on these sites, you are directed to the merchant website to complete the transaction. This saves you time and you get the possible deal available.
These are added benefits you get for being a bit more vigilant. The normal boons of online shopping are already there like saving time and energy as you can shop at your convenience. You can compare different products and prices just by a click of a mouse.

subset: commoncrawl


text:
hes 2 weeks from superb bit i guess he still has a year of training left
hes 2 weeks from superb bit i guess he still has a year of training left
Munroe from Central Districts is superb bat comp stam comp conc. And can open.
If not, Mcmillan is a strong/prof bat with strong stamina (strong/prof bowling too but obviously wouldn't make it). About 75% to Superb.
1. A Munore - superb bat
2. G Ashton – superb bat, superb keeping
3. C Allen - remarkable bat, quality bowl
4. T Kennedy - superb bat
5. G Canning - superb bat, quality bowl
6. K Challinor – superb bat
7. J Thorp – superb bat
8. R Mcmillan - strong bat, (Backup: strong bowl)
9. K Sekani – quality bowl
10. G Bardle - remarkable bowl
11. A Islam – superb bowl
- Munore added, i thought he might be a chance, can Smith post his stats, so i can have better look.
- Mcmillian added aswell, just waiting on another superb bat so he can drop down to eight. Probably will be Pak.i cus i think he is superb but im not sure.
- T Kennedy added and some changes to the batting
Last edited by chaminda_00; 24-09-2006 at 05:54 AM.
The man, the mountain, the Mathews. The greatest all rounder since Keith Miller. (Y)
Jaffna Jets CC (Battrick & FTP)
RIP WCC and CW Cricket
Member of the MSC, JMAS and CVAAS
I've got a Superb 19 yo, he should be quality soon but his secondaries aren't much.
Stats please...Originally Posted by The Baconator
Got Spin.Originally Posted by ash chaulk
Theres my superb 19 yr old bat
Tony Kennedy - 19 yo, BT Rating=16,423
RH Batsman, RM Bowler, superb batting form, superb bowling form, energetic
A destructive player with worthless leadership skills and worthless experience.
Stamina: mediocre Wicket Keeping: woeful
Batting: superb Concentration: strong
Bowling: worthless Consistency: worthless
Fielding: worthless
Brenoevo, what are Jayasuriya & Kennedy like these days?
Wayne Stevens - 18 yo, BT Rating=25,590
RH Batsman, RFM Bowler, superb batting form, superb bowling form, low
A defensive player with abysmal leadership skills and abysmal experience.
Stamina: strong Wicket Keeping: feeble
Batting: mediocre Concentration: competent
Bowling: superb Consistency: proficient
Fielding: feeble
He'll make it next season, let's say that much.Originally Posted by ash chaulk
I thought he might have popped to superb, might get infront of Islam, on batting, but then again do u really need a mediorce bat at 9 or 10 with an extra batsmen at eight.Originally Posted by ash chaulk
There both quality bats with superb conc and prof stamina. Iv stopped training them now cause i pulled a resp 17yr old bat at the start of the season and can see more potential in training him as by the time he turns 20 with 2 bat nets he'll be around exceptional.Originally Posted by Robertinho
no you dont need another batsmen if it was real lifeOriginally Posted by chaminda_00
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text:
Framework: HI025 – hi025_0808_v2
8 - Primary or Horizontal Divisions
8 - Secondary or Vertical Divisions
Part of series: HI025 Hierarchy 025 – Spiral Staircase
Series Keywords: spiral, stairs, steps, climb, ascend, descend
This staircase can show a progression of steps or levels, but because it pivots around a base so the steps can relate directly to a goal, project, organization, or other. This makes the staircase in this series able to convey different concepts than a stand-alone, straight staircase.
There are two versions of staircases. Each of the two versions has variations of 1 to 10 stairs:
- Version 1 (v1) contains staircases that pivot on bases of different heights, e.g., the 3-step staircase pivots on a base that is the height of the third step.
- Version 2 (v2) contains animated staircases that build to as many as 10 stairs and pivot on a base that is the height of the top stair, e.g., if the staircase builds to 5 stairs, the height of the base is as high as the fifth stair.
The text on the stairs in the series example is done in PowerPoint 2007. The text has been rotated to match the surface of the stair. PowerPoint 2003 does not provide this formatting option. Notice that there is a PowerPoint 2007 animated example for this series. If you are working in PowerPoint 2007, you can download this file and you will have the prerotated text to use in your slides with spiral staircases.
The text for the base (cylinder) is WordArt that bends around an object. This is possible in both versions of PowerPoint. You can download the animated example and use the WordArt there as a starting point if you wish.
Refer to the tutorial for added information about formatting and usage. There are some layering techniques that will be helpful to understand when working with v2 staircases.
Key search words: spiral, stairs, steps, climb, ascend, descend


text:
Estate Tax Distributions to all Governmental Units
for Settlement Periods during Calendar Year 1994
The Tax Commissioner administers Ohio's estate tax, but the
tax is locally collected by the treasurer of the county in
which the decedent resided. Payment by the estate
administrator is due no later than nine months from the date
of the decedent's death. After the semi-annual settlements,
each county remits 36 percent of the tax collection to the
state, less various administration fees. The remaining
portion is distributed to local jurisdictions based on the
percentage of estate assets held there.
Total estate taxes distributed to all local entities amounted
to $155.6 million in calendar year 1994, excluding
distributions for Medina and Wyandot counties. The amount
remitted to the state amounted to $85.2 million in the same
time period. The distributions range from a high of $13.5
million in Cincinnati, Hamilton County, to a low of -$56,207
in the Village of Cadiz, Harrison County.
When an estate has property in more than one county, the
county that the return was filed in must distribute the taxes
due to those jurisdictions outside of that county. In
reporting, most counties cite each jurisdiction outside of
their county along with the amount distributed to them. Some
counties, however, just make an "outside-of-county" entry and
do not specifically list these jurisdictions. This type of
entry is accounted for and labeled "unidentified" at the end
of this table.
The figures shown are compiled from a survey of county
auditors conducted by the Ohio Department of Taxation.


text:
First Quinta wireless conference system installed in Vietnam
The Thang Long Investment and Trade Company (TLSC), has recently installed the Quinta high-class wireless conference system for an affiliate of the Ministry of Public Security with its partner K&T.
It was the first installation of the new, fifth generation wireless conference system Quinta in Vietnam.
beyerdynamic is a world-class manufacturer based in Germany with over 88 years of experience in the audio industry. The product range comprises microphones, audio headsets, microphone headsets for TV commentators as well as conference and interpreter systems.
The future-proof Quinta system ensures an interference-free and secure transmission in three available frequency band (2.4/ 5.2/ 5.8 GHz). A switching of the channels is either automatically or manually, in any case absolutely silently. The system can be integrated into an existing or new building computer networks that comply the IEEE 802.1 IT standard (AVB). The numerous functions with which the meeting can be controlled, include the simple operation and configuration via Smartphone or tablet PC.
For the chairman, delegate and double delegate microphone units there are gooseneck microphones with XLR connector in four different lengths available. The solid housing and the robust, scratch-resistant paint give the microphone units a high-quality look. In short with Quinta beyerdynamic emphasizes its high quality "Made in Germany".
The TLSC, which was established in 2004, is currently the official partner and distributor of conference technology for beyerdynamic in Vietnam.


text:
Werth, LaRoche rally Nats for walk-off win vs. Halos
WASHINGTON -- It looked like the Nationals were going to lose their third straight game, but big hits by Jayson Werth and Adam LaRoche helped them walk off with a 5-4 win over the Angels on Wednesday night at Nationals Park. It was their seventh come-from-behind victory this season.
The Angels had a 4-1 lead going into the bottom of the ninth inning, with closer Ernesto Frieri on the mound. Several Nats players went over to shortstop Ian Desmond and asked how to hit Frieri. In Monday's 4-2 loss, Desmond hit a ninth-inning home run against the right-hander.View Full Game Coverage
WASHINGTON -- It looked like the Nationals were going to lose their third straight game, but big hits by Jayson Werth and Adam LaRoche helped them walk off with a 5-4 win over the Angels on Wednesday night at Nationals Park. It was their seventh come-from-behind victory this season.
The Angels had a 4-1 lead going into the bottom of the ninth inning, with closer Ernesto Frieri on the mound. Several Nats players went over to shortstop Ian Desmond and asked how to hit Frieri. In Monday's 4-2 loss, Desmond hit a ninth-inning home run against the right-hander.View Full Game Coverage
"Desi was just running his mouth, so that kind of got us going," Werth said. "It's good. I like the way we are playing."
Frieri couldn't close the game, allowing Jose Lobaton's leadoff homer. Then -- after striking out Zach Walters -- Denard Span singled and Anthony Rendon followed with a walk.
That brought up Werth, who was ahead in the count, 3-0. Manager Matt Williams gave him the green light to swing.
"I trust him. He is a veteran player. I trust the fact he has done it a couple of times this year. One was a home run. The other was a double," Williams said. "He is comfortable with it. He knows what pitch to pick, and how to go about doing it."
Werth picked a fastball and doubled down the left-field line, scoring Span and Rendon and tying the game at 4.
"It's one of those situations where if you hit into a double play right there, it's probably the worst play you have ever seen. If you get a hit, it's the best. So I'm glad it worked out, that's for sure," Werth said. "When I played for Charlie Manuel, he always talked about aiming for the foul pole. I was just aiming for the foul pole. It worked out."
Frieri was mad at himself for not being able to save the game for Los Angeles.
"I'm mad at myself, because I keep missing pitches, and if I keep doing that, I'm going to keep getting hurt like that," he said. "I don't know, man. I'm fighting, I'm trying to get better. But if I don't make my pitch, I'm going to get hurt. Even when I'm ahead in the count, I can't be leaving balls down the middle, because I'm going to get hurt."
Frieri was taken out of the game in favor of Fernando Salas. LaRoche then came to the plate, swung at the first pitch and singled to left-center field to score Werth for the game-winner.
"He left a fastball over the plate," LaRoche said. "In that situation, I was trying to hit something hard. He has a good changeup. I can't really sit on the fastball. I tried to put a good swing on something.
"The damage was really done before my at-bat. Lobaton comes up and puts a run on the board quick. I can't imagine anybody thinking [Werth] was going to swing 3-0 right there. It surprised all of us. He came up huge. Big win, needless to say. We needed that one. "
Early in the game, Gio Gonzalez provided the only offense for Washington, and it occurred in the second inning, when he singled to center field off right-hander Jered Weaver, scoring Danny Espinosa and giving Washington a 1-0 lead.
"A blind squirrel finds a nut once in a while," Gonzalez said. "It was one of those lucky swings. I was out in front. Got lucky. It 's one of those things we can smile about and turn the page."
Entering the sixth, Gonzalez had retired 10 consecutive batters and appeared to be on his way to a quality start. But before long, the Angels had a 2-1 lead and Gonzalez was on the hook for a loss.
Mike Trout walked to start the sixth. Albert Pujols followed and doubled down the third-base line, scoring Trout to tie the score at 1. Williams then took Gonzalez out of the game in favor of Aaron Barrett, because Gonzalez was experiencing tightness in his left shoulder. It was cold, and Williams didn't want to take any chances. The injury is not considered serious and Gonzalez is expected to make his next start against the Phillies next week.
"After the fifth inning, it started getting a little colder. It felt a little tight," Gonzalez said. "Matt started seeing it in the sixth inning. I feel fine now. I did all my exercises. I feel great. Next question."
Barrett was able to get Howie Kendrick to ground out. However, Erick Aybar singled to left field, scoring Pujols.
In the next inning, David Freese scored on a wild pitch by Barrett.
A strange play occurred in the sixth inning, when after LaRoche singled to lead off, Bryce Harper tried to bunt for a hit on a 2-2 count, but he fouled out to catcher Chris Iannetta.
Williams defended Harper, saying the Angels gave him the opportunity to bunt.
"I'm not opposed to him laying a bunt down with two strikes. We've seen guys do it before," Williams said. "If he gets that bunt down, it's a base hit. I'm not concern about that. He is trying to do things to help us win."
Weaver pitched well enough to win the game. He pitched six innings, allowed the one run on seven hits, and struck out two batters.
Gonzalez was charged with two runs over five-plus innings, while Drew Storen earned the win with one-third of an inning pitched.
"If they showed anything, it is that they fight. It gives [us] more opportunities. We got a couple of big hits from the middle of the order," Williams said.
Bill Ladson is a reporter for MLB.com and writes an MLBlog, All Nats All the time. He also could be found on Twitter @WashingNats.


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Giancarlo Mazzanti Arquitectos, Bogota Architects
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Giancarlo Mazzanti Arquitectos, Bogota Architects
Giancarlo Mazzanti Arquitectos, Bogota Architects Office, Colombian Design Studio
Giancarlo Mazzanti : Bogotá Arquitectos
Contemporary Colombian Buildings, South America: Interview & Film News
14 Aug 2019
Architect Giancarlo Mazzanti
New interview with Bogotá-based architect Giancarlo Mazzanti
Giancarlo Mazzanti on Social Spaces for Learning
Plane-Site's production team traveled to Bogotá to speak with Giancarlo about his experimental approach to design research and learn more about his ?spaces for learning?.
Many of these projects are situated in low-income neighborhoods and act as catalysts for social change, bringing joy and pride to marginalized communities:
Giancarlo Mazzanti on Social Spaces for Learning from PLANE—SITE on Vimeo.
Giancarlo Mazzanti – Major Building España Library, Santo Domingo, Colombia
Date built: 2007
photo : Sergio Gómez
Biblioteca Pública Santo Domingo
The Project is located on one of the hillsides of the city that have been affected by harsh violence since the 80´s because of several drug traffic networks that operate within its perimeter. It is part of the government's social master plan program that aims to give equal economic and social opportunities to the population.
Giancarlo Mazzanti – Key Projects
Major Projects by Giancarlo Mazzanti Arquitectos, alphabetical:
El Porvenir School, Bogotá, Colombia
Date built: 2010
Habitar 72 + Habitar 74 : residential buildings, Bogotá
Leon de Grief Lib...
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Livingston County Center at Parker Campus, Howell (LIVCEN)
402 Wright Road
Howell, MI 48843
Directions Livingston County Center Website
(517) 545-3522
With over 100 courses every fall and spring semester, the Livingston County Center at the Parker Campus is a great place to pursue your college studies. Classes are smaller, making it easier to get to know your classmates and instructors. For registration assistance, just stop in. Visit Livingston Center for hours of operation and services.
Student Affairs Division
Gannon Building
2nd Floor - StarZone
Phone: (517) 483-1957
Additional contact information »


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Janell Christiansen
Jimmie was Funny and nice people from calling in to payment
Jimmie was Funny and nice people from calling in to payment
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Nobody wants to have an HVAC emergency that requires them to replace it. To extend the lifetime of your heating and air conditioning units, save money, and avoid future problems, having them serviced regularly...
Water filters not only provide you with clean, fresh, safe drinking water straight from your sink but help to save the environment as well. Adding water filtration may be a larger upfront installation cost,...
A water pressure regulator not only helps to reduce the water pressure coming from the main water line, but also helps it to flow at a safe level as well. To make sure that...
How to Unclog the Garbage Disposal There are times when you may forget to not put bones, coffee grounds, eggshells, bread, fibrous or stringy fruits and veggies, grease, and oil down the garbage disposal. ...
Very Satisfied. Michael is always on time, Michael has been doing all of the plumbing related to our remodel. He always does exactly what he commits to do within the estimated cost. He keeps me informed if any wrinkles pop up, and his plumbing skills are excellent as we have had no problems or issues arise after he has left. Very professional! Highly recommend.
August 2019
We had other companies come out for estimates on what was considered a major job with jackhammers and replacing cast iron pipes under the house… having been stung before, we wanted another estimate just to make sure we were making an informed decision. The prior estimates went up to $21,000!
Bill Howe sent us an amazing veteran plumber, Dashawn and his co- worker Anthony. Dashawn did a thorough check of the problem, explained what he thought was needed … he did a final check and we were able to see for ourselves what was the problem. He recommended a complete cleaning to de-scale the pipes, and not the major work we dreaded. Thanks to his solid analysis and patience in making sure we saw what he saw, the issue was resolved completely, and saved us thousands of unnecessary work. None of the other estimates included the cleaning; only the major repairs. Thanks to Dashawn's knowledge, professionalism and honesty, we're Bill Howe customers.
August 2019
This was our second excellent experience with Bill Howe. We had a blocked kitchen sink drain 8/5. I was able to get a same day appointment which is AMAZING. Person on phone was very courteous and set me up right away. Tony arrived at the house (called me 30 minutes out so I was able to be ready). He diagnosed the issue (blocked drain, not the garbage compactor) and was able to snake it and ran the water for 10 minutes to be sure the drains were clear. He cleaned up everything and you would not have known he was even there. Thank-you Tony and Bill Howe, you always know how.
August 2019
Cory was our technician on this job. We had a time sensitive repair issue and Bill Howe understood and assisted with getting the job done quickly.
Cory was on time to start the job and finished it quickly. Upon his leaving our home he's was courteous, very knowledgeable, explaining what the problem was and how he fixed it. Very good communication and this is the way it's has always been when we use Bill Howe plumbing.
Great company to work with.
Thanks Cory.
August 2019
Heating & Cooling
I had a shower that would not drain. The employee arrived on time and fixed the problem quickly and effectively at a very fair price. I was very happy with everything. Thank you and I highly recommend your plumbing service.
January 2019
I am extremely happy with the service we received from Bill Howe Plumbing. We were given an incorrect diagnosis of our main sewer line from another vendor, but Bill Howe's servicemen came out for a second opinion, correctly diagnosed our problem and resolved our back up issues. We will continue to use Bill Howe for all our future plumbing needs!
Thank you Bill Howe Plumbing.
July 2019


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how would you do it like a video and people watch and comment?
#226 2017-10-08 13:46:21
#227 2017-10-08 18:25:18
sure. I'd watch/participate, depending on time.
I do have a few questions:
1. How can I land on planets covered by shielding? I want to explore them.
2. How can I raise skills? I know mining/exploring are the obvious ones.
3. how can I find, say, something to give me more FTL range? I'd requisission it but don't have the creds or know what to search for.
4. How on earth do people find blueprints?
#228 2017-10-08 18:26:20
5. WHy are all the mission categories empty all the time?
and finally 6. What do I have to do to get notified when my light drive is fully charged? I feel like I'd turn that on if possible. It took me 3 days to jump 200 ly because I simply forgot about it for four hours....i'd not mind a way to notify me of light charge full
#229 2017-10-09 14:32:58
It sounds like a Q&A stream might me in order then, it will most likely be in the evening for the GMT timezone. To answer your questions Merin.
1: You cannot land on planets with shields, you'll either need to use bombardment weapons or mercenaries (unlock at level 20) to destroy the shield.
2: You raise skills by your character either reading or by performing the tasks. So if you want to improve your fighter skill you would need to engage in combat more often, for craftsmanship you would need to craft weapons or equipment etc...
3: You can either craft an overcharger or what for a merchant to sell one.
4: Blueprints can be bought at merchants.
5: The missions are shared between all players and are a commodity in themselves, they are also limited by factors such as your hold size, the distance and skills - depending on the mission.
6: This has been requested by a few different players, so is something we will look into.
#230 2017-10-09 15:39:51
Enjoying the game, so far.. An idea just poped into my head and I'd like to share it..
Do you guys think it would be good to add npc crews to ships? These would work as a sort of resource like rations.. They would be used to opperate larger ships and to explore other planets (as in, you send the crew down and receive say hourly reports of whatever they find).
How does that sound? Alsso, you'd need to keep your crew happy by feeding and entertaining them.
#231 2017-10-10 01:37:17
hey i just seen that there's a new facebook messenger IOS update. has anyone got it and have the remaining accessibility issues been fixed? I've heard that they certainly are better than they were am just wondering about the rest of them?
#232 2017-10-10 10:01:51
By what I've understood your idea, it is the same as mercenaries, and it is unlockable at level 20. However, there is certainly no reference to mercenaries that would explore individual planets on landing. However, since I am not na admin of the game, I cannot guarantee that there is no exploring mercenary. Anyway, here is the link to the information:
https://textspaced.zendesk.com/hc/en-us … 5000963174
I would certainly like to join the Q&A, after being informed about its date and time. Thanks in advance!
#233 2017-10-12 19:43:03
We've scheduled our live Q&A this Saturday at 8PM BST. Hopefully we'll be able to answer all of your questions.
#234 2017-10-12 22:43:34
for those that may not be able to attend could someone summerise the q&a afterwards and post it here?
#235 2017-10-13 23:17:27
Hi Darren,
Facebook stores live videos as normal videos after they are broadcast, so you should be able to listen back if you like.
#236 2017-10-14 06:47:39
hi guys...
I think I am stuck on a wierd bug,.
so I went to check my status on the computer menu, and hit cancel button after that, but the game do not return me to the main ship menu. so, now, everytime I hit cancel, or type something, those stats are keep comeing, makeing it kindda spammy, haha
#237 2017-10-14 07:17:31
I am not sure if this will work for you, but when I was experiencing a bug that dealt with the game's main menu, typing helm_enter would help, as suggested to me when I submitted a bug report directly in the game's page:
#238 2017-10-14 07:44:26
ah thanks!
#239 2017-10-15 07:04:13
#240 2017-10-15 08:32:57
try typing helm_enter?
I was on that loop too
#241 2017-10-15 08:36:42
Hi all,
Hopefully you should be all broken out of your loops now. Ever since the Facebook outage we have issues with loops and repeating messages that appear to occur at random to random people. We are working with Facebook to try and solve the issue.
#242 2017-10-20 13:27:34
Hello, I think I found a bug.
I was able to build an hideout.. I was transfering some cargo, when sudenly, my game just got back to the main menu.. I wasn't able to dock with my hideout because it said a ship was already there. Not just that but I can't access any other menues. So I can't mine, can't access the settings, among other things. Could you please help me?
#243 2017-10-20 14:25:39
Did you try using the "helm_enter" option as an alternative to access the menus? As a user's point of view, this is the help I can give. As for reporting bugs, how about doing it in the official channel for it, since they can look for reports more quickly there?
#244 2017-10-24 17:31:02
Hi caio,
You didn't respond to our support ticket - can we assume the issue is now resolved?
#245 2017-12-13 21:29:19
Hi guys,
We have had TextSpaced in open beta now since October 1st and are happy that we have reached 600 active players. I thought it would be good to detail how all the suggestions in this thread have helped shape the game this far.
Is there a way to see where star systems are in relation to each other
A text map was added to the helm.
Once I buy this scout ship though, what else is there to do? Mining is getting kind of dull...
Lot's more mission types have been added, more ships to strive for, more story and master skills - more everything really.
a way to designate a sector to easily return to it...
Lots of smart macros were added to the game including one for waypoints, type "waypoint 1" to go to your first waypoint for example.
Yeah the random race assignment kind of made me sad because I was looking forwards to playing a droid
There is a special story to meet the "life broker" who will provide services you may find interesting.
but there didn't seem to be any information about the book apart from the title...
Books now contain lore, you can review the lore from the "books read" section. Additionally some books contain special notes and ancient books can be purchased at the Mechanical Fleet or found.
I'm bout 2000 rl hours from a shipright...
Travel was changed so its now instant if you have enough charge.
Basically do I need a bigger ship or can I build a hideout with a scout?
The material requirements were lowed for a hideout and additional structures added to help in starting out building your faction.
Is it actively being worked on in terms of content? If it is, are there change logs, or some sort of way to be alerted of changes to the game?
All the change logs are here: https://textspaced.zendesk.com/hc/en-us … ouncements and the help centre is always having new articles added. Features have been planned for the next 5 years and TextSpaced has it's first intern to help out.
I'd vote terraforming, just for the awesome lore implications that has.
Terraforming has been made an explorer master skill.
does textspaced have any of my favorite weapon -- the grey goo
An exotic weapon was created called MEHEM.
what about colonising missions and you can have taxation on colonies within your faction? then once you have enough colonists you can create jobs for them IE ship building or mining for the faction.
The master skill of colonist was added to the game, once you choose colonist as your master skill you can create colonies and try to help them thrive. If you manage to start a settlement, get it to colony status then ultimately civilisation status, you'll see NPC ships appearing under your own faction name.
well why not have npc fighter wings and npc capital ships?
Replication bays are being added to the game so that you can covert a bay on your ship to a replication bay and have your own fleet of ships (biggest ship has 8 bays)
Would it give you all too much work if the game accepted typed input and not only menu input for the options? That way, instead of needing to go through "Helm." -> "Dock", you could only type in "Dock"
Lots of smart macros have been added including the "dock" smart macro.
1 of the things you could do was to post advertisements for things wanted or for products etc that you could sell.
There are a couple of player created mission types now, and a mission from the Descendants that asks for players to craft a specific weapon.
...t resulted in my character obtaining a strange glowing cube, but I can't see it anywhere. Would it be possible to add some sort of personal inventory
A personal inventory was added and new suits and weapons can be purchased from a faction owned fortress.
as far as I'm aware, you can't sell ships
Ships can now be sold at the Shipwright. Spare ports (up to 9) can also be stored at a Shipwright.
You can't buy resources, only sell them
You can now buy commodities. If you become a master miner you can also buy and sell ingredients, which are further divisions of a commodity. The ingredients work on a separate exchange which is far more localised and prone to change - this empowers you to find your own trade routes, buy low and sell high as well as benefit more from mining.
Are there message boards, public advertisement things, etc?
Being able to use in-game credits to have your faction show in a 'promoted' section is coming very soon.
these minefields a lot less of a spamming/trolling hazard
Minefields have be re-tooled so that they deny capture of structure. Once ship on structure/structure on ship combat is present, they will also deal heavy damage to ships attacking structures.
Been kind of sitting here for about a day, waiting for an engineer ship to load...this sucks.
All ships can now be purchased at any time, some requiring certain skills to be met. Also, if you spend enough time in a ship you can unlock an enhanced version.
or, failing that, better scanners so I can see farther out to make the tedium more barable?
A map was added and a new type of ship called the 'explorer' class of ship, it has a passive ability of scan pulse that discovers three sectors either side of your current location.
These are just the tip of the iceberg of changes that have happened, but again thank you all for your feedback; it has been very valuable.
#246 2017-12-14 13:22:57
Awesome! I've always been a fan of this game.. I couldn't continue to play there due to rl getting in the way, but now I will sertainly get back to playing!
#247 2018-02-14 09:42:48
Hi all.
We have some bad news I'm afraid. We have received communication from Facebook about Messenger that states the following:
"Our team is currently not supporting accessibility for Messenger Platform. They have put supporting accessibility on the roadmap for the long-term but in the short term there won't be a fix for this."
This came to a surprise to us, as originally they had said they would be focusing on accessibility. This means that the mobile version of Facebook Messenger wouldn't be guaranteed to be accessible for some time, so the web version is the more viable solution. We wish this was better news and will continue to try and push them on this issue. We will also explore other platforms for the future.
On a more brighter note, we thank you all here who continue to play and pushing us over the 700 active player mark. Its been a great journey so far.
#248 2018-02-14 14:47:32
so will you develop a web based version of this game? shame really and in a way am glad I didn't get too invested in it as I won't be able to miss playing it if accessibility is going to be cut.
#249 2018-02-16 09:12:03
Hi Darren,
This only affects the app version of Messenger, and we still have hope Facebook will fix these issues. The web version should still work fine and in fact the majority of our blind players (around 500) use this version.
#250 2018-02-16 15:27:28
ah so the web version is fine still? is there a way of overriding the app version of messenger does anyone know so I can still play it on the phone? every time I try using it, the app version takes over.


text:
Children's and Household Tales (German: Kinder- und Hausmärchen) is a collection of German origin fairy tales first published in 1812 by Jacob and Wilhelm Grimm, the Brothers Grimm. The collection is commonly known today as Grimms' Fairy Tales (German: Grimms Märchen).
Language: English
Published in: 1812
Translator: Edgar Taylor and Marian Edwardes
Word count: 101,584 words (≈ about 7 hours)
Source: http://en.wikisource.org
Copyright: Public Domain
The Harvard Classics, originally known as Dr. Eliot's Five Foot Shelf, is a 51-volume anthology of classic works from world literature, compiled and...
The first stories told at the dawn of history, these stories have been retold throughout the generations and continue to inspire us today.


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Lincoln Electric Sportsmen of the Year
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Congratulations to RMR's 2013 Lincoln Electric Sportsmen of the Year! Visit http://bit.ly/
LincolnElectric to learn more about Lincoln Electric's products and services.
Lovendahl-Rider Racing | Bill Larson | Austin Schoenfeld | Bill Matern | The Morgan Family
Lovendahl-Rider Racing: "The recipients of this award are big supporters of the Raceway, they were always attending other events, not just the oval. The family has been racing for years and always have a smile on their faces whenever they are at RMR. They are very deserving of this award and I appreciate their racing at our facility." Dan Breach
Bill Larson: "Bill is willing to help anyone at anytime and is very personable, respected and liked by his racing competitors. That is why I'm sure he was voted the Lincoln Electric Sportsman of the Year in the Street Legal Series." Ron Craft
Bill Matern: "Bill is a gentleman in all the ways, well liked and respected by everyone that knows him. He is a person that wants to win or lose a race, and has waited on his competitors many times in the past years. For that reason is why he was voted Lincoln Electric Sportsman of the Year and Bracket Racer of the Year in the Summit ET Series." Ron Craft
The Morgan Family: "Out here on the Motocross track we have quite a few deserving people for the "Lincoln Electric Sportsman of the Year Award". The Morgan Family stuck out because it wasn't just an individual but the entire family was there to help fellow riders and families. From the youngest all the way to the parents, they are awesome!" Curt Stewart


text:
The Engagements was a book that I picked up, looking for a quick chick lit escape this summer. What happened instead, was a book that has stuck with me since I put it down and surprised me with its depth and beauty. It was the kind of book that I couldn't wait for someone else to read and the one that I drove my husband crazy sharing about in the evenings because of the smartly woven plot twists.
Each chapter alternates stories with each story deeply rooted in the engagement ring from the inception of the idea of selling the idea of an engagement ring to women to what this ring signifies to each character in the book.
The stories unfold beginning with Frances Gerety, an advertising copyright who works on the De Beers account and who is the creator of the, "A Diamond is Forever," campaign that De Beers is still known for today. She is the one who changes what the diamond ring means to women and, yet, receives very little recognition for her efforts in her career. Fans of Mad Men, in particular, will fall in love with Frances and her struggles to get to the top of her ad agency, much like the beloved Peggy Olson in our favorite television series.
All of the other chapters weave in stories from different decades of what the diamond ring means to the other characters in the book. Evelyn has been married to her husband for 40 years, ever since her husband slipped off her first wedding ring and put his own in its place. Delphine has seen both the beauty and the awfulness in relationships and finds herself in a love triangle she never expected. James is a paramedic whose constant struggle to prove to his family and wife that he is good enough when he feels in his heart he is not, and the story of Kate who has been with her partner for a decade, but refuses to be married because of her feminist beliefs.
The novel spans nearly a hundred years with these stories and, perhaps, the most surprising thing for me was how utterly enchanted I was with every single story and character. I am guilty of favoring certain characters more than others, yet with this book, every story and every chapter was so great that I enjoyed all of them equally.
Don't let the slow start on this book discourage you- I promise that it will be worth it to see this one through to the end! This is my favorite book by J. Courtney Sullivan yet and I have a feeling it will be for you too!
MomAdvice Rating- 4 Stars out of 5 Stars
If you are looking for a little reading inspiration, be sure to check our MomAdvice fan page for a weekly check-in on what everyone is reading each week on our Facebook Fan Page. I hope you will swing by on Fridays and share about the books you are working on or request recommendations with one another!
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