Soil respiration in drylands Essay

Chapter 1. Introduction

Soil respiration in drylands is ill understood and mostly understudied, despite the fact that deserts cover more than 30 % of the Earth ‘s land surface ( Thomas, 1997 ) . Given the current involvement in planetary C cycling, and in peculiar the grade to which dryland dirt C cycling may lend to it ( Lal 2004 ; Luo & A ; Zhou 2006 ) , there remains a dramatic dearth of cognition.

It is merely within the past decennary that the wide R & A ; ocirc ; lupus erythematosuss played by Biological Soil Crusts ( BSCs ) in dryland environments have begun to be earnestly appreciated ( Belnap & A ; Lange 2003 ) . Interest remains preponderantly focussed on their R & A ; ocirc ; le in keeping dirt stableness and their influence on hydrological procedures. More late, consciousness has increased that BSCs play a ‘boundary mediating ‘ R & A ; ocirc ; le, partly regulating the exchange between dryland dirts and the ambiance ( Belnap et al, 2003a, 2003c ; Viles, 2008 ) . Of existent importance is our understanding that BSCs can both breathe CO2 through respiration, but besides sequester C ( C ) through photosynthesis. The latter, in peculiar, is merely possible when crusts are sufficiently hydrated so as to be metabolically active.

This survey focuses on the boundary mediating R & A ; ocirc ; lupus erythematosus of BSCs, and partly arises from the renewed involvement in and recent versions of Noy-Meir ‘s ( 1973 ) ‘pulse-reserve ‘ theoretical account ( Schwinning & A ; Sala, 2004 ; Schwinning et Al, 2004 ; Ogle & A ; Reynolds, 2004 ; Humxan et Al, 2004a ; Austin et Al, 2004 ) . The paradigm displacement in fact-finding attacks to drylands, now understood as bing in a non-equilibrium province ( Vetter 2005 ) , underwrites the experimental principle of the survey.

Broadly, the survey investigates CO2 surface flux, and the environmental factors that may act upon flux activity. The survey involves sustained in situ sampling from the southern Kalahari Desert, near to the boundary line between Botswana and South Africa and dressed ores on placing distinguishable CO2 pulsations after fake precipitation events. The factors that might act upon the magnitude of pulsation ( precipitation sum, Photosynthetically Active Radiation, temperature ) are considered. Equally, the survey considers those factors that might bring on photosynthetic C consumption by BSCs. Subsoil respiration is besides sampled, and temporal and spacial variableness in flux activity is besides investigated.

Literature Review

Despite their frequently low vascular works screen, it would be misguided to presume that drylands are barren of important biological life ( Belnap & A ; Lange, 2003 ) . BSCs are now understood to be present in all waterless and semi-arid parts ( Thomas & A ; Dougill, 2007 ; Zaady et Al, 2000 ) , representing more than 70 % of life screen ( Belnap et al, 2003 ) , and their R & A ; ocirc ; lupus erythematosus in critical geomorphic and ecological procedures is progressively recognised ( Belnap et al, 2003a ; Viles, 2008 ) .

BSCs are “ an confidant association between dirt atoms and blue-green algaes, algae, microfungi, lichens, and nonvascular plants… which live within, or instantly on top of, the topmost millimeters of dirt ” ( Belnap et al 2003a, p.3 ) . Cyanobacteria based crusts in peculiar have the ability to bring forth exopolysaccharide secernments, which are good recognised to increase dirt cohesive strength and cut down erodibility ( Thomas & A ; Dougill, 2007 ) . Correspondingly, with the increased focal point on land debasement in drylands, important attempts have been made to research the relationship between BSCs and Aeolian and fluvial eroding, with noteworthy parts by Belnap & A ; Gilette ( 1998 ) , Argaman et Al ( 2006 ) , Bowker et Al ( 2008 ) and McKenna Neuman & A ; Maxwell ( 2002 ) .

Biological Soil Crusts as a mediating boundary

Less attending has been directed towards the ability of BSCs to modulate the boundary between the ambiance and dirt ; they are basically a illumination ecosystem between the two ( Bamforth, 2008 ) . Belnap et Al ( 2003c ) have produced some seminal work, though surveies remain focused towards the possible R & A ; ocirc ; lupus erythematosuss of BSCs in dryland hydrological rhythms. In peculiar, Belnap ( 2006 ) found the influence of BSCs on infiltration rate, surface run away and sediment production to be mostly a map of the phase of sequence of the crust. This determination has been reaffirmed by Aguilar et Al ( 2009 ) through their research in northern Mexico. Dougill & A ; Thomas ( 2004 ) suggest a three-stage categorization system for BSCs as based on their phase of sequence ( with Stage Three being the most developed ) . Indirectly, Harper & A ; Belnap ( 2001 ) and Berkley et Al ( 2005 ) discourse the influence of BSCs on vascular works mineral consumption, happening a important correlativity between BSC presence and the bioessential mineral content of vascular workss. This is thought to impact overland hydrology, as BSCs are likely to back up the ‘islands of birthrate ‘ theoretical account ( Garcia-Moya & A ; McKell, 1970 ) . Consequently, of peculiar note is the work of Housman et Al ( 2006 ) who report that the C and nitrogen arrested development abilities of BSCs are besides differentiated as a map of successional phase. Housman et Al ‘s ( 2006 ) survey is declarative of a broader grasp of BSCs ‘ ability to gaining control, transform or debar atmospheric inputs ( Belnap et al, 2003c ; Viles 2008 ) . This should non be underestimated ; they have been called ‘small-scale mantles of birthrate ‘ ( Garcia-Pichel et al 2003b ) . Correspondingly, there has been a heightened involvement in the Carbon ( C ) and Nitrogen ( N ) arrested development potency of BSCs.

Biological Soil Crusts: Respiration and photosynthesis

Despite the floaty involvement in dryland dirt respiration, Cable & A ; Huxman ( 2004 ) emphasis that cognition of the R & A ; ocirc ; le BSCs play in ecosystem C cycling is still in its babyhood. Given that drylands occupy over 30 % of the land surface globally, dirt crust blue-green algae are likely to hold important populations ; Garcia-Pichel et Al ( 2003a ) estimation there to be 54 ten 1012gC cyanophyte biomass in dryland crusts globally. The realization that CO2 transmutation by BSCs contributes significantly to C budgets in drylands has provided farther drift for research ( Belnap et al 2003c ) ; Lange in peculiar has written prolifically ( 1994, 1997, 1998, 1999, 2003a ) . Evans and Lange ( 2003 ) argue that the more evolved, lichen and moss dominated BSCs, potentially have photosynthetic rates on a par with vascular workss ( 120-370kgC ha-1 yr-1 ) although Belnap et Al ( 2003c ) emphasis that cyanobacterial-dominated crusts have lower photosynthetic potency ( 4-23kgC ha-1 yr-1 ) .

Concerted attempts have been made to characterize the biochemistry that underlies BSCs ‘ respiratory and photosynthetic activity. Concentrating in peculiar on cyanophyte crusts, they are known to be poikilohydric, i.e. they are merely metabolically active given sufficient wet handiness ( Zaady et Al, 2000 ; Belnap et al 2003b ; Ustin et Al, 2008 ) . BSCs are thought by some to be the most effectual of any photosynthetic being ( Badger & A ; Price 2003 ) , their marked photosynthetic ability being mostly attributable to their individual cell CO2 concentrating mechanism. Further biochemical grounds of environment version is provided by Ehling-Schulz & A ; Scherer ( 1999 ) , who argue that blue-green algae use three different schemes to antagonize harm from overexposure to UV. In a seminal paper Garcia-Pichel & A ; Pringault ( 2001 ) present findings that cyanobacteria vertically migrate within the dirt profile in response to wetting and drying events ; this silent reponse to H2O is thought to be alone, and probably a get bying mechanism towards utmost dryland desiccation-hydration rhythms. Bamforth ( 2004 ) continues, happening that Protozoa are the dominant signifier of microfauna that contribute to metabolic working in hot comeuppances ( though subsequently research ( Bamforth 2008 ) found markedly different Protozoa in cold desert BSCs ) .

There has besides been a move to set up the dominant environmental variables commanding BSC respiration and photosynthesis. Temperature is understood to be a critical variable, with photosynthetic activity of a rehydrated cyanobacterial crust found to be significantly higher at 25 & A ; deg ; C and 35 & A ; deg ; C than 5 & A ; deg ; C and 15 & A ; deg ; C ( Zhao et al, 2008 ) . Given the important temperature scope in many comeuppances, BSCs besides show singular version to extreme temperature ranges. San Jos & A ; eacute ; & A ; Bravo ( 1991 ) study maximal net photosynthesis at 45 & A ; deg ; C in Venezuelan Savanna algal dirt crusts, whilst Lange et Al ( 1998 ) discuss three different species ‘ varied net photosynthesis and dark respiration in response to a broad temperature scope in the Colorado Plateau, Utah.

The relationship between wet, respiration and photosynthesis

Given blue-green algae ‘s poikilohydric nature, wet is besides thought to be a basically of import variable. Lange et Al ( 1994 ) study, nevertheless, that whilst certain blue-green algaes can set about net photosynthesis from high humidness H2O vapor entirely, green algae and green algal lichen-formed crusts are reliant on liquid H2O to go metabolically active. Variable responses to wet are hence to be expected within BSC taxa, as reiterated in a later survey which found C arrested development as a map of wet to be significantly differentiated between four different species of BSC in southern Utah ( Lange et al, 1998 ) . It is hence unsurprising that Belnap et Al ( 2004 ) conclude that any change in precipitation governments will probably change the physiological operation of BSCs.

Dryland dirt outflow

Work undertaken on dryland dirt respiration and C segregation is still instead limited ( Schlesinger 1999, 2000 ; Luo & A ; Zhou, 2006 ) . Raich & A ; Schlesinger ( 1992 ) see CO2 flux in dirt respiration at the planetary graduated table and its relationship to flora and clime ; Conant et Al ( 2004 ) more specifically analyze the legion environmental controls on dirt respiration in semi-arid dirts, where dirt wet and temperature are noted as dominant factors. An earlier survey besides found that dirt microbic respiration is mostly moisture limited, with increased dirt temperature by and large taking to greater dirt respiration ( Conant et al 2000 ) . In a pioneering survey, Kieft et Al ( 1987 ) straight link the wetting of a dry waterless dirt to a mensurable release of organic C from the dirt biomass, though it is improbable to be a simple cause and consequence relationship ; Liu et Al ( 2002 ) emphasis the common determination that the relationship between dirt CO2 outflow and dirt wet is extremely variable, declarative of complex mechanisms finding dryland dirt outflow.

At the planetary graduated table, Lal ( 2004 ) has been peculiarly vocal in his research into C segregation in dryland ecosystems, and notes that drylands have been suggested as the ‘missing sink ‘ in the planetary C budget ( Lal 2003 ) . However, there remains really existent uncertainness refering dryland C pools, good characterised by the recent lively exchange between Wohlfahrt et Al ( 2008 ) and Schlesinger et Al ( 2009 ) . Whilst Wohlfahrt et Al ( 2008 ) contend that their research demonstrates that an country of the Mojave was a net CO2 sink for 2005 and 2006, Schlesinger et Al ( 2009 ) strongly question the cogency of their findings. Stone ( 2008 ) summarises Xie et Al ‘s ( 2008 ) recent farther part to the argument, proposing that high dirt salt or alkalinity positively drive CO2 segregation in a cold arid desert.

‘Pulse-Reserve ‘ Paradigm

The existent nature of dryland precipitation is thought to be critically of import in regulating both undersoil outflow and BSC activity ; episodic H2O handiness irrefutably affects dryland component cycling ( Austin et al, 2004 ) . Indeed, BSCs are exposed to significantly different precipitation governments. Veste et Al ( 2008 ) see the respiratory activity of BSCs in Israel ‘s Northern Negev coastal desert, where regular dews contribute cumulatively important sums of H2O to the ecosystem. Lalley et Al ( 2006 ) note similar fog precipitation in the Namib desert ; this is potentially biologically important, estimated at four times the 19mm average one-year rainfall.

In recent old ages at that place has been renewed involvement in precipitation forms and their influence on dryland ecosystem processes ( Huxman et Al, 2004a, B ) . The influential ‘pulse-reserve ‘ paradigm draws on the fact that dryland precipitation “ is non merely discontinuous but besides stochastic ” ( Noy-Meir, 1973: 31 ) . It suggests that pulsations of BSC activity are tightly coupled to precipitation events. Noy-Meir ‘s ( 1973 ) hypothesis has received considerable attending, and its focal point on important temporal and spacial precipitation variableness is supported by the more recent non-equilibrium hypothesis ( Vetter, 2005 ) . In recent old ages it has been modified ; Schwinning et Al ( 2004 ) emphasis both the importance of biologically important pulsations ( pulling on Beatley ‘s ( 1974 ) work into response thresholds ) and hysterisis, the construct that ecosystems have ‘memory ‘ of past precipitation events that governs response to future events. To this terminal, Reynolds et Al ( 2004 ) propose that sequences of precipitation events, as opposed to single pulsations, are more relevant for understanding dryland C cycling and the function BSCs drama in interceding it. Schwinning & A ; Sala ( 2004 ) further modify Noy-Meir ‘s ( 1973 ) original paradigm, reasoning that pulse deepness and pulse continuance are critical in understanding BSC response to pulse events. Indeed, they argue that the length of metabolic activity should represent the pulsation, non the original precipitation event ; accordingly, pulse events are likely to hold different continuances for different species.

Empirical grounds for both BSC activity and undersoil respiration supports the pulse-reserve paradigm. Both Huxman et Al ( 2004a ) and Ogle & A ; Reynolds ( 2004 ) theoretical account ecosystem component responses to precipitation pulsations, being careful to emphasize that the influence of pulse events on dryland C kineticss is significantly governed by antecedent wet conditions. Fierer & A ; Schimel ( 2003 ) suggest a mechanism for rapid undersoil CO2 production following a fake pulsation event, though Huxman et Al ( 2004b ) find high spacial variableness in CO2 outflow in Arizona, USA, following fake pulsations. Similarly high, small-scale variableness is reported by Maestre & A ; Cortina ( 2003 ) in a Mediterranean semi-arid steppe. In a research lab based experiment, Satoh et Al ( 2002 ) further reaffirm cyanophyte version to dehydration, demoing rapid photosynthetic activity upon wetting, though Harel et Al ( 2004 ) conclude that there is really small research into the mechanisms that allow photosynthetic recovery after hydration.

Biological Soil Crusts and subsoil outflow in the Kalahari, Southern Africa

To day of the month, the pulse-reserve theoretical account has been small studied in Southern African drylands. Nicholson ( 1994, 2000 ) reiterates the current intense spatial and temporal precipitation variableness in Southern Africa, and Schwinning et Al ( 2004 ) note the likeliness of clime alteration in drylands coercing greater precipitation extremes with more fickle fluctuations. This is peculiarly true of the Kalahari, where Thomas et Al ( 2005 ) envisage future clime alteration further cut downing moisture handiness in the part. Despite Weltzin et Al ( 2003 ) reaffirming the criticalness of research to understand how ecosystems respond to alterations in precipitation, there are really few probes into undersoil and BSC responses to precipitation pulsations in the part. Of the few surveies identified, there is a clear division in spacial graduated table studied. Whilst Wang et Al ( 2007, 2009 ) comparison sites over a 600km Kalahari Transect, Thomas & A ; Hoon ( 2010 ) and Thomas et Al ( 2008 ) focal point in peculiar on a more waterless site in southern Botswana. In decision, cognition refering the spacial distribution of BSCs ( Dougill & A ; Thomas, 2004 ; Berkley et Al, 2005 ; Thomas & A ; Dougill, 2007 ) and their ability to intercede and react to precipitation pulse events in this critical country is highly limited, peculiarly at finer spacial graduated tables.


This survey aims to foster the work of Wang et Al ( 2007, 2009 ) , Thomas et Al ( 2008 ) and Thomas & A ; Hoon ( 2010 ) . There is at present an unsatisfactory famine of research into in situ dryland dirt CO2 fluxes, the huge bulk of surveies to day of the month being unreal research lab simulations. Those that are field based are on the whole little graduated table and temporally harsh, and consequence in unacceptable perturbation of Biological Soil Crust unity. Additionally, our apprehension of the temporal and spacial variableness that characterise surface and subsurface fluxes in the Kalahari is still mostly in its babyhood.

This survey hence investigates the effects of fake precipitation events on both BSC respiration and photosynthesis and undersoil respiration at two sites in the southern Kalahari, with three wide purposes:

  • To detect how Biological Soil Crust-mediated surface efflux/uptake varies internally within sites.
  • To detect how Biological Soil Crust-mediated surface efflux/uptake and subsurface respiration vary between two nearby ( 350 meters ) sites on different substrates.
  • To set up what environmental factors affect Biological Soil Crust-mediated surface efflux/uptake and subsurface respiration.

Chapter 2. Location of Research: The Southern Kalahari Desert

The Kalahari ‘s clime is capable to a distinguishable seasonality, crudely distinguished between moisture ( October to April/May ) and dry. Liing within the southern hemisphere semitropical high force per unit area belt, winter precipitation in the northern Kalahari is peculiarly influenced by fluctuations in the places of both the Inter Tropical Convergence Zone and the Zaire Air Boundary ( Tyson 1986 ) . Temperatures vary both seasonally and spatially ( with often big diurnal temperature governments during the dry season ) , and systematically high temperatures coupled with low humidness consequence in high possible evapotranspiration ; Upington, South Africa has 3805mm one-year possible evapotranspiration, for illustration ( Thomas & A ; Shaw 1991 ) .

The intense temporal and spacial precipitation variableness that characterises the Kalahari is of acute importance to the survey. Though efforts to climatically delimit countries of the Kalahari are fraught with trouble due to inter-annual and inter-decadal variableness, general features can be noted. In peculiar, there is a south-west north-east gradient with one-year precipitation increasing in easterly and northern waies, runing from waterless to semi-arid/subhumid ( Thomas & A ; Shaw, 1991 ; Dougill & A ; Thomas 2004 ; Wang et Al 2007 ) . Precipitation variableness is besides found to increase to the South. This consequences in a important scope in precipitation from the waterless South to the more humid north-east, as shown in both Fig. 2.1 and Table. 2.1.

Though Pike ( 1971 ) concludes that half of Kalahari storms generate less than 10mm of rainfall, the nature of precipitation remains ill documented. Whilst little graduated table spacial variableness can frequently be seen as a map of convective activity, longer term temporal variableness is besides noteworthy. Tyson ( 1986 ) notes a statistically important 18 twelvemonth rhythm of rainfall fluctuations over southern Africa ( Thomas & A ; Shaw 1991 ; Tyson 1986 ) .

The Kalahari ‘s dirts have received light scientific attending, mostly due to their low agricultural potency ( Thomas & A ; Shaw 1991 ) , but most surveies report a close relationship between geology and dirts. Kalahari sand dirts in peculiar consist of over 95 % all right sand-sized, Aeolian deposited deposit ( Thomas & A ; Shaw 1991 ; Wang et Al 2007 ) . Predominantly deep and structureless, soil organic C and entire N content have their highest concentrations ( C max = 1.5 % , N max = 0.1 % ) in the top 10-20 centimeter of the dirt profile ( Wang et al 2007 ) . Surveies of Biological Soil Crust ( BSC ) prevalence have, until really late, been wholly neglected in the Kalahari. However, among recent surveies Dougill & A ; Thomas ( 2004 ) study cyanophyte BSC screen runing from 19 % to 40 % , at four locations within the Kalahari, and Berkley et Al ( 2005 ) assert the belief that BSC distribution in the Kalahari is likely to be related to Acacia mellifera ( which has a peculiarly high prevalence in the Kalahari ) and Grewia flava populations, happening crust denseness positively correlated to Acacia mellifera canopies.

The research for this survey was undertaken on two sites at Berry Bush Farm, about 7km north E of Tsabong, in south west Botswana ( 25 & A ; deg ; 56’51S 22 & A ; deg ; 25’40E ) , as shown in Fig 2.2. The two survey secret plans are within an enclosed country of farming area that has a history of graze ( most late a twelvemonth before the survey ) , and flora is preponderantly a mix of grass, woody bushs and trees ( Thomas & A ; Hoon 2010 ) . One secret plan was sited on Kalahari sand ( afterlife referred to as KS ) , with & gt ; 97 % grain size of all right sand, and a pH 5.9 +/- 0.4 ( Thomas & A ; Dougill 2004 ) . The other ( hereafter referred to as Calcrete ) was sited on a surface desiccated calcrete pan, with important Aeolian sedimentations of Kalahari sand. The huge bulk of crusts in the country are cyanophyte, typically 3-4mm midst and scope in morphological development from Type 1 to Type 3, following Thomas & A ; Dougill ‘s ( 2007 ) categorization system. Coverage varies widely ; from 19-40 % in pastoral countries to over 95 % in undisturbed wildlife direction zones. KS had about 55 % coverage, and Calcrete 75 % .

Berry Bush Farm has a precipitation record dating from September 1996. Fig 2.4 shows a histogram of precipitation events from September 1996 to June 2009.

Berry Bush ‘s precipitation informations have been part-processed by the writer to bring forth Table 2.2 which besides shows a comparing with the much longer record at nearby Tsabong. The average precipitation event for Berry Bush is 10.66mm, though mean is a notoriously hapless measuring in dryland environments. The % difference from the 1934-1988 mean is extremely variable. 66.26 % of all precipitation events are 10mm or less, with 85.22 % of precipitation in distinct events of 20mm or less. This strengthens the principle for the fake precipitation events. Convective driven storms are irregular, low frequence high magnitude events. Over the analysed period there have merely been 10 events of =50mm ( stand foring merely 2.72 % of all precipitation events ) . Whilst the fake 50mm event for the undersoil pore infinite CO2 degrees represents a statistically rare event, the precipitation delivered in convective storms of =20mm represents, on norm, 47.76 % of one-year precipitation. Their importance can non be overlooked.

As Fig 2.5 illustrates, there is a pronounced seasonality of temperature at the survey site, coupled with a important diurnal temperature scope.

This is peculiarly marked in the winter months ( as in the survey period ) , where it is non uncommon for temperatures to fall below stop deading during the dark.

Chapter 3. Methodology

Field environmental variables

All research was conducted from 25/06/09 to 07/07/09. The Kalahari Sand site was studied from 25/06/09 through to 30/06/09, the Calcrete site from 02/07/09 through to 07/07/09. At each site a figure of background variables were measured.

Photosynthetically Active Radiation ( PAR ) was measured every four proceedingss utilizing a Delta-T QS2 Quantum detector, connected to a information lumberman ( Delta-T, UK ) . Air temperature, humidness and dew point were logged every 20 proceedingss utilizing a Measurement Science USB502 detector ( Adept Science, UK ) . A detector had been left on the KS site the old twelvemonth, entering from 12/08/09 to 23/06/09 ( as shown in Fig. 2.5 ) . Precipitation has been continuously recorded at the survey site, utilizing a basic rain gage, since September 1996 ( see Table 2.2 ) by a research worker trained by the Botswanan Meteorological Organisation.

Soil surface CO2 flux

Six portable in situ closed Chamberss ( ISCC ) were used to quantify dirt CO2 flux. It is good established that crusts are extremely sensitive to alterations in wet, temperature and perturbation ( Lange 2003a ; Zaady et al. , 2000 ) . Indeed, their photosynthetic ability is dependent on care of their structural unity, in peculiar the chlorophyll set ( Plate 3.4 ; Thomas 2009 pers. comm ) . Given the built-in importance to the survey of keeping the unity of the crust surface, specially designed ISCC were used ( besides referred to as ‘cells ‘ ) .

True in situ surveies look intoing dryland dirt CO2 fluxes are rare ; most are reliant on crust remotion and laboratory simulation. Until now, those undertaken in situ have tended to change the ambient environment ( significantly disrupting the soil/environment equilibrium ) or intolerably damage the crust surface, sabotaging the cogency of consequences. The ISCC used in the survey are alone, as patented by Hoon et Al ( 2009 ) . Their shaping features are shown in Table 3.1.

Each site was sampled over a 6-day period. Of the six ISCC at each site, one was a control cell, and the other five experimental cells. When non being used, the ISCC palpebras were removed to guarantee care of ambient environmental conditions. Each site was subjected to simulated precipitation pulsations. The fake precipitation governments ( Table 3.2 ; they represent a spectrum of average sums derived from analysis of Berry Bush ‘s 13-year precipitation record ) were applied to all experimental cells at 06:45, utilizing either a pipette or works spray, dependant on sum. Sample periods started at 07:00, 10:00, 13:00, 14:00, 16:00, 19:00, 22:00. A generic sample period was as follows: Attach ISCC palpebra. Sample ISCC internal CO2. Wait 45 proceedingss. Inject syringe, and manually ‘mix ‘ ( inject/purge x 2 to guarantee ISCC CO2 was adequately assorted, avoiding gas stratification. ) Sample CO2. Remove palpebra.

Subsoil Pore CO2 concentrations

A dirt cavity, next to each secret plan, was dug to look into alterations in pore infinite CO2 concentrations as a map of fake rainfall events. The methodological analysis used mostly followed that of Thomas & A ; Hoon ( 2010 ) . The square cavity ( 2 x 2m ) was about 1m deep. One face of the cavity was smoothed, and measured to a deepness of 75cm. Marks were made linearly every 5cm from 5cm to 75cm deepness, ensuing in 15 trying points. At each point, dirt temperature and dirt wet content were measured utilizing a Delta T multivariable investigation, inserted horizontally to about 20cm.

CO2 samples were taken at 6cm, 15cm, 30cm and 75cm. This involved infixing a gas sampling syringe ( Fig 3.1 ) into a predrilled sampling hole in the dirt cavity face at the four deepnesss, which were so left in situ for 24 hours. A 9mm sample was so taken utilizing a gas syringe, which was injected into a pre-evacuated Exetainer & A ; reg ; 4.5mm vial ensuing in a somewhat over-pressurised sample.

Samples were taken for 6 back-to-back yearss to see the response of pore infinite CO2 concentrations to a 50mm fake precipitation pulsation at 07:00 on the 2nd twenty-four hours. When non in usage, the cavity was covered with galvanised aluminum sheeting to forestall undue vaporization from the exposed dirt face. Thomas & A ; Hoon ( 2010 ) give a more elaborate account of the methodological analysis ‘s principle.

Lab Analysis

The Exetainer & A ; reg ; phials incorporating the CO2 samples for both surface dirt outflow and subsurface dirt pores were analysed on return to the United Kingdom. This was manually undertaken utilizing a portable Gas Chromatograph ( GC 3000, Agilent, USA ) “ using a silicon channel clip of flight and thermic conduction sensor ( TOF/TCD ) and high pureness He ( 99.999 % ) bearer gas. ” ( Thomas & A ; Hoon 2010:133 ) Each sample had a 45 2nd tally clip.

Scaning Electron Microscopy ( SEM ) images were taken in early November 2009 of the Kalahari Sand crust samples harvested on 07/07/09.

Chapter 4: Consequences and Analysis

Local Climate

The dry season winter June-July period selected for this survey was designed to maximize the effects of fake precipitation events. The moisture season normally ends in April, but 2009 was an anomalous twelvemonth. Precipitation recorded for the two-month period in 2009 is the highest on the 13-year record. Given the intense temporal variableness the information period is excessively limited to be considered representative. However, the part ‘s moisture season normally ends in early April ( Torrance, 1971 ) , and surely by May. The past two old ages can hence be considered anomalous. This anomalousness has great significance for the research conducted, as antecedent dirt wet was significantly elevated. Whilst no precipitation was recorded in the two six-day survey periods ( 25/06/09 – 07/07/09 ) , a 34.4mm convective storm occurred on the 10/06/09, the badness of which the landholder called “ unprecedented ” for the clip of twelvemonth. This is shown in Fig 4.1.

Consequently, the grade to which the effects of really low magnitude simulated precipitation events could be observed was questionable, given the unnatural antecedent wet conditions.

Diurnal variableness in temperature, humidness and dew point are shown in Figs. 4.2a ( Kalahari Sand site ) and 4.2b ( Calcrete pan site ) .

KS site has a maximal diurnal temperature scope of 35 & A ; deg ; C ( -3 & A ; deg ; C to 32 & A ; deg ; C ) , whilst the Calcrete site has a somewhat reduced scope of 30.5 & A ; deg ; C ( -1 & A ; deg ; C to 29.5 & A ; deg ; C ) . Nonetheless, both sites reflect the terrible conditions to which the crusts are exposed. Particularly revealingly, both sites show strong association between temperature and dew point in the early hours of each experimental twenty-four hours. In the early forenoon on yearss 3 ( see Plate 4.1 ) , 4, 5 and 6 KS records temperatures low plenty to allow dew formation. This quantitatively supports field observations ( Plate 4.1 ) , and suggests a antecedently unappreciated beginning of wet for BSCs in the Kalahari ( though Danin & A ; Orshan ( 1999 ) have investigated dew extensively in Israel ‘s Negev desert, and Ullmann & A ; Budel ( 2003 ) reexamine the extended literature on fog and dew inputs to BSC communities in the Namib ) .

Scaning Electron Microscope ( SEM )

Figs 4.3 a – e. are SEM images taken of cyanophyte dirt crust harvested in situ from the Kalahari Sand site on 07/07/09. These images are thought to be alone, for any old literature detailing SEM analysis on BSCs by and large uses unreal, laboratory environmental chamber cultivated dirt crusts.

Figs. 4.3 a-c show a harsh graduated table overview of the sample. They clearly illustrate the cyanophyte strands that bind together the vitreous silica grains, therefore playing an instrumental function in keeping the structural unity of BSCs. Fig 4.3d, and to a lesser extent figures 4.3a, B and degree Celsius seem to bespeak that the cyanophyte strands stabilise the dirt by enfolding vitreous silica grains, therefore fostering stableness by organizing a cohesive ‘mat ‘ around the vitreous silica grains.

The samples were analysed on 02/11/09, a little less than four months after being harvested. It is hence sensible to anticipate that the blue-green algae would be mostly dehydrated, with badly reduced exopolysaccharide content. However, 4.3e obviously demonstrates relentless adhesion of the vitreous silica grain to the cyanophyte sheath, despite dehydration. The binding function of cyanophyte strands is a rich country for farther survey.

CO2 flux

Fig. 4.4 illustrates the ambient atmospheric CO2 concentration. Each value represents the average CO2 concentration for all six cells at the start of each 45 min trying period ; basically a placeholder for background CO2 concentration.

It is evident that there is important fluctuation over clip. It is apparent that the KS site shows more fluctuation over clip than the Calcrete site. It besides appears that the KS site varies diurnally. This would match with observations that the KS site was located in close propinquity to an Acacia mellifera bush, which is likely to hold contributed to any CO2 values. Vascular works populations were non present in such propinquity on the calcrete site.

The deliberate tendency line appears to worsen exponentially. The ascertained diminution is sensible, given the precipitation informations for the two months prior to the survey. Following the comparatively high precipitation recorded in June, the continued elevated background CO2 most likely reflects a CO2 outflow associated with the most recent precipitation event. This supports the modelled diminution in background CO2 over the survey period, during which no precipitation was recorded. The CO2 degree appears to be levelling at 0.045 by around twenty-four hours nine ( 225 hours ) of the research, which represents an expected background degree during the dry season.

Surface CO2 flux: Kalahari Sands

Fig 4.5 illustrates the hourly mean for CO2 flux for all six experimental cells at the Kalahari site, over the class of the experimental period. Positive values represent outflow ( CO2 respired to the ambiance ) , whilst negative values represent segregation ( CO2 drawn down ) . Photosynthetically Active Radiation ( PAR ) is besides shown.

Four values ( around 24, 48, 72 and 120 hours ) show significantly raised standard divergences. They have each been skewed by an unnatural value ( all from different cells ) . The anomalous CO2 samples at these points were re-analysed by Gas Chromotography, yet were still unnatural. They most likely represent operator mistake, as they are excessively skewed to be conceivably natural. Though included for the intents of this graph, they have been excluded from statistical computations.

The mean should, in theory, be a hapless contemplation of the fluxes recorded at the cells, given that Cell 1 was the control and had no fake precipitation. However, as Figures. 4.6a – vitamin E show, the elevated ancestor dirt wet clearly restricted the ability to detect the effects of fake precipitation events. As such, no important difference was found under a Repeated-Measures ANOVA between Cell 1 flux and experimental cell fluxes. Whilst the ANOVA shows no difference between the experimental cells ( and therefore implies both low variableness and extremely replicable methodological analysis ) , that no difference can be established between the control and experimental cells is a really unexpected consequence.

Some general observations can be made. All experimental cells ( saloon Cell 6, Fig 4.6e ) show a strong period of segregation at 48 hours ( Cell 2 being much higher than the others ) . This is surprising, as given that this was night-time photosynthesis would non be expected. There is definite grounds of diurnal cycling, with most periods of net outflow happening during daytime, from around 10:00 boulder clay 16:00. Qualitatively, the graphs seem to bespeak a grade of variableness between cells ( the control cell is non shown, as no difference can be established between the control and experimental cells ) , though a Repeated-Measures ANOVA concludes that no important difference can be established between any of the cells. Variance therefore can non be statistically established on the Kalahari Sand site for the experimental cells. However, the ascertained fluxes contrast strongly with those of Thomas et Al ( 2008 ) , as no pronounced CO2 pulsations can be identified after fake precipitation events. Whilst this may non hold been expected at 0.5mm or 2mm, it is uncovering that no sizeably bigger CO2 pulsations can be identified after the larger fake precipitation events. This once more implies that the antecedent wet affected the ability to use fake precipitation events as an independent parametric quantity.

This is reiterated through comparing with Fig. 4.7, which compares Cell 1 ( the control ) with a average value for all experimental cells. This averaging is justified given the deficiency of statistically important difference observed between the experimental cells.

There are extended periods of comparative para in response ( though Cell 1 is more overdone ) between control and experimental fluxes ; notably, from 30 to 60 hours and 78 to 90. However, whilst at 48 hours both experimental and control cells show C segregation, the control cell net C addition is about dual that of the experimental cells. The 2nd noteworthy point of going, around 68 hours, once more illustrates control cell net C segregation, yet net experimental cell C outflow. The contrary is true at about 120 hours, where experimental cells show net segregation, and the control cell net outflow. The possible grounds for this are considered in Chapter 5.

On norm, the Kalahari Site experienced net CO2 outflow on all except the 2nd twenty-four hours, as shown in Fig 4.8 ( the mean day-to-day flux for all six cells ) . Whilst the fake precipitation may be triping cyanophyte photosynthesis, any net segregation appears to be being countered by CO2 respired from both surface and subsurface beginnings.

One peculiar country of involvement is the relationship between CO2 flux and temperature. It was hypothesised that a strong association would be between the two variables, given cyanophyte activity ‘s dependance on a threshold temperature to be metabolically active ( Lange 2003a ) . Fig. 4.9 illustrates the relationship between the two.

If old findings by Lange ( 2003a ) were to be replicated, it would be expected that photosynthesis, and therefore C segregation would increase as a map of increased temperature. However, there is no statistically important correlativity between flux and temperature. A general observation can be made that outflow seems to increase in a loose association with temperature, though this could quite probably be a contemplation of subsurface microbic respiration spreading to the surface, dissembling any cyanophyte photosynthetic effects. Extricating the influence of the two remains a really existent trouble.

Surface CO2 flux: Calcrete Pan

Fig 4.10 demonstrates tight diurnal yoke, with each value holding a comparatively low standard divergence in comparing with KS. The two exclusions are around 48 hours, where larger standard divergences and an unexpected addition in outflow can be considered unnatural. However, field notes at this trying point record trouble with the instrumentality, significance that the last sampling point before 48 hours ( the 14th ) was extended by about 18 proceedingss. This would explicate the ascertained addition in outflow, due to the unintended longer sampling clip period likely increasing CO2 concentration in the dirt Chamberss.

As with the KS, a mean has been used for all six cells. This is justified through consideration of Fig 4.11, which shows the single flux for all six cells.

As Fig. 4.11 indicates, there is a comparatively strong association between all cells. Of most involvement is that no important difference between cells can be identified in a Repeated-Measures ANOVA. As such, fluctuation can non be established between the experimental cells, and no difference can be found between the control and experimental cells. Whilst this suggests assurance in the methods used and their replicability, the deficiency of difference between experimental and control cells emphasises the challenges of utilizing fake precipitation as an experimental variable due to the antecedent wet conditions.

As Figs. 4.12a – vitamin E demonstrate, there are clear diurnal fluctuations in flux, which is in line with preexistent literature. Peak outflow stays mostly changeless throughout the period ( dropping somewhat in all cells on the 20mm simulation twenty-four hours ) . This is implicative of higher photosynthetic activity due to greater wet handiness stifling undersoil CO2 outflow. Peak flux recovers on the 6th twenty-four hours, when no extra H2O was applied.

A uncovering comparing can hence be made between the control and experimental cells, as shown on Fig. 4.13. The mean of all experimental cells shows close para to the control cell for the first three yearss of the experimental period. However, the association weakens as the experiment progresses.

Noteworthy divergences include the two periods of pronounced segregation for the experimental cells on the 4th and 5th twenty-four hours. This is non replicated by the control cell, and to a big extent this is in line with the theory that greater fake precipitation should bring on photosynthetic activity in the experimental cells, therefore pulling down C. The association appears so to be reinstated on the 6th twenty-four hours, where no precipitation is simulated ( and therefore photosynthesis dampened ) .

Fig 4.14 shows the day-to-day mean flux for all six cells, bespeaking a net outflow every twenty-four hours ( although about impersonal on Day 5 ) . This mostly reinforces theory that flux will worsen with increased precipitation, due to the increased C consumption by photosynthetic blue-green algae. The pronounced addition in flux on Day 6 may reflect a clip slowdown of the effects of the infiltration of the 20mm H2O added on the old twenty-four hours, therefore increasing undersoil respiration.

The relationship between temperature and flux ( for all cells ) is illustrated in Fig.4.15, which shows a strong association between the two variables. A Pearson trial shows a positive ( 0.652 ) correlativity, which is important at 99 % . One facet is of peculiar involvement. At the three trying points around 72 hours ( night-time ) , the temperature is over dual ( 10.7 & A ; deg ; C ) that of old darks at the same sampling times ( a mean of 4.7 & A ; deg ; C ) . The increased outflow seen in the last sample on the 3rd twenty-four hours may good reflect higher respiratory activity due to these out of the blue higher temperatures.

Surface CO2 flux: Comparison between Kalahari Sand and Calcrete sites

Given that no precipitation was simulated on the control cells, flux would non be expected to change significantly either between yearss or sites. As Fig. 4.16 shows, nevertheless, there is great discordance between the sites.

A T-test indicates that statistically important difference can be established between the two, at 99.9 % assurance. Given that all other conditions remained equal, this implies high flux variableness between the two survey sites. The Calcrete site shows comparatively replicable and consistent diurnal flux cycling, with similar extremum fluxes and merely somewhat differing peak segregation. KS, nevertheless, is far more variable, proposing a extremely complex mechanism driving flux, given that all other environmental factors stay comparatively changeless. Peak outflow is subdued in comparing with calcrete, with notably greater segregation over the six twenty-four hours period.

Fig 4.17 compares the KS and Calcrete mean fluxes for the experimental cells. For the first two experimental yearss, there is a little association between the two. Whilst the Calcrete site shows higher degrees of respiration, there is similarity in the way of tendency.

However, as the degree of fake precipitation additions on both sites, disparity additions between flux activity. Calcrete still maintains a grade of diurnal association, yet KS seems to lose any association. In some respects, the disassociation that begins at around 48 hours is because the Calcrete site shows an unexpected continued rise in respiration at the terminal of the 2nd twenty-four hours. Were a lessening to hold occurred, so association may hold been maintained for somewhat longer. However, the extremely varied responses in the last three yearss are most likely a contemplation of overdone activity due to higher fake precipitation. Overall, Fig. 4.17 does strongly connote that there is high fluctuation in surface dirt CO2 flux over a little spacial graduated table, although no statistically important difference could be established between the two.

Subsoil CO2 flux: Kalahari Sands

Fig 4.18 shows KS dirt cavity wet over the class of the 6-day experimental period ( H2O equivalent to a 50mm precipitation event was added, utilizing a spile, at 07:00 on the 2nd experimental twenty-four hours ) . All measurings are taken at 13:00.

Prior to wetting, wet conditions increase somewhat with deepness, demoing a crisp diminution below 60cm deepness ; this deepness appears to be a threshold for meaningful infiltration. Infiltration progresses over the class of the 6 yearss, with wet top outing in the top half of the dirt profile on the 3rd twenty-four hours.

Fig 4.19 shows temperature fluctuation at the same site ( all measurings taken at 13.00 ) . Whilst temperature decreases with deepness to around 30cm ( due to cut down radiative warming from sunstroke ) , it mostly balances out beyond this deepness. Day 2 shows a pronounced divergence, though given that this was the wetting twenty-four hours, reduced dirt temperature is a sensible observation.

Fig. 4.20 illustrates soil pore CO2 concentration on Day 1, the dry twenty-four hours. CO2 concentration systematically increases with deepness ( perchance a contemplation of increased Soil Organic Carbon ) .

The peculiarly big addition between 30cm and 75cm is most likely a contemplation of the increased propinquity to vascular works root webs. The celebrated CO2 concentration addition with deepness is temporally stable, with somewhat higher concentrations at earlier sample times at all deepnesss ( the 75cm concentration at 19:15 is most likely an anomalousness ) .

Fig 4.21 demonstrates CO2 concentration after the fake precipitation event. Note the immediate rise in CO2 concentration at 27 hours, at 15cm. This is an look of increased microbic respiration, although is merely observed at 6cm and 15cm ( perchance due to inadequate infiltration and temperature restriction ) . At 30 hours, increased CO2 concentration values are still found at the same deepness, though it is harder to demarcate, with farther clip, important differences between CO2 concentrations. That the highest Carbon dioxide values at 75cm are found halfway through the experiment is besides sensible, as by this clip at that place would hold been equal infiltration to excite respiration. Generally, though little, a CO2 pulsation can be noted following the wetting event, though it is smaller than in old literature ( Thomas et al 2008 ) .

A figure of statistical analyses were performed on the undersoil informations.

The moderate to strong negative correlativity between deepness and wet ( -0.572 ) is expected. However, a likewise moderate negative correlativity ( -0.508 ) between wet and CO2 concentration is surprising. The really strong positive correlativity ( 0.919 ) between deepness and CO2 is every bit dramatic. Similarly, no important correlativity between dirt temperature and CO2 concentration is an unannounced result. The weak negative correlativity between dirt wet and temperature is most likely explained by the confusing influence of surface vaporization.

A One-way ANOVA and a Post-Hoc Tukey trial besides indicate that CO2 concentration varies significantly with deepness. Whilst no important difference can be found between values at 6cm and 15cm, important differences can be found between all other groups. Again, 75cm is significantly different from all other deepnesss at 95 % assurance, farther underscoring the likely influence of vascular root respiration significantly increasing undersoil pore CO2 concentration.

Subsoil CO2 flux: Calcrete Pan

Soil wet fluctuation with deepness and clip at the calcrete dirt cavity is shown in Fig. 4.22 ( the same wetting event was applied on twenty-four hours two as in the KS site ) .

The immediate rise in wet station wetting ( Day 2 ) is noticeable, and this rapidly subsides with clip. In general, wet degrees are higher than KS ; whilst KS has a peak wet of 9.6 % , the upper profile on the Calcrete is 17.6 % after wetting. There is besides greater disparity at deepness at the Calcrete site, with the high value at 70cm testament to the greater infiltration capacity. By and large, the calcrete site has more variable dirt wet, both with deepness and clip.

Fig 4.23 shows dirt temperature discrepancy with deepness and clip. The findings are really similar to the KS site, although Day 2 is an exclusion. The upper profile values are about dual those of the old dry twenty-four hours, and the fake precipitation event does non look to hold induced any chilling.

Fig 4.24 shows CO2 concentration discrepancy with depth/time on the dry twenty-four hours. The absolute CO2 concentrations are really similar to those observed at the KS site. The ascertained consequences are mostly consistent with theory ( Luo & A ; Zhou 2006 ) , demoing CO2 concentration increasing throughout the twenty-four hours, reflecting the heater dirt temperatures. This so subsides in the afternoon. There is a noticeable rise in CO2 concentration with deepness ( as at the KS site ) , which is likely to be a contemplation of vascular works root respiration.

Fig 4.25 illustrates CO2 concentration discrepancy with deepness and clip after wetting. By and large, the effects of a pulse event can be observed, with CO2 concentration mostly increasing with clip at the upper profiles. However, such a clear relationship is non found at 30cm and 75cm. More regular sampling was possible than on the KS site, as the Calcrete dirt cavity had greater structural unity. The addition in CO2 concentration at 15cm at 24 hours is similar to that seen in KS at 27 hours at the same deepness, declarative mood of heightened microbic respiration.

Post-hoc trials on a One-way ANOVA indicate that there is a important difference ( 95 % assurance ) between CO2 concentrations at all deepnesss at all times. This compares with KS, where no important difference was found between 6cm and 15cm CO2 concentrations. The ANOVA consequences support the determination that deepness and CO2 concentration are positively correlated ( 0.883, 99 % assurance ) over all six yearss.

A figure of statistical trials for correlativity were undertaken, as shown in Table 4.2:

No correlativity was found between dirt wet and temperature, although deepness and dirt wet are reasonably negatively correlated ( -0.419 ) . A surprising negative correlativity ( -0.452, ) was identified between CO2 concentration and wet, which goes against theory that increased wet will excite higher microbic respiration, and hence higher CO2 concentration.

Chapter 5. Discussion

The CO2 fluxes observed from both the KS and Calcrete sites do non look to back up the ‘pulse-reserve ‘ paradigm. Fake precipitation had small discernable consequence on extremum flux, irrespective of the entire precipitation sum. This survey ( excepting anomalous values ) , reports extremum flux at KS of a small over 50 milligrams C M2 h1, observed on both the 2mm and 20mm simulations. At the Calcrete site the consequences are lower, the flux over the class of the six yearss top outing at somewhat under 40 milligrams C M2 h1. This contrasts with the findings of Thomas & A ; Hoon ( 2010 ) , who report a peak flux from a similar Kalahari Sand site in July 2007 of 339.2 milligrams C M2 h1. Whilst this was recorded after fake rainfall of 120mm, higher than in this survey, fluxes of 65.6 milligrams C M2 h1 were observed after merely 1.4mm application. In a wider context, the positive outflows observed at both the KS and Calcrete sites are perceptibly lower than those reported in Raich & A ; Tufekcioglu ‘s ( 2000 ) planetary analysis. The lower flux is likely a contemplation in portion of the limited organic content of the dryland dirt, as suggested by Thomas & A ; Hoon ( 2010 ) . As such, the lower fluxes observed in this survey are extremely declarative of an ecosystem that is limited by C, instead than wet handiness.

This theory is supported by the exponentially worsening values recorded for the ambient atmospheric CO2 ( see Fig. 4.4 ) which are in line with the impression that the survey site was retrieving from a big pulse event, most probably associated with the 34.4mm rainstorm of 10/06/09. At both sites, flux does non look to change as a map of fake precipitation sum. It would justifiably be assumed that a important difference could be faithfully established between the control and experimental cells, yet none exists ( as shown in Figs. 4.7 and 4.13 ) . Whilst Thomas et Al ( 2008 ) , Thomas & A ; Hoon ( 2010 ) and Wang et Al ( 2009 ) suggest that CO2 fluxes in Kalahari dirts are likely to be wet limited for the bulk of the twelvemonth, the same decision can non be supported on the footing of this survey ‘s findings. Rather, the extremely elevated antecedent wet conditions seem to hold undermined the efficaciousness of using fake precipitation events as an independent experimental parametric quantity. Though the ecosystem is normally moisture limited in June, it appeared to be mostly C limited in June 2009 owing to high rainfall prior to experimentation.

Given the low vascular works populations, the BSCs are likely to be the lone important beginning of C at the dirt surface. Belnap et Al. ( 2003b ) study that C arrested development by blue-green algae during photosynthesis can ensue in dirt organic C additions of up to 300 % , being secreted as extracellular polymeric substances. Whilst Evans & A ; Lange ( 2003 ) recognize the potentially important part of BSCs to carbon shops in drylands, the Kalahari remains noteworthy for its comparative deficiency of informations. The antecedent wet conditions following the rainfall prior to this survey seem to bespeak, nevertheless, that the dirt C shops had been mostly dog-tired prior to experimentation. This would partly explicate why few pulsations were observed following the application of H2O ; surely, it is non possible to distinguish between different magnitudes of precipitation events. This does non propose that the pulse-reserve paradigm is invalid, but raises the likeliness of the Kalahari being both wet and C limited. In many respects hence, whilst the survey failed to fulfill its purpose of showing precipitation event driven CO2 fluxes, it gives an penetration into the impact of low frequence, high magnitude unseasonal storm events on C cycling.

A cardinal focal point of the survey was to analyze variableness both within sites and between the two sites. The findings of the T-Test show that the six-day flux consequences for the control cells at each site are significantly ( 99.9 % assurance ) different. This is deserving some consideration. As all other conditions remained equal, the graduated table of the disparity seen between the two is striking. Numerous environmental variables, both surface and subsurface, are likely to hold contributed towards this. Surely, the propinquity of the KS site to a ample Acacia Mellifera works may hold influenced samples. As Dougill & A ; Thomas ( 2004 ) note, crusts develop preferentially under this works, so C exchange is improbable to be spatially even. This has peculiar deductions given the recent shrub invasion recorded in the Kalahari ( Dougill 2002 ) . The root web is likely to hold impacted undersoil pore CO2 values, and surface vascular CO2 exchange is once more likely to hold affected trying. Field observations besides suggest that the crusts at the two sites were in a different phase of sequence. Despite the fence at the KS site, guaranting that the crusts were protected for the twelvemonth prior to analyze, they were mostly destroyed in summer 2008 during research ( Thomas 2009, pers comm ) . However, although the Calcrete site was non protected, pastoral agriculture had stopped on the site about 20 months prior to this probe. There were big countries of clearly integral, Stage 3 crust ( with noticeable surface roughening and microtopography ) , whilst merely Stage 2 crust was identified at KS. The influence of crust sequence on its function as an atmospheric boundary has been noted ( Housman et al 2006 ; Zaady et Al, 2000 ) , though Zaady et Al ( 2000 ) suggest that higher degree of sequence may take to higher flux rates. This suggestion was non borne out in this survey ( average flux was higher at the KS site ) , where more consistent fluxes were observed on the Calcrete site between all cells. This determination is in line with the theory that greater homogeneousness in BSC surface crust may take to more replicable CO2 exchange ( Housman et al, 2006 ) . Equally, the consequence of the Calcrete site ‘s more consistent BSC surface coverage on H2O infiltration should non be overlooked. On the 5mm and 20mm simulation yearss, infiltration varied greatly between cells on the KS site, yet no such fluctuation was noted at the Calcrete site. Indeed, infiltration took longer at the Calcrete site, which is potentially a contemplation of both the more developed BSC population and the significantly higher antecedent dirt wet ( pre-wetting, 8.5 % as compared to KS 3.5 % ) . It is executable that the accordingly more unvarying infiltration may partly explicate the more consistent observed fluxes between cells. However, this is merely a partial account, for the influence of fake precipitation pulsations is likely to hold been reduced by the high ancestor dirt wet.

Whilst the pulsations of positive CO2 flux may be less than expected, some interesting observations can be made on negative flux ( carbon consumption ) . At the Calcrete site, some distinction can be established between the control and experimental cells. Whilst negative flux does happen at the start of Day 3 for the control cell ( -9.7 milligram CO2 m2 h1 ) , it is less than half the average experimental negative flux ( -20.7 milligram CO2 m2 h1 ) . However, field observations for this twenty-four hours observe important dewfall that dark, as supported by the temperature/dew point informations. Whilst non quantified, it is likely that sufficient dew was present to set up low metabolic activity for the control cell, therefore explicating the C consumption. Sing the experimental cells, some negative flux may hold been expected on the 2nd twenty-four hours ( 0.5mm event ) , for Lange ( 2003a ) notes that optimum photosynthesis, and therefore CO2 arrested development, can happen with 0.5mm rehydration ( though this is extremely species dependant ) . The fact that the negative flux is recorded instantly after wetting once more confirms blue-green algae ‘s rapid metabolic response to hydration, as suggested by Garcia-Pichel & A ; Pringault ( 2001 ) . Surely, the stronger negative flux for the experimental instead than command cells at 2mm rehydration implies photosynthetic reactivation. Sing experimental Day 5 ( 20mm event ) , another important negative flux can be identified. As Fig. 4.12e reiterates, this flux is unusually consistent ( both over clip and in magnitude ) between all experimental cells. This determination is testament to the high temporal declaration of the trying methodological analysis developed by Hoon et Al ( 2009 ) . Such nuance of flux would non be evident in the surveies of Wang et Al ( 2007, 2009 ) and others, which use one flux measuring as representative for the whole twenty-four hours.

The high segregation counters old consequences found by Thomas et Al ( 2008 ) . By and large, at higher wetting events, any increased photosynthesis which may take to a CO2 drawdown is masked by increased undersoil respiration. This does non look to be the instance in this survey, beef uping statements that the ecosystem may be C exhausted. Equally, the strength of the CO2 segregation is surprising, for Lange ( 2003b ) studies photosynthetic suppression at suprasaturation. However, Lange ‘s work is mostly focused on lichenous BSCs ; our consciousness of species differentiated responses is at best presently limited, and farther research is needed to find optimum hydration for photosynthesis for the blue-green algae at this survey site.

The KS site shows more variable periods of CO2 consumption, and consumption is less consistent between cells. Most surprisingly, uptake appears to happen at all experimental cells merely before 48 hours. The trying point was at 22:00, so photosynthetic activity is non an obvious account. Had merely one cell experienced segregation so this observation could possibly be dismissed, yet given that it is replicated over five cells it is deserving some consideration. Mairs ( 2009, pers comm ) studies in situ consequences from the Kalahari, where BSC blue-green algae were unnaturally darkened, and wetted with a 5mm event every other twenty-four hours. Using IR Spectrometry, it was found that after six yearss surface chlorophyll content really increased in darkened conditions, explained by the ability of chlorophyll to glide vertically within the dirt column on EPS sheaths. It is possible that the experimental ISCC used at the KS site may hold limited photosynthetically active radiation to the BSC, due both to shadowing from the Acacia Mellifera and shadowing from the ISCC themselves. Consequently, a time-lag may hold occurred in segregation. However, the fact that the control cell shows important segregation at 22:00 on the 2nd and 19:00 on the 3rd experimental twenty-four hours ( -61.6 milligram CO2 M2 h1 and -76.4 mg CO2 M2 h1 severally ) further complicates the relationship. As all other conditions remain equal, there is no obvious theory to explicate this. It is possible that light independent ( dark ) photosynthesis may be happening ; Belnap ( 2003 ) discusses nitrogen arrested development in cyanophyte BSC, where under research lab conditions dark arrested development has been observed. Equally, Stone ( 2008 ) cites episodes of night-time CO2 consumption in China ‘s Gurbantunggut Desert. This is mostly attributed to the high alkalinity of the dirt, yet the pH at the KS site was 5.9, so this theory is improbable to be applicable in this case. This is an country rich for future research, as our apprehension is really much in its babyhood.

Over the class of the six yearss, both of the survey ‘s sites show net outflow. The six cell daily intend ( Fig 4.8 ) shows net CO2 consumption on the 2nd twenty-four hours at the KS site, although this is skewed by the out of the blue high consumption of the control cell. However, the discrepancy at the KS site as shown by the mistake bars, is much higher than the Calcrete site, so any decisions should be probationary. The Calcrete site ( Fig 4.14 ) shows a diminution in the average net flux from the 2nd to fifth experimental twenty-four hours, which is consistent with theory that increased wet will take to increased photosynthetic consumption. As such, the rise in the 6th twenty-four hours may good be explained by the 20mm event being sufficiently big to infiltrate to a deepness whereby it could do subsoil respiration, and therefore a larger efflux the undermentioned twenty-four hours ( by which clip surface wet would hold declined to the extent that it might restrict photosynthesis ) . Sing the overall segregation at both the KS and Calcrete sites, there is a contrast to the work of Wohlfahrt et Al ( 2008 ) , who report net one-year CO2 consumption in the Mojave. However, there is likely to be seasonal discrepancy in flux activity, given the important scope in both temperature and wet handiness, so any decisions from this survey are by no agencies representative.

The environmental variables regulating BSC metamorphosis and dirt respiration have been widely studied ( Belnap & A ; Lange, 2003 ) . Whilst this survey recorded Photosynthetically Active Radiation ( PAR ) , equipment failure resulted in informations loss for the full KS site. As such, a placeholder was used, taking the PAR measurings from mid-July 2008. Consequently, i