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Environmental controls on the distribution of

bacterial tetraether membrane lipids:

Constraints on the MBT-CBT paleothermometer

Francien Peterse

ISBN 978-90-8570-839-1

On the cover: Mangshan Loess Plateau (courtesy of Maarten Prins).

Environmental controls on the distribution of

bacterial tetraether membrane lipids:

Constraints on the MBT-CBT paleothermometer

Omgevingsinvloeden op de verdeling van

bacteriele tetraether membraanlipiden en toepassing van de MBT-CBT paleothermometer (met een samenvatting in het Nederlands)

Proefschrift

ter verkrijging van de graad van doctor aan de Universiteit Utrecht ingevolge het besluit van het college voor promoties in het openbaar te verdedigen op donderdag 15 september 2011 des middags te 4.15 uur door

Francien Peterse

geboren op 6 juni 1980 te Amsterdam

Prof. dr. ir. S. Schouten

and the Royal Netherlands Institute for Sea Research (NIOZ). ha die pa, voor jou

Summary

Samenvatting

Part I

soils: Implications for the MBT-CBT temperature proxy.

Chapter 4:

Assessment of soil n-alkane

brane lipid distributions as tools for paleoelevation reconstruction.

Part II

Chapter 5:

Absence of seasonal patterns in MBT-CBT indices in mid-latitude soils.

Chapter 6:

Constraints on the application of the MBT-CBT paleothermometer in peat and soils.

Chapter 8:

Revised calibration of the MBT-CBT paleotemperature proxy based on branched tetraether membrane lipids in surface soils. the last deglaciation.

References

About the author

Table of contents

9 12 15 25
33
39
49
69
81
95
119
135
142
144
9

Summary

tion of Branched Tetraethers (MBT) index and the Cyclisation of Branched Tetraethers (CBT) index. The

investigated and possible constraints on its applicability need to be ex amined.

relation is similar to that based on the global soil calibration set, the intercept is different. This difference

can be explained by the use of in situ soil temperature in the relation for the geothermally heated soils

dance of Acidobacteria

suggested as a proxy for paleoelevation, in addition to soil n-alkane D values. To assess their suitability

D of n-alkanes

both proxies can be subject to relatively large uncertainties, but that a combination of both proxies likely

results in a more reliable paleoelevation reconstruction. 10

potential seasonal bias in MBT-CBT-derived air temperature estimates, for example caused by a season of

over an annual cycle. MBT-CBT-reconstructed temperatures remained constant throughout the year and

organic matter may be predominantly produced in situ. The MBT-CBT proxy should therefore only be used

for marine coastal sediments at sites receiving a substantial input of soil organic matter relative to the

amount of marine organic matter, i.e. close to a river mouth.

primarily produced in this part of the peat. This is supported by the increase of absolute concentrations of

Acidobacteria

in these soils, albeit in different distributions.

distributed surface soils to recalibrate the proxy. As only 26% of these surface soils contained all nine

2 2

Summary

11

maining scatter in the calibration, but reconstructed MATs for soils from arid regions tend to substantially

independent proxy data. Application of the MBT-CBT proxy on a loess-paleosol sequence in China resulted in a paleotempera- 12

Samenvatting

bevat. Uit een empirisch onderzoek van meer dan 130 bodems is gebleken dat de samenstelling van

reconstructie van temperatuurveranderingen in het stroomgebied van de rivier in het verleden. Voordat de

Acidobacteriën, gesuggereerd als mogelijke

hetgeen deze suggestie ondersteunt.

Samenvatting

13 D- D

ing richting een bepaald seizoen laten zien, mogelijk veroorzaakt door een periode van optimale groei-

langer is dan een jaar. om na te gaan of de MBT-CBT proxy ook toepasbaar is in Arctische gebieden, gekenmerkt door een lage door het koude klimaat en de geringe mate van bodemvorming relatief laag is, komt de met de MBT-CBT marien organisch materiaal, zoals bijvoorbeeld bij riviermondingen het g eval is. 14

Acido-

bacteriën 2 2

stelde achterliggende fysiologische mechanismen. Lokale omgevingsfactoren of de invloed van seizoenen

kunnen de overgebleven variatie in de kalibratie niet verklaren, maar berekende MATs voor bodems uit peratuurreconstructies laat zien dat de trends niet veranderen, maar dat de absolute temperaturen en overeenstemming zijn met onafhankelijke proxy data.

randeringen in luchttemperatuur in fase liepen met de zomerinsolatie op het noordelijk halfrond en toont

hetgeen overeenkomt met de uitkomst van andere studies die de continentale luchttemperatuur recon- gedreven zijn door verschillende factoren.

Samenvatting

Chapter 1

General outline and introduction

16

1.1 Climate variability

changes in the mixture of atmospheric greenhouse gases. Especially variations in the concentration of CO

2

have appeared to be a strong driver for climate change. Since the industrial revolution, human activity

has increased the level of CO 2

earth"s surface (IPCC, 2007). For the prediction of the exact consequences of continuing rise of the CO

2 level

for our future climate, global climate models are used. These models are generally validated by simulating

2 concentrations in the past, like the Early Eocene (56-48 2

for the output data of these models only go back a limited amount of time. The global temperature record,

for example, only covers just over the past 100 years (Peterson and Vose, 1997). Extension of instrumental temperature records can be achieved by the application of so-called cli-

linked to an environmental parameter. Application of these proxies on a sedimentary record provides valu-

able information on past climatic variations, e.g. temperature, crucial for the testing of climate models.

responsible for the global redistribution of heat, and thereby form an important control on the earth"s

climate (Wang et al., 2004).

1.2 Reconstruction of past temperatures

There are various methods to reconstruct past sea surface or air temperatures. SSTs, for example,

are recorded in the oxygen isotopic composition of calcite or opal containing shells that are preserved in

marine sediments. The ratio of 18

O and the lighter

16 O that is incorporated in the shells depends, amongst (). The elemental composition of the shells also provides information on past ). A temperature

signal can furthermore be obtained from fossil organic molecules present in sediments. For example, the

37:2
and C 37:3
long-chain unsaturated ketones depends on tem- perature (Brassell et al. 1986 k" 37
index (Unsaturation ratio of long chain (C 37
these alkenones are present in a sedimentary record (Fig. 1.1). SSTs are also recorded by the distribution of isoprenoid tetraether membrane lipids of marine Thaumarchaeota. on the temperature of their environment (). The number of cyclopen- TEX 86
in the relative abundance of the membrane lipids into SST (Fig. 1.2).

Chapter 1

17

Fig. 1.1

k" 37
k" 37
and annual mean

Müller and Fischer, 2004).

compared to those in the marine realm. Nevertheless, there are several proxies that are regularly applied

is, for example, the inventory of pollen assemblages (e.g., Colinvaux et al., 1996

preserved in lake sediments and peat bogs, and quantitative temperature estimates can be derived from

calibrate glacial assemblages (). This method furthermore assumes that plant-climate in- interactions are sensitive to changes in e.g. CO 2 concentration (Street-Perrott et al., 1997). A second plant construct paleotemperatures. Baily and Sinnot (1915

into a quantitative method based on linear regressions of extant untoothed species and MAT, consequently

enabling the reconstruction of paleotemperature by analyzing the margins of fossil leaves ( regression models do not take adaptation to changes in the environment into account (

al., 2010). Also the effects of preferential preservation of (un)toothed leaves and the extinction of species

are not considered (). By making use of the temperature sensitivity of the oxygen isotopic composition of precipitation, ter location the longest record has been retrieved, extending 800,000 years ( ). Obviously, ice core records are limited to ice covered sites, although these are most

stalagmites can be used to reconstruct past temperatures. Also the isotopic value of the carbon and trace

18 perature signal (). A last example of a method that used oxygen isotopic values to infer

paleotemperatures is based on animal bone and tooth enamel. Bones and teeth contain biogenic apatite,

e.g. Longinelli, 1984). Thus, past on fossil bones and teeth ()

Fig. 1.2

86
and the reation of annual mean 86
values (Kim et al., 2008). [GDGT-2]+[GDGT-3]+[Cren'] [GDGT-1]+[GDGT-2]+[GDGT-3]+[Cren'] TEX= 86
Lakes provide generally suitable settings to obtain continuous, high resolution paleotemperature

records. The climatic information can be retrieved from the fossils of aquatic organisms that are preserved

tions in their distribution in lake sediments (), although Velle et al., 2005). After the detection of Thaumarchaeotal isoprenoid tings (), also applications of the TEX 86
on lake sediments have resulted in quantitative past continental temperature reconstructions ( ), and present studies indicate that TEX 86

Blaga et al., 2011).

Chapter 1

19 In summary, many of the available continental temperature proxies are besides temperature also

be globally applicable, provide quantitative temperature estimates, have temperature as main environ-

2 concentration.

1.3 The MBT-CBT continental paleothermometer

Fig. 1.3) membrane lipids (Weijers et al., 2007c-

magnetic resonance spectroscopy (

Weijers et al., 2006a

mixed archaeal (membrane spanning tetraethers, ether lipids) and bacterial (branched alkyl chains)

Weijers et al., 2006a). The bacteria that

the phylum of Acidobacteria may be their source (Weijers et al., 2009).

Fig. 1.3.

Fig. 1.4

Methylation of Branched Tetraethers (MBT) and the Cyclisation of Branched Tetraethers (CBT), enables The Roman numerals in the equations refer to the abundance of the molecules i n Fig. 1.3. 20

Fig. 1.5a, b):

2 2

by the CBT index then results in a function to estimate MAT, solely based on the distribution of branched

Fig. 1.5c):

2

coastal sediments may yield an integrated temperature record for a river catchment area. For example,

(i.e. Eq. 1.5) in a sediment core retrieved from the Congo River fan (Weijers et al., 2007a). The timing and

Weijers

et al., 2007b). Thus, the MBT-CBT proxy seems to be a promising tool to obtain paleoclimatic information that can

contribute to a better understanding of the timing and the driving forces of climatic changes in the past.

of the MBT-CBT proxy.

Fig. 1.4.

tures (Weijers et al, 2007c). Roman numerals refer to the structures in Fig. 1.3.

Chapter 1

21

Fig. 1.5.

Weijers et al., 2007c).

1.4 Scope and framework of this thesis

Weijers et al., 2007c

be determined. In the second part, the proxy is further constrained and tested in a variety of settings in

order to test the applicability of the proxy.

Chapter 2 describes the

cf. Fig. 1.4 that this is an important factor controlling this proxy.

Chapter 3. Analysis of core lipids (CLs) and derivatives of their intact and presumably living precursors,

22

Acidobacteria

In Chapter 4 -

D analyzed in the same

D and the MBT-CBT proxy may result in more

reliable paleoelevation reconstructions. The second part of this thesis decribes the constraints on the application of the MBT-CBT proxy as paleothermometer. In Chapter 5 is thus in the order of one year or more. applied on modern Svalbard soils and coastal sediments (Chapter 6 of marine OM, i.e. close to river mouths.

Chapter 7

Acidobac-

teria

In Chapter 8

improvement only of the calibration accuracy. Furthermore, the statistically derived indices no longer

Chapter 1

23

Chapter 9

yielded a continuous air temperature record for East Asia that covers the past 34,000 years, based on

18

O records indicates that the onset of

settings. this. This supports the general suitability of the MBT and CBT indices as paleothermometer and proxy

relatively long, and that, as a consequence, absolute temperatures generated by the proxy are not biased

recovered from areas that receive a substantial input of terrestrial OM relative to the amount of marine

be applied in a variety of sites and time periods and can yield valuable unique paleoclimatic informa-

tion.

Distribution of branched tetraether lipid in geo-

thermally heated soils: Implications for the MBT-

CBT temperature proxy

Organic Geochemistry 40, 201-205.

Chapter 2

Abstract

in soils, the Cyclisation of Branched Tetraethers (CBT) index and the Methylation of Branched Tetraethers

2

geothermally heated soils is similar to that of a global soil calibration set, although the intercept for the

26

2.1 Introduction

Fig. 2.1

al.(2000 cyclopentane moieties (Fig. 2.1Weijers et al., 2006a).

based on the fact that branched dialkyl glycerol diethers have been found in thermophilic bacteria (Lang-

) and that non-isoprenoid lipids have not been

2000) suggested bacteria to be the most likely producers

2006a

Fig. 2.1.

In an empirical study, Weijers et al. (2007c) analyzed about 130 soils from 90 different locations

They found that the number of methyl groups relates to the annual mean air temperature (MAT) and soil

The numbers refer to the structures in Fig. 2.1Table 2.1

core recovered close to the river mouth of the Congo Basin resulted in a temperature record for tropical

Chapter 2

27
Africa representing the last 25,000 yr (Weijers et al., 2007a-

Weijers et al., 2007b2008a)

uncertainty exists over their calibration and the impact of other environmental factors. For example, the

MAT for each sample in the soil data set of Weijers et al. (2007c MAT values slightly overestimated the in situ measured annual MAT (), suggest- parameters at each sampling site are similar.

2.2. Material and methods

2.2.1. Soil location and sampling

Table 2.2). From the edge of Leonard"s spring, soil samples th digital thermometer. The 2 2

Table 2.1. 2007c)

and Californian hot springs. 28

2.2.2. GDGT analysis

6 2 SO 4 2 g of an internal C 46
). The extracts 2 O 3 2 , the m (Ø 4 mm) prior to analysis using

2.2.3. HPLC/APCI-MS

20002007a

l. Single ion monitoring (SIM) of the M ions of the internal C 46

Weijers et al., 2007

ferent response factors (). a

Table 2.2.

a

Roman numbers refer to structures in Fig. 2.1.

b

Chapter 2

29

2.3. Results and discussion

2.3.1. Branched GDGT distribution in soil transects

both springs (Table 2.2 at 250 cm (Table 2.2).

Fig. 2.2

sites, the clayey soil had a greyish blue colour and a sulfurous smell, indicating anaerobic conditions. The

that anaerobic soil bacteria are producers of these membrane lipids (Weijers et al., 2006a). Fig. 2.3

III are more abundant and occur in about equal amounts. This change in distribution pattern for both hot

Weijers et al., 2007c).

Fig. 2.2.

30

2.3.2. CBT and MBT indices

2.1), i.e. for Leonard"s spring MBT values changed from 0.44 close to the spring to 0.26 at the farthest

2 Fig.

2.4a; Table 2.1

than that reported by Weijers et al. (2007c 2

Table 2.1). In

situ Fig. 2.4b; Table 2.1 as the global soil calibration set of Weijers et al. (2007c FP FP In contrast to the CBT, the MBT index related to in situ soil temperature and, although to a much Fig. 2.4c and d; Table 2.1). This result is similar to that of Weijers et al. (2007cquotesdbs_dbs19.pdfusesText_25