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# The evolution of the stellar populations in low surface brightness galaxies

## 1 Introduction

Deep searches for field galaxies in the local universe have revealed the existence of a large number of galaxies with low surface brightnesses that are hard to detect against the night sky (see the reviews by Impey & Bothun 1997 and Bothun, Impey & McGaugh 1997). Many different kinds of LSB galaxies have so far been found, including giant LSB galaxies (e.g. Sprayberry et al. 1995) and red LSB galaxies (O’Neil et al. 1997), but the most common type seems to be “blue LSB galaxies”: late-type, disk-dominated spirals with central surface brightnesses $`\mu _0(B)\begin{array}{c}>\\ \end{array}23`$ mag arcsec-2.

In this paper, we will concentrate on these late-type LSB spirals. Observations show that they are neither exclusively dwarf systems nor just the faded counterparts of “normal” high surface brightness (HSB) spirals (de Blok et al. 1996, hereafter dB96). In many cases, LSB galaxies follow the trends in galaxy properties found along the Hubble sequence towards very late types. These trends include increasingly blue colours (e.g. Rönnback 1993; McGaugh & Bothun 1994; de Blok et al. 1995 \[hereafter dB95\]; Bell et al. 1999), decreasing oxygen abundances in the gas (e.g. McGaugh 1994; Rönnback & Bergvall 1995; de Blok & van der Hulst 1998a), and decreasing Hi surface densities (from type Sc onwards; e.g. van der Hulst et al. 1993, hereafter vdH93). Despite the low gas densities, LSB spirals rank among the most gas-rich disk galaxies at a given total luminosity as their Hi disks in general are extended (Zwaan et al. 1995; dB96). The fact that LSB galaxies still have large reservoirs of gas together with their low abundances suggest that the amount of star formation in the past cannot have been very large (vdH93; Van Zee et al. 1997). Clearly, LSB galaxies are not the faded remnants of HSB spirals.

The unevolved nature of LSB spirals can be interpreted in various ways. For instance, LSB spirals could be systems whose stellar population is young and in which the main phase of star formation is still to occur. Alternatively, the stellar population in LSB spirals could be similar to those in HSB galaxies but with a young population dominating the luminosity. This has been explored and modelled in studies by e.g. Knezek (1993), Jimenez et al. (1998), dB96, Gerritsen & de Blok (1999) and more recently in two papers by Bell et al. (1999) and Bell & de Jong (1999). These studies all indicate a scenario in which LSB galaxies are unevolved systems with low surface densities, low metallicities and a constant or increasing star formation rate (cf. Padoan et al. 1997, however, for a different opinion).

In this paper we address these various scenarios for the evolution of LSB galaxies by detailed modelling of their spectro-photometric and chemical properties. Model results are compared directly with the observed colours, gas phase abundances, gas contents, and current star formation rates of LSB galaxies. Using simple model star formarion histories we confirm the results of previous studies and show that the ratio of young stars to old stars in LSB galaxies is larger than typically found in HSB galaxies.

In reality, however, star formation is unlikely to proceed as smoothly as the various population synthesis models suggest. Gerritsen & de Blok (1999) point out that small surges in the star formation rate are likely to be very important in determining the observed colours of LSB galaxies. In this paper we will pay special attention to the influence of small bursts of star formation on the optical colours, and show that they can have a significant influence on the inferred properties of LSB galaxies.

This paper is organized as follows. We briefly compare observational data of LSB galaxies with those of spirals and dwarf galaxies in Sect. 2. In Sect. 3, we describe the ingredients of the galactic evolution model we used. Some general properties of the model are presented in Sect. 4. The comparison with observational data is presented and discussed in Sect. 5. The impact of small amplitude star formation bursts in LSB galaxies is investigated in Sect. 6 and predicted star formation rates are compared with the observations in Sect. 7. A short discussion of where LSB galaxies fit in the grand scheme of galaxy formation and evolution is presented in Sect. 8. A short summary is given in Sect. 9.

## 2 Observational characteristics of LSB galaxies

### 2.1 Sample selection

We use the sample described in dB96. This consists of 24 late-type LSB galaxies (inclinations up to $`60^{}`$), taken from the lists by Schombert et al. (1992) and the UGC (Nilson 1973). The sample is representative for the LSB galaxies generally found in the field. We selected a subsample of 16 LSB galaxies for which good data are available. For these systems, optical data have been taken from dB95, Hi data from dB96, and abundance data from McGaugh & Bothun (1994) and de Blok & van der Hulst (1998a).

The galaxy identification and Johnson $`UBV`$ and Kron-Cousins $`RI`$ are given in columns (1) to (6) in Table 1 (a Hubble constant of $`H_0=100`$ km s-1 Mpc-1 was used). Column (7) lists the $`B`$-band luminosities; column (8) the Hi-mass-to-light ratio; column (9) and (10) the Hi and dynamical masses respectively. The gas fraction $`\mu _{\mathrm{rot}}`$ in column (11) is given by $`M_{\mathrm{gas}}`$/$`(M_{\mathrm{gas}}+M_{,\mathrm{max}})`$ where $`M_{,\mathrm{max}}`$ denotes the mass of the stellar component obtained from maximum disk fitting of the rotation curve (dB96) (see Sect. 2.6). The mean \[O/H\] abundances are listed in column (12).

### 2.2 Magnitudes and colours

We compare in Fig. 1$`a`$ and $`b`$ magnitudes and broadband colours of LSB galaxies to those of late-type HSB face-on spirals (de Jong & van der Kruit 1994) and dwarf galaxies (Melisse & Israel 1994; Gallagher & Hunter 1986, 1987). The usual distinction between HSB and LSB galaxies is purely an artificial one since normal galaxies along the Hubble sequence show a continuous range in central surface brightness, i.e. ranging from values around the Freeman value for early type systems to the very faint values observed for the late-type LSB galaxies in our sample (e.g. de Jong 1995; dB95). For the purpose of this paper we will continue to use the distinction as a shorthand, where “HSB galaxy” should be interpreted as “typical Hubble sequence Sc galaxy.” Fig. 1$`a`$ and $`b`$ show that LSB galaxies are generally bluer and fainter than their HSB counterparts (e.g. McGaugh 1992; vdH93; dB95).

### 2.3 Abundances

Estimates of the ISM abundances in LSB galaxies predominantly rely on abundance determinations of their Hii regions. Oxygen abundances of Hii regions listed in Table 1 are taken from McGaugh (1994) and de Blok & van der Hulst (1998a).

We assume that the Hii-region abundances on average are a reasonable indicator of the ISM abundances within a given LSB spiral. The intrinsic scatter in \[O/H\] among different Hii regions within a given LSB galaxy is usually of the order 0.2 dex (e.g. McGaugh 1994). Bell & de Jong (1999) find that the gas metal abundances in LSB galaxies generally trace the stellar metallicities quite well, with the stars being $`0.4`$ dex more metal-poor.

In Fig. 1c we compare mean \[O/H\] abundances of Hii regions in LSB galaxies with those in HSB spirals (Zaritsky et al. 1994) and dwarf galaxies (Melisse & Israel 1994; Gallagher & Hunter 1986, 1987). On average, LSB galaxies generally follow the correlation between the characteristic gas-phase abundance and luminosity as found for HSB spirals (Zaritsky et al. 1994), but with a large scatter. The range in \[O/H\] at a given $`B`$ magnitude is nearly one dex. This scatter is probably related to evolutionary differences among the LSB galaxies of a given $`B`$ magnitude (e.g. in the ratio of old to young stellar populations) and/or the Hii regions they contain.

### 2.4 Extinction

Observational studies by (amongst others) Bosma et al. (1992), Byun (1992), Tully & Verheijen (1997) show that late-type spirals appear transparent throughout their disks. This supports the idea that face-on extinction in LSB galaxies is relatively low, i.e. $`E_{BV}\begin{array}{c}<\\ \end{array}0.1`$ (e.g. McGaugh 1994). This is consistent with findings by Tully & Verheijen (1998), who studied a large sample of galaxies in the UMa cluster and found the LSB galaxies in their sample to be transparent, whereas non-negligible extinction was present in the HSB galaxies.

In this study we will ignore the possible effects of dust on the colours and magnitudes. See Sect. 5.3 for a discussion.

### 2.5 Gas masses

We compare in Fig. 1d the observed atomic gas masses $`M_\mathrm{g}1.4M_{\mathrm{HI}}`$ (corrected for helium) of LSB galaxies with those of HSB galaxies and dwarfs. Strictly speaking we should also take the molecular gas mass into account. However, so far no CO-emission has been detected in late-type LSB galaxies (Schombert et al. 1990; de Blok & van der Hulst 1998b, but see Knezek 1993 for a discussion on giant LSB galaxies). The amount of molecular gas in LSB galaxies is probably small, even though relatively high CO/H2 conversion factors may apply in these low metallicity galaxies (Wilson 1995, Mihos et al. 1999). We conclude that, on average, LSB galaxies are considerably more gas-rich (up to a factor $`3`$) than HSB galaxies of the same $`B`$ magnitude (see also Fig. 1g).

### 2.6 Gas fractions and mass-to-light ratios

We show in Figs. 1g and h the distribution of the mass-to-light ratio $`M_{\mathrm{HI}}/L_B`$ vs $`(BV)`$ and vs $`\mu _{rot}`$, respectively. LSB galaxies exhibit considerably higher $`M_{\mathrm{HI}}/L_B`$ ratios than HSB spirals (cf. Table 1 and McGaugh & de Blok 1997).

We would however like to know the ratio of gas mass to stellar mass, i.e. the gas fraction $`\mu _1M_{\mathrm{gas}}`$ / $`(M_{\mathrm{gas}}+M_{\mathrm{stars}}`$ of a galaxy. This involves the conversion of the observed galaxy luminosity to stellar mass. However, the mass-to-light ratio of the underlying stellar population is generally not well known and, in fact, is an important quantity to determine. Assuming a fixed value of the mass-to-light ratio will however introduce artificial trends since this ratio is expected to vary among galaxies having different star formation histories (e.g. Larson & Tinsley 1978).

As a partial solution we have determined the gas fraction $`\mu _{\mathrm{rot}}M_{\mathrm{gas}}/(M_{\mathrm{gas}}+M_{,\mathrm{max}})`$ where $`M_{,\mathrm{max}}`$ denotes the mass of the stellar component obtained from maximum disk fitting of the rotation curve (dB96). This method likely overestimates the contribution of the luminous stellar disk to the observed total mass distribution by about a factor of 2 (e.g. Kuijken & Gilmore 1989; Bottema 1995) and, therefore, also provides a lower limit to the actual gas fraction. In summary, while $`\mu _1`$ is a model quantity, $`\mu _{\mathrm{rot}}`$ is determined from observations.

Figs. 1e and 1f show the distribution of the present-day gas fraction $`\mu _{\mathrm{rot}}`$ and total gas mass vs $`(BV)`$. It can be seen that LSB galaxies and dwarfs exhibit larger gas fractions than HSB spirals. Taking into account the factor of $`2`$ overestimate, values for LSB galaxies are typically $`\mu _1\begin{array}{c}>\\ \end{array}0.5`$ and $`\mu _10.1`$ for HSB spirals.

We conclude that the available evidence supports of the view that LSB galaxies are relatively unevolved systems compared to HSB spirals. This agrees well with the fact that LSB galaxies usually display properties intermediate to those of HSB spirals and dwarf galaxies.

## 3 Model description and assumptions

We here summarise the galactic evolution model used (for a more extensive description see van den Hoek 1997, hereafter vdH97). We concentrate on the stellar contribution to the total galaxy luminosity in a given passband (other contributions are neglected). For a given star formation history (SFH), we compute the chemical enrichment of a model galaxy by successive generations of evolving stars. To derive the stellar luminosity in a given passband at a given age, we use a metallicity dependent set of theoretical stellar isochrones as well as a library of spectro-photometric data.

### 3.1 Chemical evolution model

We start from a model galaxy initially devoid of stars. We follow the chemical enrichment during its evolution assuming stars to be formed according to a given star formation rate (SFR) and initial mass function (e.g. a power law IMF: d$`N`$/d$`mM(m)m^\gamma `$). Both stellar and interstellar abundances as a function of galactic evolution time $`t_{ev}`$ are computed assuming that the stellar ejecta are returned and homogeneously mixed to the ISM at the end of the individual stellar lifetimes (i.e. relaxing the instantaneous recycling approximation; see Searle & Sargent 1972). A description of the set of galactic chemical evolution equations used can be found in e.g. Tinsley (1980) and Twarog (1980), see also vdH97. This model is a closed-box model.

We follow the stellar enrichment of the star forming galaxy in terms of the characteristic element contributions of Asymptotic Giant Branch (AGB) stars, SNII and SNIa. This treatment is justified by the specific abundance patterns observed within the ejecta of these stellar groups (see e.g. Trimble 1991; Russell & Dopita 1992). A detailed description of the metallicity dependent stellar lifetimes, element yields, and remnant masses is given by vdH97 and van den Hoek & Groenewegen (1997). We compute the abundances of H, He, O, and Fe, as well as the heavy element integrated metal-abundance $`Z`$ (for elements more massive than helium), during the evolution of the model galaxy. Both the SFH, IMF and resulting element abundances as a function of galactic evolution time are used as input for the spectro-photometric evolution model described below.

Boundary conditions to the chemical evolution model are the galaxy total mass $`M_{\mathrm{tot}}`$, its evolution time $`t_{\mathrm{ev}}`$, and the initial gas abundances. Note that solutions of the galactic chemical evolution equations are independent of the ratio of the SFR normalisation and $`M_{\mathrm{tot}}`$. Primordial helium and hydrogen abundances are adopted as $`Y_\mathrm{p}=0.232`$ and $`X=0.768`$ (cf. Pagel & Kazlauskas 1992). Initial abundances for elements heavier than helium are initially set to zero.

We list the main input parameters in Table 2, i.e. the adopted IMF-slope, minimum and maximum stellar mass limits at birth as well as the progenitor mass ranges for stars ending their lives as AGB star, SNIa, and SNII, respectively. For simplicity, we assume the stellar yields of SNIb,c to be similar to those of SNII. Furthermore, we assume a fraction $`\nu ^{\mathrm{SNIa}}=0.015`$ of all white dwarf progenitors with initial masses between $`2.5`$ and 8 $`M_{}`$ to end as SNIa. These and other particular choices for the enrichment by massive stars are based on similar models recently applied to the chemical evolution of the Galactic disk (e.g. Groenewegen et al. 1995; vdH97).

### 3.2 Spectro-photometric evolution model

The total luminosity of a galaxy in a wavelength interval $`\mathrm{\Delta }\lambda `$ is determined by: 1) the stellar contribution $`L_{}`$, 2) the ISM contribution $`L_{\mathrm{ism}}`$ (e.g. Hii-regions, high-energy stellar outflow phenomena, etc.), and 3) the contribution due to absorption and scattering $`L_{\mathrm{ext}}`$:

$$L_{\mathrm{gal}}^{\mathrm{\Delta }\lambda }(t)=L_{}+L_{\mathrm{ism}}L_{\mathrm{ext}}$$
(1)
where each term in general is a complex function of galactic evolution time. We concentrate on the stellar contribution and neglect the last two terms in Eq. (1). Then, the luminosity in a wavelength interval $`\mathrm{\Delta }\lambda `$ at galactic evolution time $`t=T`$ can be written as:

$`L_{\mathrm{gal}}^{\mathrm{\Delta }\lambda }(t=T)=`$ (2)
$`{\displaystyle _{t=0}^T}{\displaystyle _{m_\mathrm{l}}^{m_\mathrm{o}(Tt)}}`$ $`L_{}^{\mathrm{\Delta }\lambda }(m,Z(t),Tt)S(t)M(m)\mathrm{d}m\mathrm{d}t`$
where $`m_\mathrm{l}`$ denotes the lower stellar mass limit at birth, $`m_\mathrm{o}(t)`$ the turnoff mass for stars evolving to their remnant stage at evolution time $`t_{\mathrm{ev}}`$, and $`L_{}^{\mathrm{\Delta }\lambda }`$ the luminosity of a star with initial mass $`m`$, initial metallicity $`Z(t)`$, and age ($`Tt`$). We assume a separable SFR: $`S(m,t)=S(t)M(m)`$ where $`S(t)`$ is the star formation rate by number \[yr-1\] and $`M(m)`$ the IMF \[$`M_{}`$-1\]. By convention, we normalise the IMF as $`M(m)dm=1`$ where the integration is over the entire stellar mass range \[$`m_\mathrm{l}`$, $`m_\mathrm{u}`$\] at birth (cf. Table 2).

Starting from the chemical evolution model described above, we compute the star formation history $`S(m,t)`$, gas-to-total mass-ratio $`\mu (t)`$, and age-metallicity relations (AMR) $`Z_i(t)`$ for different elements $`i`$. Thus, at each galactic evolution time $`t_{\mathrm{ev}}`$ the ages and metallicities of previously formed stellar generations are known. To derive the stellar passband luminosity $`L_{}^{\mathrm{\Delta }\lambda }`$ we use a set of theoretical stellar isochrones, as well as a library of spectro-photometric data. Stellar evolution tracks provide the stellar bolometric luminosity $`L_{}^{\mathrm{bol}}`$, effective temperature $`T_{\mathrm{eff}}`$, and gravity $`g`$, as a function of stellar age for stars with initial mass $`m`$ born with metallicity $`Z_{}`$. We compute Eq. (2) using a spectro-photometric library containing the stellar passband luminosities $`L_{}^{\mathrm{\Delta }\lambda }`$ tabulated as a function of $`T_{\mathrm{eff}}`$, $`\mathrm{log}g`$, and $`Z_{}`$ (see below).

The turnoff mass $`m_\mathrm{o}(Tt)`$ occuring in Eq. (2) depends on the metallicity $`Z(t)`$ of stars formed at galactic evolution time $`t_{\mathrm{ev}}`$. For instance, the turnoff mass for stars born with metallicity $`Z=10^3`$ at a galactic age of $`t_{\mathrm{ev}}=14`$ Gyr is $`m_\mathrm{o}0.8`$ $`M_{}`$ (e.g. Schaller et al. 1992). This value differs considerably from $`m_\mathrm{o}0.95`$ $`M_{}`$ for stars born with metallicity $`Z=Z_{}`$. Such differences in $`m_\mathrm{o}`$ affect the detailed spectro-photometric evolution of a galaxy by constraining the mass-range of stars in a given evolutionary phase (e.g. horizontal branch) at a given galactic evolution time. In the models described below, we explicitly take into account the dependence of $`m_\mathrm{o}(t)`$ on the initial stellar metallicity $`Z_{}`$ (see vdH97)

### 3.3 Stellar evolution tracks and spectro-photometric data

We use the theoretical stellar evolution tracks from the Geneva group (e.g. Schaller et al. 1992; Schaerer et al. 1993). These uniform grids cover large ranges in initial stellar mass and metallicity, i.e. $`m=0.05120`$ $`M_{}`$ and $`Z=0.040.001`$, respectively. These tracks imply a revised solar metallicity of $`Z_{}=0.0188`$ with $`Y_{}=0.299`$, and $`\mathrm{\Delta }Y/\mathrm{\Delta }Z=3.0`$ for a primordial He-abundance of $`Y_\mathrm{p}=0.232`$. For stars with $`m>7`$ $`M_{}`$, these tracks were computed until the end of central C-burning, for stars with $`m=25`$ $`M_{}`$ up to the early-AGB, and for $`m<1.7`$ $`M_{}`$ up to the He-flash. For stars with $`m\begin{array}{c}<\\ \end{array}0.8`$ $`M_{}`$, we used the stellar isochrone program from the Geneva group (Maeder & Meynet, priv. comm.; see also vdH97 ).

To cover the latest stellar evolutionary phases (i.e. horizontal branch (HB), early-AGB, and AGB) for stars with $`m\begin{array}{c}<\\ \end{array}8`$ $`M_{}`$, we extended the tracks from Schaller et al. with those from Lattanzio (1991; HB and early-AGB) for $`m12`$ $`M_{}`$, and from Groenewegen & de Jong (1993; early-AGB and AGB) for $`m18`$ $`M_{}`$. These tracks roughly cover the same metallicity range as the tracks from the Geneva group. Corresponding isochrones were computed on a logarithmic grid of stellar ages, covering galactic evolution times up to $`t_{\mathrm{ev}}14`$ Gyr. Isochrones are linearly interpolated in $`m`$, log $`Z`$, and log $`t`$.

The spectro-photometric data library that we use is based on the Revised Yale Isochrones and has been described extensively by Green et al. (1987). These data include stellar $`UBVRI`$ Johnson-Cousins magnitudes covering the following ranges in $`T_{\mathrm{eff}}[\mathrm{K}]=2800`$ to 20000, log $`g`$ \[cm s-2\] $`=0.5`$ to 6, and log ($`Z/Z_{})2.5`$ to $`+0.5`$. Corresponding spectro-photometric data for stars with $`T_{\mathrm{eff}}>20000`$ K have been adopted from Kurucz (1979) at solar metallicity, covering $`T_{\mathrm{eff}}=2000050000`$ K.

Although a detailed description of the tuning and calibration of the adopted photometric model is beyond the scope of this paper, we note that the model has been checked against various observations including integrated colours and magnitudes, luminosity functions, and colour-magnitude diagrams of Galactic and Magellanic Cloud open and globular clusters covering a wide range in age and metallicity (see vdH97).

## 4 General properties of the model

Before discussing the modelling of LSB galaxies, we first describe the general behaviour of the models assuming a few different star formation scenarios.

We start from a model galaxy with initial mass $`M_\mathrm{g}(t=0)=10^{10}`$ $`M_{}`$, initially metal-free and devoid of stars. The chemical and photometric evolution of this galaxy is followed during an evolution time $`t_{\mathrm{ev}}=14`$ Gyr, assuming one of the theoretical star formation histories discussed below. Unless stated otherwise, we assume that stars are formed according to a Salpeter (1955) IMF (i.e. $`\gamma =2.35`$) with stellar mass limits at birth between 0.1 and 60 $`M_{}`$ (cf. Table 2).

The following star formation histories are considered: 1) constant, 2) exponentially decreasing, 3) linearly decreasing, 4) exponentially increasing, and 5) linearly increasing. Normalised SFRs and resulting age-ISM metallicity relations are shown in Fig. 2. For each model, the amplitude of the SFR is chosen such that a present-day gas-to-total mass-ratio $`\mu _1=0.1`$ is achieved.

In columns (2) to (6) of Table 3, we list the functional form of the SFR, average past and current SFRs, the ratio of current and average past SFRs $`\alpha `$, and the present-day mass averaged ISM oxygen abundance. The average past SFR is roughly the same for all models ending at $`\mu _1=0.1`$ and is $``$SFR$`=0.9`$ $`M_{}`$yr-1. In contrast, present-day SFRs range from SFR1 = 0.07 to 3 $`M_{}`$yr-1 and in fact determine the contribution by young stars to the integrated light of the model galaxy. Current oxygen abundances predicted are \[O/H\]$`{}_{1}{}^{}+0.25`$ and are mainly determined by $`\mu _1`$, the IMF, and the assumed mass limits for SNII (cf. Table 3). Column (7) gives the number of stars formed, Column (8) their total $`B`$ band luminosity. Column (10) shows the resulting Hi mass-to-light ratio. Column (11) gives the slope of the IMF used, while Column (11) refers to additional Notes.

Let us consider the photometric evolution of the constant and exponentially decreasing SFR models in some more detail. Fig. 3 shows the evolution of the total number of MS and post-MS stars for exponentially decreasing SFR model A1. The current total number of MS stars is roughly $`410^{10}`$, compared to $`10^8`$ post-MS stars (all phases) and $`310^4`$ for AGB stars only. The present-day mean luminosity of stars on the MS is $`10^1L_{}`$ compared to $`10^4L_{}`$ for AGB stars. The current bolometric galaxy luminosity is determined mainly by MS stars ($`L_{\mathrm{MS}}410^9L_{}`$). In particular, AGB stars ($`L_{\mathrm{AGB}}10^8L_{}`$) are relatively unimportant.

We show in Fig. 4 the $`B`$ and $`I`$-band magnitudes of stars in different evolutionary phases (again for the exponential model). As for the bolometric galaxy luminosity, MS stars generally dominate in the $`B`$-band. However, in the $`I`$-band, RGB and HB stars are nearly as important as MS stars, at least in later stages of galactic evolution. Due to the cooling of old, low-mass MS stars as well as the increasing contribution by RGB and HB stars with age, the current total $`I`$-band magnitude is considerably brighter than that in the B-band. This qualitative model behaviour is insensitive to the adopted star formation history (for e-folding times larger than $`35`$ Gyr) but instead is determined by the assumed IMF and the stellar input data used. Thus, constant star formation models exhibit a similar behaviour apart from being brighter by about one magnitude in all passbands at late times (cf. Fig. 4).

Fig. 5 illustrates the sensitivity of broadband colours to the star formation history for constant and exponentially decreasing SFR models. The colours considered here redden with galactic age. In general, differences between colours such as for different SFR models are less than the variations of these colours with age for a given model. Assuming a galactic age of e.g. 8 instead of 14 Gyr has limited effect on the resulting galaxy colours (e.g. less than 0.1 mag in $`BV`$), even though absolute magnitudes are substantially altered. Both age and extinction effects can result in substantial reddening of the colours of a stellar population in almost the same manner and it is difficult to disentangle their effects on the basis of photometry data alone. Clearly, galaxy colours alone are not well suited to discriminate between different SFR models, even when internal extinction is low and other reddening effects are negligible (see below).

## 5 Results

As our emphasis will be on a comparison between the evolution of HSB and LSB galaxies, we will first focus on the relatively well-known evolution of HSB galaxies, using a model chosen to emphasise the observed properties of HSB galaxies. Using this as a starting point, we explore other models to find the best model for LSB galaxies.

### 5.1 HSB galaxies: $`\mu _1=0.1`$ models

Models using exponentially decreasing SFRs and different e-folding times for different Hubble types (e.g. Larson & Tinsley 1978; Guiderdoni & Rocca-Volmerange 1987; Kennicutt 1989; Bruzual & Charlot 1993; Fritze-v. Alvensleben & Gerhard 1994) are usually found to be in good agreement with observations of HSB galaxies. Therefore we will mainly focus on the exponential SFR models discussed above. As the gas-richness is one of the obvious differences between HSB and LSB galaxies, we will first focus on models with $`\mu _1=0.1`$ to describe HSB galaxies.

Fig. 6 shows the results of the exponentially decreasing SFR model: SFR $`\mathrm{exp}(t/\tau _{\mathrm{sfr}})`$ with $`\tau _{\mathrm{sfr}}=5`$ Gyr ending at $`\mu _1=0.1`$ at $`t=t_{ev}=14`$ Gyr for several initial galaxy masses $`M_\mathrm{g}(t=0)10^810^{10}`$ $`M_{}`$. The precise value of $`\mu _1`$ was achieved by scaling the amplitude of the SFR accordingly. Galaxy colours and abundances are not affected by this scaling while absolute magnitudes and final gas masses scale with $`M_\mathrm{g}(t=0)`$ (as indicated in Fig. 6).

Present-day ($`BV`$) and ($`RI`$) colours for this model are 0.6 and 0.5, respectively. HSB galaxies with $`(BV)\begin{array}{c}>\\ \end{array}0.6`$ and $`(RI)\begin{array}{c}>\\ \end{array}0.5`$ can be explained only when substantial amounts of internal dust extinction are incorporated. The reason is that even though a single stellar population born with metallicity $`Z=0.02`$ may become as red as $`(BV)1.1`$ and $`(RI)0.8`$, the luminosity contribution by young stellar populations (with ages less than a few Gyr) results in substantial blueing of the galaxy colours. For the same reason, values of $`(BV)\begin{array}{c}>\\ \end{array}0.65`$ are not predicted by dust-free models independent of the adopted (exponential) SFR (as long as $`\tau \begin{array}{c}>\\ \end{array}4`$ Gyr) or IMF. Therefore, considerable reddening of the younger stellar populations is required to explain the colours of HSB galaxies in this model. Figs. 6c and d show that both the \[O/H\] abundance ratios and present-day gas masses of HSB galaxies can be explained reasonably well for $`M_\mathrm{g}(t=0)>10^{10}`$ $`M_{}`$. These conclusions agree with results from many previous models (with different stellar input data) for HSB galaxies (e.g. Larson & Tinsley 1978; Guiderdoni & Rocca-Volmerange 1987).

It is clear from Fig. 6 that this model is not able to explain the properties of the LSB galaxies in our sample. The model colours are too red while the predicted \[O/H\] abundances are too high. As LSB galaxies have higher gas-fractions we thus need to broaden the range of gas-fractions.

### 5.2 LSB galaxies: $`\mu _10.1`$ models

We now consider models with gas fractions $`\mu _1=`$ 0.025, 0.1, 0.3, 0.5, 0.7, and 0.9. First we will examine the constraints which individual properties such as abundance and gas-fraction impose on the models, before combining these in a final model.

#### 5.2.1 Colours and magnitudes

Results for a model with initial mass $`M_\mathrm{g}(t=0)=10^{10}`$ $`M_{}`$ are shown in Fig. 7. While the age distribution of the stellar populations is identical in all the models shown, present-day ($`BV`$) and ($`RI`$) colours are found to decrease by 0.2 and 0.1 mag, respectively, when going from models ending at $`\mu _1=0.025`$ to $`0.9`$. This bluing effect is due to the decrease of stellar metallicities for models ending at increasingly higher gas fractions.

The LSB galaxies can be distinguished in two major groups by means of their colours. First, LSB galaxies with $`(BV)\begin{array}{c}>\\ \end{array}0.5`$ which can best be modelled with exponentially decaying SFR models ending at values $`\mu _1=0.50.7`$ at age 14 Gyr. These results can be shifted towards brighter or fainter $`B`$ magnitudes by changing the initial gas masses accordingly. This leaves the resulting colours unaltered (cf. Fig. 6).

Second, relatively blue LSB galaxies with $`(BV)\begin{array}{c}<\\ \end{array}0.4`$ cannot be fitted by exponentially decreasing SFR models with $`\tau \begin{array}{c}<\\ \end{array}5`$ alone (assuming $`t_{\mathrm{ev}}=14`$ Gyr), regardless of their current gas fraction $`\mu _1`$. There are several possible explanations: a relatively young “surplus” stellar population may influence the galaxy colours resulting from an underlying exponentially decreasing SFR. Alternatively, these LSB galaxies may have started forming stars recently (e.g. $`58`$ Gyr instead of 14 Gyr, cf. Fig. 7a), or they may have more slowly decaying or constant SFRs which generally result in present-day colours of $`(BV)\begin{array}{c}<\\ \end{array}0.4`$ and $`(RI)\begin{array}{c}<\\ \end{array}0.35`$ (but the latter option appears to be excluded because of their abundances and gas fractions; see below).

#### 5.2.2 Abundances

The observed range in \[O/H\] abundances for LSB galaxies is well explained by exponentially decaying SFR models with $`\mu _1\begin{array}{c}>\\ \end{array}0.3`$ (cf. Fig. 7c). Constant SFR models ending at $`\mu _1\begin{array}{c}>\\ \end{array}0.3`$ are also consistent as the abundances of elements predominantly produced in massive stars are in general determined by the present-day gas fraction $`\mu _1`$ and are insensitive to the detailed underlying star formation history (e.g. Tinsley 1980). However, these latter models appear to be excluded by the measured gas masses (see 5.2.4 and 5.2.5) (for a closed box model). Though the abundances of metal-poor LSB galaxies with \[O/H\] $`\begin{array}{c}<\\ \end{array}1`$ can be well explained by models ending at $`\mu _1=0.9`$, it is likely that the exponential models presented here are an over-simplification for these gas-dominated systems. It is more likely that they have experienced a low and sporadic star formation rate.

#### 5.2.3 Gas content

Present-day gas masses observed in LSB galaxies can be well fitted by exponentially decaying SFR models ending at $`\mu _10.5`$ (Fig. 7d). This is consistent with the range of $`\mu _1\begin{array}{c}>\\ \end{array}0.3`$ derived from the \[O/H\] data. Exponentially decreasing SFR models ending at $`\mu _1\begin{array}{c}>\\ \end{array}0.6`$ are inconsistent with the observations unless we assume that LSB galaxies have started forming stars only a few Gyr ago.

Fig. 8 displays the gas mass vs $`(BV)`$ for constant and exponentially decaying SFR models. Exponentially decreasing SFR models are able to explain simultaneously values of $`\mu _10.5\pm 0.2`$, $`(BV)\begin{array}{c}>\\ \end{array}0.5`$, and $`M_\mathrm{g}\begin{array}{c}>\\ \end{array}10^9`$ $`M_{}`$, as observed for the majority of the LSB galaxies in our sample. Over the entire range of possible gas fractions, constant (or increasing) SFR models are only able to explain the bluest galaxies.

Fig. 8 demonstrates that exponentially decreasing SFR models ending at $`\mu _10.5`$ are in good agreement with the observed hydrogen mass-to-light ratios $`M_{\mathrm{HI}}/L_B`$, in contrast to constant (or slowly decreasing) SFR models (expect for the bluest systems).

The conclusion that constant and slowly decreasing SFR models are inconsistent with the observed photometric and chemical properties of LSB galaxies is based on the assumption of negligible amounts of dust extinction in these systems. However, if a considerable fraction of the LSB spirals in our sample would suffer from extinctions of $`E_{(BV)}=0.10.25`$, the $`M_{\mathrm{HI}}/L_B`$ ratios predicted by constant SFR models would increase by a factor $`1.42.5`$ after correction for internal extinction. In this manner, constant and slowly decreasing SFR models ending at gas fractions $`\mu _1=0.30.7`$ after $`14`$ Gyr could also explain relatively red LSB galaxies with $`(BV)\begin{array}{c}>\\ \end{array}0.5`$ and $`M_{\mathrm{HI}}/L_B`$ ratios as large as $`1.5`$. Even though we argued in Sect. 2 that internal extinction in LSB galaxies is unlikely to exceed $`E_{(BV)}0.1`$, knowing the dust content of LSB galaxies is obviously of crucial importance in modelling their evolution.

#### 5.2.4 Summary: the best model

To summarise, we find that exponentially decreasing SFR models ending at $`\mu _1=0.30.7`$ are in agreement with the colours, magnitudes, \[O/H\] abundances, gas contents, and mass-to-light ratios observed for LSB galaxies with $`(BV)\begin{array}{c}>\\ \end{array}0.45`$. Blue LSB galaxies with $`(BV)\begin{array}{c}<\\ \end{array}0.45`$ cannot be fitted by exponentially decreasing SFR models without an additional light contribution from a young stellar population. Alternatively, such LSB galaxies may have experienced constant SFRs, or may be much younger than 14 Gyr.

### 5.3 HSB/LSB: Just an extinction effect?

Since the models require an internal extinction in HSB galaxies up to $`E_{(BV)}0.5`$ (corresponding to $`A_V1.5`$ mag), independent of the SFH, one could argue that extinction may be the main explanation for the difference in colour observed between LSB and HSB galaxies.

As shown in this paper the models indicate that the underlying stellar populations in LSB and HSB galaxies are still distinctly different due to differences in the chemical evolution of these systems. Therefore, extinction effects, although important, cannot be the entire explanation for the colour differences observed.

## 6 The effects of small star formation bursts

### 6.1 Luminosity contribution by HII regions in LSB galaxies

The presence of Hii regions in all the LSB galaxies in our sample suggests that recent star formation is a common phenomenon in these systems, and may thus influence the observed properties. In the previous section we found that for some LSB galaxies a simple exponential SFR was unable to explain the observed properties and that an additional “burst” of young stars was needed. Here we examine the effects of such bursts on the colours of LSB galaxies. To this end, we selected all Hii regions that could be identified by eye, either from H$`\alpha `$ or $`R`$-band CCD images, and added their total luminosity in the $`I`$-band. We restrict ourselves to the $`I`$-band data as to be least affected by extinction that may be present in or near Hii regions (McGaugh 1994). We define $`\eta _I`$ as the ratio of this Hii region integrated luminosity and the total luminosity of the corresponding LSB galaxy. As we are probably incomplete at low Hii region luminosities, this ratio gives a lower limit for the Hii region contribution to the total luminosity.

In columns (1) and (2) of Table 4, we list the LSB galaxy identification and number of Hii regions selected. The number of Hii regions identified within individual LSB galaxy ranges from a few to $`25`$. For the ensemble of Hii regions in each LSB galaxy, we tabulate the absolute $`I`$ magnitudes as well as the corresponding ratios $`\eta _I`$ of the Hii region integrated luminosity and total LSB galaxy luminosity, in columns (3) and (4). Mean ($`BV`$) colours for the Hii regions and for the LSB galaxy as a whole are given in columns (5) and (6), respectively.

For most of the LSB galaxies in our sample, the contribution of the Hii regions to the total light emitted by LSB galaxies does not exceed $`\eta _I=0.050.1`$. However, as Hii regions are likely to contain an increased amount of dust (McGaugh 1994) the actual contributions may be higher by a factor of $`2`$ in $`I`$. Thus, the values of $`\eta _I`$ provide lower limits to the actual luminosity contributions of the Hii regions. For some LSB galaxies, e.g. F568-V1 and F577-V1, the Hii region contribution is found to be as high as $`\eta _I=0.2`$. These systems contain a only modest number of Hii regions so that their Hii regions on average may be larger and/or brighter than those present in several other LSB galaxies.

Figure 10 shows the resulting Hii region contributions $`\eta _I`$ for the SFR models discussed in Sect. 4. If we assume a maximum age $`\tau _{\mathrm{HII}}=5`$ Myr for the Hii regions observed, then this implies that stars more massive than $`25`$ $`M_{}`$ are associated with these regions, according to the stellar evolution tracks from the Geneva group (see below).

Corrections for dust extinction within the Hii regions and LSB galaxy as a whole, respectively, will shift the observations in the directions as indicated in Fig. 10 (assuming a mean Galactic extinction curve). From the $`I`$ band observations we can conclude that the values of $`\eta _I`$ predicted by smoothly varying SFR models are systematically too low for most of the LSB galaxies in our sample.

This is true in particular for F568-V1, F577-V1, and U628 for which values of $`\eta _{I,R}\begin{array}{c}>\\ \end{array}0.20.25`$ suggest that star formation has been recently enhanced by factors $`510`$ relative to the SFRs predicted by exponentially decreasing or constant SFR models.

### 6.2 Effects of small amplitude star formation bursts

We investigate whether the observed values of $`\eta _I0.25`$ in the $`I`$ band, as observed for several LSB galaxies discussed above, can be explained by small-amplitude bursts of star formation.

We assume a Gaussian star formation burst profile with a given amplitude $`A_b`$ and dispersion $`\sigma `$. We follow its evolution during 1 Gyr with a time resolution of $`0.1`$ Myr at time of burst maximum and of $`2`$ Myr at roughly 5$`\sigma `$ from burst maximum. We superimpose the star formation burst on an exponential SFR model, as discussed in Sect. 5.1.

We initially assume a galactic evolution time at burst maximum of $`t_\mathrm{b}=13`$ Gyr, burst maximum amplitude $`A_\mathrm{b}=8`$ $`M_{}`$ yr-1, burst duration $`\mathrm{\Delta }t_\mathrm{b}=5`$ Myr, maximum Hii region lifetime $`\tau _{\mathrm{HII}}=5`$ Myr, and an initial galaxy mass of $`10^{10}`$ $`M_{}`$. For simplicity, we neglect any influence of the burst on the chemical evolution of the model galaxy.

We show the result in Fig. 11a. At $`t=t_b=13`$ Gyr the contribution by young stars increases rapidly. Simultaneously, the $`(RI)`$ colours become significantly bluer. After burst maximum, the contribution by young stars decreases again and galaxy colours start to redden until the effect of the burst becomes negligible and colours and magnitudes evolve as prior to the burst.

In this manner, a characteristic burst loop is completed as shown in Fig. 11a. The shape of this loop is determined by: i) the burst amplitude, ii) the extinction within the Hii regions, iii) the contribution by the old stellar population to the integrated galaxy light, iv) the duration (and profile) of the burst, v) the maximum time during which young stars produced by the Hii regions can still be distinguished from the surrounding field population.

Burst amplitude and IMF: Fig. 11a demonstrates that values of $`\eta _I0.25`$ can be well explained by bursts with amplitude $`A_\mathrm{b}=8`$ $`M_{}`$yr-1 superimposed on an exponentially decreasing SFR model ending at $`\mu _1=0.1`$. Decreasing the burst amplitude by factors 2.5 and 5, respectively, results in the smaller loops shown in Fig. 11a. The actual burst amplitude required to explain the observations depends on many quantities as described below

Dust extinction: Fig. 11b illustrates the effect of dust extinction on $`\eta _I`$ and $`(RI)`$. A selective extinction of $`E_{BV}=(0.25,0.5,1)`$ mag for the Hii regions results in a reduction $`\eta _I`$ by a factor (1.6, 2.5, 6.5) and a reddening $`E_{RI}=`$ (0.14, 0.27, 0.56) mag, assuming a Galactic extinction curve. For values of $`E_{BV}\begin{array}{c}>\\ \end{array}0.5`$ mag, we find that the bluing effect on $`(RI)`$ by young massive stars formed during the burst is neutralised almost entirely by extinction. For intense bursts reddening of the galaxy colours may even occur. If variations in the mean extinction of the ensembles of Hii regions among LSB galaxies are small (e.g. less than a factor two), it is difficult to see how extinction alone can provide an adequate explanation for the large variations observed in $`\eta _I`$.

Global star formation history: From Fig. 11c it is clear that the burst effect on the galaxy magnitudes and colours increases when the contribution by the old stellar population is decreased. Thus, colours and magnitudes are less affected by bursts imposed on constant or even increasing SFRs compared to those imposed on exponentially decreasing SFR models: the smaller loop sizes just reflect that the mean age of the underlying stellar population is relatively low.

Current gas fraction: Fig. 11d demonstrates how the present-day gas fraction $`\mu _1`$ affects the effect of the burst. The luminosity contribution by the old stellar population decreases for increasing values of $`\mu _1`$, and thus, as discussed above, we find that the effect of a star formation burst for galaxies with $`\mu _1=0.9`$ (i.e. unevolved systems) is as large as that of a ten times stronger burst for $`\mu _1=0.1`$ (i.e. highly evolved systems). Thus, the burst amplitude required to explain the observations strongly depends on the present-day gas-to-total mass-ratio.

Burst duration: Fig. 11e shows that the duration of the burst affects the impact of the burst as well. For $`\mathrm{\Delta }t_\mathrm{b}=1`$, 5, and 10 Myr, the variation in $`\eta _I`$ is $`0.25`$ while the resulting galaxy $`(RI)`$ colours become bluer. Burst durations in excess of $`\mathrm{\Delta }t_\mathrm{b}5`$ Myr are unlikely since this would require dust extinctions $`E_{(BV)}\begin{array}{c}>\\ \end{array}1`$ mag in order to provide agreement with the observed $`(RI)`$ colours (cf. Fig. 11b). Such large extinction in Hii regions are probably excluded by the observations (e.g. McGaugh 1994). Thus, relatively narrow burst profiles are needed to explain extreme values $`\eta _I0.2`$ (as in F568-V1; cf. Table 4).

Maximum age of Hii regions: Fig. 11f shows that when the maximum lifetime of the Hii regions is increased from $`\tau _{\mathrm{HII}}=5`$ to 50 Myr, partial agreement with the observations can be achieved without invoking star formation bursts. We note that the resulting Hii region contributions do not increase linearly with $`\tau _{\mathrm{HII}}`$ as short lived massive stars dominate the luminosity contribution of all stars formed during the past $`\tau _{\mathrm{HII}}`$ yr.

Values of $`\tau _{\mathrm{HII}}\begin{array}{c}>\\ \end{array}50`$ Myr would mean that stars down to masses of $`m_{\mathrm{ion}}\begin{array}{c}<\\ \end{array}7`$ $`M_{}`$ would contribute to the Hii regions identified (e.g. Schaerer et al. 1993; $`Z=0.001`$). Observational estimates for $`m_{\mathrm{ion}}`$ are usually in the range $`1015`$ $`M_{}`$ (Wilcots 1994; García-Vargas 1995) and correspond to $`\tau _{\mathrm{HII}}\begin{array}{c}>\\ \end{array}15`$ Myr. Even though these values would imply that our adopted value of $`\tau _{\mathrm{HII}}=5`$ Myr is too low, this excludes extreme values of $`\tau _{\mathrm{HII}}=200`$ Myr which would be required to explain the observed range in $`\eta _I`$ exclusively in terms of variations in $`\tau _{\mathrm{HII}}`$ and/or $`m_{\mathrm{ion}}`$. Therefore, variations in $`\tau _{\mathrm{HII}}`$ (or equivalently $`m_{\mathrm{ion}}`$) and/or extinction may provide an explanation only for part of the variations in the Hii region integrated luminosity contributions observed among LSB galaxies.

In conclusion, the observations suggest that small amplitude, short bursts of star formation are important in at least several of the LSB galaxies for which accurate photometry data is available. Such recent episodes of enhanced star formation may play an important role in affecting the colours of the blue LSB galaxies.

### 6.3 Quantitative effect of small amplitude bursts on galaxy colours and magnitudes

Table 5 lists the effect of a 5 Myr star formation burst on the galaxy integrated magnitudes and colours for various burst amplitudes. For an exponentially decreasing SFR (model A1 in Table 3; assuming $`M_\mathrm{g}(0)=10^{10}`$ $`M_{}`$, $`\mu _1=0.1`$, Salpeter IMF), we find that a burst with amplitude $`A_\mathrm{b}=0.8`$ $`M_{}`$ yr-1 results in maximum colour variations $`\mathrm{\Delta }(BV)`$ and $`\mathrm{\Delta }(RI)`$ of $`0.18`$ and $`0.04`$ mag, respectively. The effect of increasing the burst amplitude by a factor 10 to $`A_\mathrm{b}=8`$ $`M_{}`$ yr-1 results in corresponding shifts of $`0.56`$ and $`0.26`$ mag, respectively. This effect is similar to that when the initial galaxy mass is reduced by a factor ten while leaving the burst amplitude unaltered (i.e. for. $`M_\mathrm{g}(0)=10^9`$ $`M_{}`$ and $`A_\mathrm{b}=0.8`$ $`M_{}`$ yr-1).

For bursts superimposed on exponentially decreasing SFRs models ending at $`\mu _1=0.1`$, the colour and magnitude shifts predicted are consistent with the observations in case of burst amplitudes $`A_\mathrm{b}\begin{array}{c}<\\ \end{array}3`$ $`M_{}`$ yr-1, assuming a typical extinction of $`E_{(BV)}=0.3`$ mag in the Hii regions. In fact, the effect of the burst is determined mainly by the total luminosity of the young stellar populations formed according to the continuous SFR during the last Gyr or so. Since the amplitude of the SFR scales with $`(1\mu _1)`$ for models ending at different gas fractions $`\mu _1`$ the impact of the burst for models ending at $`\mu _1=0.5`$ is about twice that given in Table 5. Similarly, for models ending at $`\mu _1\begin{array}{c}>\\ \end{array}0.9`$, the burst effect becomes roughly ten times stronger compared to the $`\mu _1=0.1`$ case (cf. Fig. 11). The effect of the burst is substantially reduced when going from exponentially decreasing to constant SFRs (cf. Table 5).

## 7 Present-day star formation rates in LSB galaxies

### 7.1 Theoretical star formation rates

#### 7.1.1 Continuous SFR

Theoretical SFRs can be derived from the models discussed in the previous section (exponentially decreasing SFR, Salpeter IMF), using:

$$\mathrm{SFR}^{\mathrm{cont}}A(\mu _1)\frac{M^{\mathrm{tot}}}{10^{10}}M_{}\mathrm{yr}^1$$
(3)
where $`A(\mu _1)`$ is the model SFR amplitude required to end at a gas fraction $`\mu _1`$ at a galactic evolution time of 14 Gyr (assuming an initial mass of $`10^{10}`$ $`M_{}`$) and $`M^{\mathrm{tot}}`$ the total galaxy mass as obtained from $`M_{\mathrm{HI}}`$ and $`\mu _1`$.

We list the required values for $`A`$ to have the models end at certain values of $`\mu _1`$ in Table 6. Using these values we find that LSB galaxies show present-day SFRs (without bursts) between $`0.01\begin{array}{c}<\\ \end{array}\mathrm{SFR}^{\mathrm{cont}}\begin{array}{c}<\\ \end{array}0.15`$$`M_{}`$yr-1. For a typical LSB galaxy (i.e. $`M^{\mathrm{tot}}=10^{10}`$ $`M_{}`$, $`\mu _1=0.5`$) we estimate SFR$`{}_{}{}^{\mathrm{cont}}=0.1`$ $`M_{}`$yr-1.

#### 7.1.2 Burst SFR

As we discussed in Sect. 6, the effect of a starburst on the galaxy integrated magnitudes and colours depends on many quantities. However, assuming a modest Hii region extinction of $`E_{(BV)}=0.25`$ mag and lifetime $`\tau _{\mathrm{HII}}=20`$ Myr, crude estimates for the maximum burst amplitude can be derived from:

$$\mathrm{SFR}^{\mathrm{burst}}20\eta _\mathrm{I}\frac{M_{\mathrm{HI}}}{10^{10}}\left(\frac{1}{\mu _{\mathrm{rot}}}1\right)M_{}\mathrm{yr}^1$$
(4)
We find that $`I`$-band contributions of $`\eta _\mathrm{I}=0.2`$ are best explained by a 5 Myr burst with amplitude SFR$`{}_{}{}^{\mathrm{burst}}=0.8`$ $`M_{}`$yr-1, assuming present-day values of $`M_{\mathrm{HI}}=210^9`$ $`M_{}`$ and $`\mu _{\mathrm{rot}}=0.5`$.

Theoretical estimates for the total current SFRs in LSB galaxies are found from $`\mathrm{SFR}^{\mathrm{tot}}=\mathrm{SFR}^{\mathrm{cont}}+\mathrm{SFR}^{\mathrm{burst}}`$ and are listed in the last two columns of Table 7. Present-day SFRs for LSB galaxies experiencing small amplitude bursts range from SFR$`{}_{}{}^{\mathrm{tot}}0.02`$ to 0.8 $`M_{}`$ yr-1.

### 7.2 Empirical star formation rates

We have also derived current SFRs for LSB galaxies using the empirical method presented by Ryder & Dopita (1994; hereafter RD) based on CCD surface photometry of galactic disks. These authors found a relationship between the local H$`\alpha `$ and $`I`$-band surface brightness in the disks of a sample of 34 of southern spiral galaxies. From this relation, RD derived a constraint on the present-day SFR integrated over the entire stellar mass spectrum as:
$$\mathrm{log}\mathrm{SFR}_1=0.26\mu _I+0.92\mathrm{log}\sigma _{\mathrm{HI}}+5.3$$

$$M_{}\mathrm{pc}^2\mathrm{Gyr}^1$$
(5)   where $`\mu _I`$ is the $`I`$-band surface brightness and $`\sigma _{\mathrm{HI}}`$ is the global mean Hi surface density \[$`M_{}`$pc-2\] within the star-forming disk. The relation between SFR1 and $`\mu _I`$ is normalised by a term related to the mean surface density $`\sigma _{\mathrm{HI}}`$ and by a constant which is partly related to the conversion of the massive star formation rate to total SFR depending on the adopted IMF (cf. Kennicutt 1983). It is unclear from the RD sample whether the relation is valid also for the lowest (stellar) $`I`$-band surface densities that are observed among LSB galaxies. However, since this relation holds over a wide range in surface brightness and massive star formation in the disks of spirals it appears rather insensitive to galactic dynamics, extinction, and molecular gas content (RD), we expect this relation to be valid also in case of LSB galaxies. At the faintest surface brightnesses (i.e. $`\mu _I\begin{array}{c}>\\ \end{array}25.6`$ mag arcsec-2), the relation may be flattening off although the effects of sky subtraction and small number statistics leave this open to question. If flattening indeed occurs, Eq. (5) provides lower limits to the actual SFRs in LSB galaxies.

Using Eq. (3), we estimate global present-day SFRs for all LSB galaxies in our sample with measured $`I`$-band magnitudes and related data. For these LSB galaxies we list the distance, apparent $`I`$-band magnitude, and radius $`R_{27}`$ of the 27 mag arcsec-2 $`B`$-band isophote, in columns (2) to (4) in Table 7. This radius corresponds to the optical edge of the LSB galaxy and is more representative for the radius within which the old disk stellar population in LSB galaxies is contained than is $`R_{25}`$ as used by RD for HSB galaxies. Accordingly, we define an effective I band surface brightness as:

$$\mu _I^{\mathrm{eff}}=m_I+2.5\mathrm{log}(\pi R_{27}^2)$$
(6)
and use this in Eq. (5). We tabulate the outermost radius of the measured Hi rotation curve $`R_{\mathrm{HI}}`$, effective $`I`$ band surface brightness $`\mu _I^{\mathrm{eff}}`$, total Hi mass derived within $`R_{\mathrm{HI}}`$, and the mean global surface Hi density $`\sigma _{\mathrm{HI}}`$ in columns (5) to (8), respectively. For a consistent comparison between effective surface brightness and global Hi surface density, one ought to measure them out to the same radius (e.g. $`R_{27}`$). However, since $`M_{\mathrm{HI}}`$ has been measured within $`R_{\mathrm{HI}}`$, we expect the former values to be more representative of the average Hi surface density in the star forming part of the disk. The mean global Hi surface densities $`\sigma _{\mathrm{HI}}`$ derived using $`R_{\mathrm{HI}}`$ vary between 2 and 5.5 $`M_{}`$pc-2 and are substantially smaller (i.e. by $`2060`$ %) than those derived using $`R_{27}`$ instead.

The empirically derived mean present-day SFRs for the LSB galaxies from our sample range from about SFR$`{}_{1}{}^{}=0.02`$ to $`0.2`$ $`M_{}`$yr-1 (cf. column 10 of Table 7). Errors arising from the Hi normalisation are estimated to be within a factor of two. This is illustrated when the same SFRs are derived assuming a fixed Hi surface density of 2 $`M_{}`$pc-2 for all LSB galaxies (cf. SFR$`{}_{}{}^{\mathrm{fix}}{}_{1}{}^{}`$ in Table 7).

The empirically derived current SFRs in LSB galaxies are in good agreement with the theoretically derived SFRs ranging from SFR$`{}_{}{}^{\mathrm{cont}}0.01`$ to 0.15 $`M_{}`$yr-1 (cf. Table 7). As discussed before, the theoretically derived present-day SFRs of individual LSB galaxies lie probably between SFRcont and SFRtot where the latter values include the contribution of small amplitude bursts (SFR$`{}_{}{}^{\mathrm{tot}}=`$SFR$`{}_{}{}^{\mathrm{cont}}+`$SFRburst). In several cases, the values of SFRtot are considerably larger than the empirical values which suggests that the burst contribution is overestimated by the models and/or that the SFRs derived empirically do not trace well local enhancements of star formation at the faint surface brightnesses of LSB galaxies.

The present-day SFRs in LSB galaxies are thus found to be considerably below the $`510`$ $`M_{}`$yr-1 derived for their HSB counterparts (e.g. Kennicutt 1992 and references therein) but significantly larger than the $`0.001`$ $`M_{}`$yr-1 observed typically in dwarf irregular galaxies (e.g. Hunter & Gallagher 1986).

## 8 Discussion

In the previous sections we have focussed on the difference between HSB galaxies and the type of LSB galaxies we have in our sample. As noted in the introduction (and also shown in Fig. 1) HSB and LSB galaxies, however, do not form separate groups. In fact there is a broad range of galaxy properties and the purpose of the discussion is to clarify what role the star formation history plays in determining the final character of a galaxy.

In general, there appears a trend along the Hubble sequence from rapidly decaying SFRs for early type galaxies to constant or even increasing SFRs for dwarf irregular galaxies (see e.g. reviews by Sandage 1986; Kennicutt 1992). On average, the observed trend corresponds to a decrease of the ratio of mean past to present SFR along the Hubble sequence. Most of the LSB galaxies in our sample belong to the group of late-type galaxies for which exponentially decreasing SFR models are in good agreement with the observations. The remaining LSB galaxies, for which slowly decreasing or constant (sporadic) SFR models are more appropriate, belong to a group of galaxies with properties intermediate to those of disk galaxies with weak or absent spiral arms and Sm/Im galaxies. Thus, in general LSB galaxies comply well with the observed trend of SFR variation with Hubble type.

The “burst” scenario discussed in section 6 indicates that current star formation in virtually all the LSB galaxies in our sample is local both in time and space and suggests that sporadic star formation has been a continuous process from the time star formation started in the disks of LSB galaxies.

The low star formation rates of $``$0.1 $`M_{}`$yr-1 derived for LSB galaxies as well as the local nature of the star formation in these systems are consistent with the idea of a critical threshold for the onset of global star formation in disk galaxies (e.g. Skillman 1987; Kennicutt 1989; Davies 1990).

Even though LSB galaxies contain large amounts of gas, only very limited amounts participate in the process of star formation. If we assume that LSB galaxies maintain their current star formation rate of $``$0.1 $`M_{}`$yr-1, their typical present-day amount of gas $``$ Mg = 2.5 109 $`M_{}`$ will be consumed within $`\tau _{\mathrm{gas}}`$30 Gyr (for a recycled fraction of 25%). Such gas consumption times for LSB galaxies are much larger than a Hubble time (e.g. Romanishin 1980). For comparison, $`\tau _{\mathrm{gas}}24`$ Gyr in HSB galaxies, assuming typically M$`{}_{\mathrm{g}}{}^{}10^{10}`$ $`M_{}`$ and SFR$`{}_{1}{}^{}5`$ $`M_{}`$yr-1, which implies that HSB galaxies will run out of gas soon (see Kennicutt 1992).

The presence of an old stellar population in many late-type LSB galaxies as indicated by their optical colours (e.g. vdH93; dB95) and as confirmed by the results from the modelling suggests that LSB galaxies roughly follow the same evolutionary history as HSB galaxies, but at a much lower rate.

The models suggest that very slowly decreasing star formation rates with $`e`$-folding times much larger than $``$ 5 Gyr probably can be ruled out for the majority of the LSB galaxies examined here. The observed colours are too blue, provided extinction is indeed negligible. This result is insensitive to the possible occurrence of infall or accretion of gas. Similarly the models indicate that the dominant stellar population in galaxies with $`(BV)0.5`$ mag cannot be as young as 10 or 5 Gyr (cf. Fig. 5). In contrast the models can not rule out a dominant luminosity contribution by stellar populations significantly younger than 5 – 10 Gyr for LSB galaxies with $`(BV)0.4`$.

First this indicates that the mean age of the stellar population in most LSB and HSB galaxies is similar even though the disks of LSB galaxies are in a relative early evolutionary stage. Although we have not explored an entire range of star formation e-folding times, values much below or in excess of $``$ 5 Gyr are not very likely. Smaller values would increase the colour contribution from the underlying older stellar population and hence produce redder colours than observed, while much larger values correspond to slowly decreasing or almost constant star formation rates implying a relatively larger contribution of the younger stellar population.

Secondly, the combined effect of extinction and metallicity on galaxy colours is sufficient to explain the colour differences observed between LSB galaxies and HSB galaxies. Since the amount of extinction depends strongly on the dust content, which in turn is coupled to the heavy element abundances in the ISM (see Sect. 2), metallicity probably is the main quantity determining the colour differences observed between LSB and HSB galaxies. In this manner the much lower rate of star formation in LSB galaxies, implying lower metallicities and dust contents, indirectly determines the blue colours of LSB galaxies compared to HSB galaxies.

## 9 Summary

We have examined the star formation histories of LSB galaxies using models which take into account both the photometric and the chemical evolution of the galaxies. For the majority of the LSB galaxies in our sample, observed $`UBVRI`$ magnitudes, \[O/H\] abundances, gas masses and fractions, and Hi mass-to-light ratios are best explained by an exponentially decreasing global SFR ending at a present-day gas-to-total mass-ratio of $`\mu _10.5`$. When small amplitude bursts are involved to decrease the predicted $`M_\mathrm{g}/L`$ ratios, models ending at $`\mu _10.7`$ may also apply. In addition to exponentially decreasing SFR models, $`15\%`$ of the LSB galaxies require modest amounts of internal extinction $`E_{(BV)}\begin{array}{c}<\\ \end{array}0.1`$ mag to explain the relatively red colours of $`(BV)0.6`$ of these systems.

A substantial fraction ($`35\%`$) of the LSB galaxies in our sample have colours $`(BV)\begin{array}{c}<\\ \end{array}0.45`$ mag and exhibit properties similar to those resulting from exponentially decreasing SFR models at evolution times of $`510`$ Gyr. Alternatively, recent episodes of enhanced star formation superimposed on exponentially decreasing SFR models may provide an adequate explanation for the colours of these systems (see Sect. 6). Recent star formation is observed, at least at low levels, in essentially all the LSB galaxies in our sample. Hence, it seems justified to assume that the disks in LSB galaxies experienced continuous (i.e. frequent small amplitude bursts of) star formation, at least during the last few Gyr.

There is nothing peculiar about the evolution of LSB galaxies. Broadly speaking their evolution proceeds like that of HSB galaxies, but at a much lower rate.

###### Acknowledgements.

We thank Andre Maeder and Georges Meyanet for providing us with the stellar isochrone data and conversion programs. We thank Roelof de Jong for making available to us the data of a large sample of face-on spirals. We are grateful to the referee Uta Fritze-von Alvensleben for constructive comments from which this paper has benefitted.


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# Structure Factor of a Lamellar Smectic Phase with Inclusions

## Abstract

Motivated by numerous X-ray scattering studies of lamellar phases with membrane proteins, amphiphilic peptides, polymers, or other inclusions, we have determined the modifications of the classical Caillé law for a smectic phase as a function of the nature and concentration of inclusions added to it. Besides a fundamental interest on the behavior of fluctuating systems with inclusions, a precise characterization of the action of a given protein on a lipid membrane (anchoring, swelling, stiffening …) is of direct biological interest and could be probed by way of X-ray measurements. As a first step we consider three different couplings involving local pinching (or swelling), stiffening or tilt of the membrane. In the first two cases we predict that independent inclusions induce a simple renormalization of the bending and compression modulii of the smectic phase. The X-ray experiments may also be used to probe correlations between inclusions. Finally we show that asymmetric coupling (such as a local tilt of the membrane) results in a modification of the usual Caillé law.

Lyotropic smectic phases ($`L_\alpha `$ phases) are liquid crystalline systems of parallel, regularly spaced lipid bilayers. Those phases form a quasi-crystalline structure in one (the $`z`$) direction, while retaining their fluid properties in the two others directions ($`xy`$ plane). They have been studied extensively in recent years, as they form one of the most convenient lyotropic structure for the experimental study of fluctuating interfaces. They have been for instance particularly useful in demonstrating the importance of the so-called entropic (Helfrich) repulsion between fluctuating membrane. Furthermore, the degree of dilution (the layer spacing) and other controlled parameters of the lyotropic $`L_\alpha `$ phase can be experimentally varied over a wide range, which allows for a precise study of the scattering intensity of the phase. This scattering intensity is expected to show several peaks corresponding to a stack of regularly spaced membrane. In these layered systems however, the thermal fluctuations destroy the long range order at finite temperature, leading to weak - power law - singularities of the peaks.

Inclusions in membrane have been actively studied for many years. Most of the earlier work was done by the biological community, for which it was a necessary step toward the comprehension of the complexity of biological (cell) membrane. In the last twenty years or so, a more physical description of membrane-inclusion complexes, in term of curvature energy and disruption of the molecular order of the membrane, have led to interesting results on membrane-induced interactions between inclusions. While this “physical” picture of the membrane-inclusion interactions leaves aside many fundamental processes driven by specific interactions, it allows for a description of universal features of the complexes in terms of elastic couplings between membrane and inclusion. This description is applicable to a certain extent to membrane with adsorbed or grafted polymers or colloids, as well as to transmembrane inclusions resembling certain membrane proteins.

In recent years, numerous experimental studies have investigated the role of added membrane proteins in lyotropic smectic phase. This lipid phase seems to be of particular interest for the study “in vitro” of the bilayer-protein association, and may have some practical application for gene therapy. From a physicist point of view, it is interesting to understand the effect of inclusions on the thermal fluctuations of flexible, fluid membranes. From a more “biological” point of view, a precise characterization of the physical action of a given protein onto a lipid membrane (anchoring, swelling, stiffening …) is of direct biological interest, and could be reach via X-ray measurements. This work have some relevance in material science as well, since for practical purposes, a lamellar phase is very rarely composed of two components (solvent+surfactants) only. The adjunction of a third or fourth component, such as cosurfactant, polymers, proteins or colloids is of common practice to tune the properties of the phase. The object of the present work is to determined the modifications of the smectic X-ray scattering due to the presence of inclusions in the lamellar phase. Ultimately, one could expect to map out different membrane-inclusion couplings by their effect on the X-ray structure factor. We consider three model inclusions, leading to different modifications of the membrane elastic properties, and we show that the qualitative analysis of the diffraction spectrum is enough to discriminate between these inclusions.

Several recent theoretical works deal with lamellar systems with flexible polymers or other soluble inclusions (so-called doped solvent lamellar phases). These works are complementary, as we are interested in insoluble inclusions such as the membrane proteins used in , while  studied effects such as polymer depletion - induced interactions between lamellae, resulting from an equilibrium between the trapped inclusions and free inclusions in pure solvent.

$`\mathrm{}`$ The standard description of Caillé for pure $`L_\alpha `$ phases has been confirmed experimentally in numerous systems. It is based on the classical smectic hamiltonian:

$$H_0=\frac{1}{2}d^3r\left(B\left(_zu\right)^2+K\left(_{}^2u\right)^2\right)$$
(1)
where $`u(𝐫)`$ is the (continuous) normal displacement of the layer at the point $`𝐫`$, $`z`$ is the coordinate normal to the layers, and $`_{}`$ is the gradient along the layers. The elasticity of the smectic phase is characterized by the two coefficients $`K`$ and $`B`$, which are related to the energy cost of bending and compressing the sample. Typically, $`K=\kappa /d`$, where $`\kappa `$ is the bending modulus of a single membrane and $`d`$ is the average layer spacing, and $`B`$ is function of the intermembrane interactions. Those modulii define the characteristic smectic length $`\lambda =\sqrt{K/B}`$. The scattering intensity $`I(𝐪)`$ is related to the Fourier transform of the density correlation function $`G_n(𝐫)`$, itself related to the exponential of the layer displacement correlation function $`g(𝐫)=q_0^2/2\left(u(𝐫)u(0)\right)^2`$, where $`\mathrm{}`$ represents a thermal average of the fluctuations, and $`q_0=2\pi /d`$ (see , sec 6.3.2). As usual, the Fourier decomposition of the layer displacement is used: $`u(𝐫)=d^3q/(2\pi )^3u_qe^{i\mathrm{𝐪𝐫}}`$. Using Eq.(1):

$$g(𝐫)=\frac{q_0^2}{V}\frac{d^3q}{(2\pi )^3}|u_q^2|\left(1e^{i\mathrm{𝐪𝐫}}\right)\mathrm{with}|u_q^2|_0=\frac{Vk_BT}{Bq_z^2+Kq_{}^4}$$
(2)
where $`V`$ is the volume of the sample. After integration over $`q_z`$, an asymptotic solution of $`g(𝐫)`$ can be found:

$`g^{(0)}(𝐫)={\displaystyle \frac{q_o^2k_BT}{4\pi \sqrt{KB}}}{\displaystyle 𝑑q_{}\frac{\left(1J_0(q_{}r_{})e^{\lambda zq_{}^2}\right)}{q_{}}}\eta \left(2\gamma +2\mathrm{log}{\displaystyle \frac{\pi r_{}}{l}}+E_1\left({\displaystyle \frac{r_{}^2}{4\lambda z}}\right)\right)`$ (3)
$`\mathrm{with}\eta {\displaystyle \frac{q_0^2k_BT}{8\pi \sqrt{KB}}}`$ (4)
where $`J_0(x)`$ is the Bessel function of the first kind, $`\gamma `$ is the Euler constant, and $`E_1(x)`$ is the exponential integral ($`l`$ is a molecular size). From Eq.(4), one can show that the scattering intensity $`I_n(𝐪)`$ near the nth peak, defined by

$$I_n(𝐪)=G_n(𝐪+n𝐪_0)+G_n(𝐪n𝐪_0)\mathrm{with}G_n(𝐫)e^{n^2g(𝐫)}$$
(5)
shows power law singularities of the form

$$G_n(𝐪n𝐪_0)\{\begin{array}{cc}(q_znq_0)^{2+\eta }\mathrm{if}q_{}=0\hfill & \\ q_{}^{4+2\eta }\mathrm{if}q_z=0\hfill & \end{array}$$
(6)
The above law gives a satisfactory description of the X-ray structure factor in the vicinity of the diffraction peaks, and has been observed experimentally (see  and references therein). A more accurate calculation of $`I(𝐪)`$ over a broad range of wave vector requires a discrete description of the lamellar phase, and the account of the fluctuations of the lipid concentration, and is not tractable analytically. As one can see, this peculiar power law divergence allows for an experimental determination of the product $`KB`$, through the parameter $`\eta `$. Low values of the parameter $`\eta `$, such as those observed for electrostatically stabilized systems ($`0.2`$) correspond to a well defined smectic organization with little fluctuations. Larger values of order unity are observed for weakly interacting neutral or screened systems stabilized by the Helfrich interaction.

$`\mathrm{}`$ Several couplings between the inclusions and the membrane are considered below. They are referred to as “pinch”, “stiff”, and “tilt”, and are schematically depicted in Fig.1 and Fig.2. The “pinch” inclusion can be thought of as exerting a force which tends to pinch ($`\beta _{pinch}>0`$, see Eq.(7) below) or swell ($`\beta _{pinch}<0`$) neighboring membranes. Since we are using a continuous description for the lamellar phase, the inclusion may reside within a membrane or between two membranes. Inclusions which would typically induce such deformations are amphiphilic proteins, with hydrophobic parts which penetrate in the bilayers and hydrophilic parts lying in the water, or simply particles larger than the layer spacing which sterically swell the membrane. A pinched lamellar structure has also been identified in the case of anionic polymers added to a lamellar phase formed by a mixture of cationic and neutral lipids. The hamiltonian corresponding to this perturbation couples the density of inclusion $`\rho (𝐫)`$ with the variation of the layer spacing:

$$\mathrm{\Delta }H_{pinch}=d^3r\beta _{pinch}\rho (𝐫)_zu$$
(7)
The “stiff” inclusion corresponds to a transmembrane protein which locally stiffens (or softens) the membrane and modifies its fluctuations. A stiffening is expected for many integral membrane proteins, or for rigid inclusions laying on the lipid bilayer. On the other hand, pores in the membrane are expected to induce a softening of the membrane (among other, less trivial phenomena). The inclusion density is coupled with a variation ($`\delta K`$) of the bending modulus:

$$\mathrm{\Delta }H_{stiff}=\frac{1}{2}d^3r\delta Kv\rho (𝐫)\left(_{}^2u\right)^2$$
(8)
where $`v`$ is the effective volume of an inclusion in the membrane.

The “tilt” inclusion corresponds to an anisotropic inclusion (such as a grafted polymer) which would locally induce a curvature to a membrane without changing the layer spacing. It has been shown that such a deformation can be described by a modification of the “pinch” hamiltonian Eq.(7); we will get back to it later. It is clear that any real inclusion will, to a certain extent, combine the three effects described above.

A single inclusion of the “pinch” or “tilt” type induces a non-zero average deformation to the membrane. However, this mean deformation evens out during the course of an experiment, and is not detectable on the scattering spectrum (except if it induces a phase separation, such as expulsion of the solvent). The effect of the inclusions on the spectrum is visible via their effect on the membrane fluctuations. In what follows, we do not investigate a possible coupling between the membrane fluctuations and the inclusion correlation function (assumption of a quenched distribution of inclusions). It has been shown for a single membrane system that “quenched” and “annealed” distributions of inclusions give similar corrections to the fluctuation spectrum of the membrane. The scattering experiments show an average of the structure factor over the positions of the inclusions, it is thus necessary to calculate the membrane correlation function after a proper average over the possible positions of the inclusions.

$`\mathrm{}`$ For the case of “pinch” inclusions, the hamiltonian Eqs.(1,7) can be rewritten in the discrete form:

$`H_{pinch}={\displaystyle \frac{1}{V}}{\displaystyle \underset{q>0}{}}\left\{Q_q\left|u_qu_q^{}\right|^2\left(\xi _qu_q^{}+\xi _q^{}u_q\right)\right\}`$ (9)
$`\mathrm{with}Q_q=Bq_z^2+Kq_{}^4\mathrm{and}\xi _q=\beta _{pinch}iq_z\rho _q`$ (10)
The structure factor is then easily obtained:

$$\frac{|u_q|^2}{V}=\frac{T}{Q_q}+\frac{\xi _q\xi _q^{}/V}{Q_q^2}=\frac{k_BT}{Bq_z^2+Kq_{}^4}+\beta _{pinch}^2\frac{q_z^2}{\left(Bq_z^2+Kq_{}^4\right)^2}\frac{|\rho _q|^2_\rho }{V}$$
(11)
where $`\mathrm{}_\rho `$ designates an average over the position of the inclusions. The first part of the left hand side corresponds to the correlation function in a pure $`L_\alpha `$ phase $`g^{(0)}(𝐫)`$ (Eq.(2)), and the second part describes the corrections due to the inclusions $`\mathrm{\Delta }g`$. This correction reflects correlations between inclusions $`|\rho _q|^2_\rho `$.

If the inclusions are independent from each other, $`|\rho _q|^2/V`$ is the average concentration of inclusions $`\overline{\rho }`$. Similarly to Eq.(4), the correction to the spatial structure factor can be written:

$$\mathrm{\Delta }g(𝐫)=2\eta \frac{\beta _{pinch}^2\overline{\rho }}{2k_BTB}\left(𝑑q_{}\frac{\left(1J_0(q_{}r_{})e^{\lambda zq_{}^2}\right)}{q_{}}+\frac{1}{4}e^{\frac{r_{}^2}{4\lambda z}}\right)$$
(12)
This correction leads to the same power law divergence of the scattering intensity as the pure $`L_\alpha `$ (Eq(6)), with the effective modulii:

$$\left(KB\right)_{pinch}=\frac{KB}{\left(1+{\scriptscriptstyle \frac{\beta _{pinch}^2\overline{\rho }}{2k_BTB}}\right)^2}$$
(13)
One notice right away that the “pinch” inclusions tend to soften the lamellar phase. We thus expect broader peaks in the scattering intensity (see Fig.1). This finding is qualitatively consistent with experiments on comparable systems. Note that results reported for the doped solvent situation tend to show qualitatively similar results for physically different reasons. In that case, the inclusions-mediated attraction between lamellae tends to reduce the compression modulus $`B`$.

Possible correlations between inclusions may be induced either by direct interactions between them, or by membrane mediated interactions. In the Latter case, it has been shown that the inclusions interact via the potential (in Fourier space)

$$U_q=\frac{\beta _{pinch}^2}{2}\frac{q_z^2}{Q_q}$$
(14)
In this case, a ”Debye-Huckel like” expansion of the correlation between inclusions (valid if $`\overline{\rho }U_q/(k_BT)1`$), shows that:

$`{\displaystyle \frac{|\rho _q|^2}{V}}{\displaystyle \frac{\overline{\rho }}{1+\overline{\rho }U_q/(k_BT)}}\mathrm{or}{\displaystyle \frac{|\rho _q|^2}{V}}={\displaystyle \frac{\overline{\rho }Q_q}{Q_q^{}}}`$ (15)
$`\mathrm{with}Q_q^{}B^{}q_z^2+Kq_{}^4\mathrm{and}B^{}B\left(1{\displaystyle \frac{\overline{\rho }\beta _{pinch}^2}{2k_BTB}}\right)`$ (16)
From Eq.(11), the corrections to the structure factor is given by:

$$\mathrm{\Delta }|u_q|^2=\frac{\beta _{pinch}^2q_z^2\overline{\rho }}{Q_qQ_q^{}}\eta _{pinch}=\eta \left(1+\frac{\beta _{pinch}^2\overline{\rho }}{k_BT\sqrt{B^{}}(\sqrt{B}+\sqrt{B^{}})}\right)$$
(17)
The effective modulii are now given by:

$$\left(KB\right)_{pinch}=\frac{KB}{\left(1+\frac{\beta _{pinch}^2\overline{\rho }}{2k_BTB}+\frac{3}{16}\left(\frac{\beta _{pinch}^2\overline{\rho }}{2k_BTB}\right)^2\right)}$$
(18)
Note that in the limit of validity of the expansion, the correction due to correlation is quite small.

$`\mathrm{}`$ The investigation of the effect of the “stiff” inclusions is more delicate, because of the non-quadratic nature of the hamiltonian Eq.(8). This leads to a coupling between different Fourier modes of the membrane displacement, in which case Eq.(2) for the real space correlation function has to be replaced by:

$$g(𝐫)=\frac{1}{2}q_0^2\frac{d^3q}{(2\pi )^3}\frac{d^3q^{}}{(2\pi )^3}u_qu_q^{}^{}\left(e^{i\mathrm{𝐪𝐫}}1\right)\left(e^{i𝐪^{}𝐫}1\right)$$
(19)
The discrete version of the hamiltonian reads:

$`{\displaystyle \frac{H_{stiff}}{k_BT}}=`$ $`{\displaystyle \underset{q,q^{}>0}{}}\left(G_{q,q^{}}^0+\mathrm{\Delta }G_{q,q^{}}\right)u_qu_q^{}^{}`$ (20)
$`=`$ $`{\displaystyle \frac{1}{2}}{\displaystyle \underset{q,q^{}}{}}\left({\displaystyle \frac{Q_q}{Vk_BT}}\delta _{q,q^{}}+{\displaystyle \frac{\delta Kv}{k_BTV^2}}q_{}^2q_{}^{}{}_{}{}^{2}\rho _{(qq^{})}^{}\right)u_qu_q^{}^{}`$ (21)
from which the mean square displacement can be calculated:

$$u_qu_q^{}^{}=\left[\left(G^0+\mathrm{\Delta }G\right)^1\right]_{q,q^{}}$$
(22)
In the limit where the correction is only a small perturbation, an expansion of the previous equation $`[G^0+\mathrm{\Delta }G]_{q,q^{}}^1=\left[G^0\right)]^1_{q,q^{}}[\mathrm{\Delta }G]_{q,q^{}}\times [G^0]^2_{q,q^{}}`$ leads to:

$$\frac{u_qu_q^{}^{}}{V}\frac{k_BT}{Bq_z^2+Kq_{}^4}\delta _{q,q^{}}\frac{\delta Kvk_BT}{V}\frac{q_{}^2q^{}{}_{}{}^{2}\rho _{qq^{}}^{}}{(Bq_z^2+Kq_{}^4)(Bq_z^{}{}_{}{}^{2}+Kq_{}^{}{}_{}{}^{4})}$$
(23)
For an homogeneous distribution of particle of average concentration $`\overline{\rho }`$, the Fourier modes are decoupled: $`\rho _q=\overline{\rho }\delta _q`$. It is then easy to show that again, the scattering intensity show peaks with power law divergence, from which the product of the effective compression and bending modulii can be extracted. The renormalization introduced by the inclusions is given by:

$$\left(KB\right)_{stiff}=\frac{KB}{\left(1\frac{\delta Kv\overline{\rho }}{2K}\right)^2}$$
(24)
Note that this results compares to the finding of Netz and Pincus, who have shown that for a single membrane with inclusions, the effective bending rigidity of the membrane is given by $`\kappa _{eff}\kappa +\delta \kappa \varphi `$ at the lowest order in the volume fraction $`\varphi `$ of inclusions.

In contrary to the case of “pinch” inclusions, a stiffening of the $`L_\alpha `$ phase is observed for inclusions increasing locally the bending rigidity of the membrane. This should lead to a sharpening of the peaks in the scattering intensity. Hence, a quick look at the intensity scattered by the sample should give us valuable information on the inclusion-membrane coupling. A stiffening of a Lα phase has been recently observed upon adjunction of polysoap molecules. This is a more complex case, since the polysoaps are fairly long molecules and can be expected to modify (increase) the interactions between membranes in addition to their effect on an isolated bilayers. The reduction of the effective $`\eta `$ has however been observed below the overlap concentration of polysoaps, where the molecules are expected to be spread flat onto the membrane.

$`\mathrm{}`$ The last coupling investigated in this work is the case of anisotropic inclusions which induce a local tilt (spontaneous curvature) on a membrane. It has been shown in a previous work that such a tilt can be conveniently mimic with the aid of an analogy between the “pinch/swell” inclusions and +/- electrical charges in electrostatics (see Fig.2). The deformation of the smectic induced by a “dipole” $`p_z\beta _{tilt}d`$ (where $`\beta _{tilt}`$, the tilt intensity, has the dimension of an energy) normal to the membrane has been calculated:

$$u_{tilt}=\frac{p_z}{16\pi \sqrt{KB}z^2}\left(1\frac{r_{}^2}{4\lambda z}\right)e^{r_{}^2/\left(4\lambda z\right)}$$
(25)
and correspond to imposing a local curvature $`C_0=\beta _{tilt}/(\pi \kappa d)`$ at the position of the inclusion ($`r_{}=0`$, $`z=d/2`$) without changing the layer spacing ($`\kappa =Kd`$ is the rigidity of an isolated membrane). It can be shown easily in the Fourier space that such a deformation can be induced by a coupling between the second order z-derivative of the layer deformation and the inclusion density:

$$\mathrm{\Delta }H_{tilt}=d^3rp_z\rho (𝐫)_z^2u$$
(26)
which, for independent inclusions, leads to the membrane structure factor::

$$\frac{|u_q|^2}{V}=\frac{k_BT}{Bq_z^2+Kq_{}^4}+p_z^2\frac{q_z^4}{\left(Bq_z^2+Kq_{}^4\right)^2}\overline{\rho }$$
(27)
and the correlation function:

$$\mathrm{\Delta }g(𝐫)=\frac{q_o^2p_z^2\overline{\rho }}{2\pi B^2}\frac{d^2q_{}}{2\pi }\frac{dq_z}{2\pi }\left(1e^{i\mathrm{𝐪𝐫}}\right)\frac{q_z^4}{(q_z^2+\lambda ^2q_{}^4)^2}$$
(28)
After integration over $`q_z`$ (the ($``$) indexes are forgotten):

$`\mathrm{\Delta }g(𝐫)={\displaystyle \frac{q_o^2p_z^2\lambda \overline{\rho }}{8\pi B^2}}`$ (29)
$`\mathrm{where}=3{\displaystyle 𝑑qq^3\left(1J_0(qr)e^{\lambda zq^2}\right)}+\lambda z{\displaystyle 𝑑qq^5J_0(qr)e^{\lambda zq^2}}`$ (30)
Both integrals can be computed after a series expansion of the Bessel function. The correction to the correlation function takes the form:

$$\mathrm{\Delta }g(𝐫)=\frac{q_o^2p_z^2\lambda \overline{\rho }}{16\pi B^2}\left(\frac{e^\omega }{(\lambda z)^2}\left(1+\omega \omega ^2\right)+\mathrm{constant}\right)\mathrm{with}\omega \frac{r_{}^2}{4\lambda z}$$
(31)
The X-ray spectrum is obtained by a combination of the above expression with Eqs.(4,5):

$`G_𝐪{\displaystyle d^3re^{i\mathrm{𝐪𝐫}}\left(\frac{l}{r_{}}\right)^{2\eta }\mathrm{exp}\left(\eta \left(E_1(\omega )+\left(\frac{l_{tilt}}{z}\right)^2e^\omega \left(1+\omega \omega ^2\right)\right)\right)}`$ (32)
$`\mathrm{with}l_{tilt}d\sqrt{{\displaystyle \frac{\beta _{tilt}^2\overline{\rho }}{2k_BTB}}}\mathrm{and}\omega {\displaystyle \frac{r_{}^2}{4\lambda z}}`$ (33)
Unlike the case of symmetrical inclusions considered earlier, the asymptotic scaling laws for $`G_𝐪`$ in the limits $`q_z=0`$ or $`q_{}=0`$ cannot be easily extracted from the above expression, since the exponential term in the integral cannot be written as a function of $`r_{}^2/(\lambda z)`$ alone. However, up to the first order in the dipole strength, $`l_{tilt}/z<1`$ for $`z>d/2`$ and we can expand the exponential in the integral Eq.(33). One can see clearly that unlike symmetrical inclusions such as “pinch” and “stiff”, the effect of a “tilt” inclusion cannot be matched by a renormalization of the smectic modulii, as they modify of the power law divergence of the scattering intensity near the Bragg positions. In fact, the inclusions add a term to the intensity, which for $`q_{}=0`$ can be written:

$`I_{tilt}(q_z)q_z^\eta l_{tilt}^2l^{2\eta }\lambda ^{1\eta }{\displaystyle }{\displaystyle \frac{dZ}{Z^{1+\eta }}}e^{iZ}{\displaystyle }{\displaystyle \frac{d\omega }{\omega ^\eta }}e^\omega (1+\omega \omega ^2)e^{\eta E_1(\omega )};Zq_zz`$ (34)
and in the other limit $`q_z=0`$:

$`I_{tilt}(q_{})q_{}^{2\eta }l_{tilt}^2l^{2\eta }\lambda {\displaystyle }{\displaystyle \frac{dR}{R^{1+2\eta }}}J_0(R){\displaystyle }d\omega e^\omega (1+\omega \omega ^2)e^{\eta E_1(\omega )};Rq_{}r_{}`$ (35)
The integrals being prefactors only, the scattering intensity should take the form:

$`I_{tilt}\{\begin{array}{cc}\alpha _1q_z^{2+\eta }\alpha _2q_z^\eta \hfill & \text{for }q_{}=0\hfill \\ \alpha _1q_{}^{4+2\eta }\alpha _3q_{}^{2\eta }\hfill & \text{for }q_z=0\hfill \end{array}`$ (36)
where the $`\alpha _i`$ are constants. The first term is the Caillé law for a pure smectic phase and the second term is the signature of the asymmetric inclusions. Note that the effect of the tilt inclusion as described by Eq.(36) is most probably very small and might be hardly detectable in experiments. Note also that as a “pinch” inclusion lead to a renormalization of the compression modulus $`B`$ (see Eq.(16)), a “tilt” inclusion can be seen as renormalizing the elastic modulus of an higher order term of the smectic hamiltonian, of the form $`K^{}(_z^2u)^2`$, which has been neglected in the reference hamiltonian Eq.(1). This reinforces the fact that their effect on the structure factor must be small. One nevertheless might hope to detect such an effect because of its dependence on the concentration of inclusion.

$`\mathrm{}`$ In conclusion, we have derived the effect of inclusions on the shape of the scattering curve of a smectic $`L_\alpha `$ phase. We have considered the three inclusions depicted in Figs.1 and 2, and have shown that the two symmetrical inclusions (“pinch” and “stiff”) renormalize the elastic modulii of the phase without modifying the power law divergence near the peaks (Caillé law). In agreement with experiments, we have obtained that the “pinch” inclusions soften the lamellar phase, while the “stiff” inclusions makes it more rigid. the “tilt” inclusions are predicted to modify the shape of the scattering curves, but their effect is small and might hardly be detectable.

As a last remark, we note that in view of our results, the study of the X-ray scattering spectrum of a lamellar phase with inclusions should be a powerful tool to investigate some changes in conformation of membrane proteins. One can argue that the bridge (pinch) between neighboring membranes created by some amphiphilic peptides and reported in  may be pulled out of one of the two membranes as the intermembrane spacing is increased. If the peptide is flexible enough, it is then expected to lay flat on one membrane. This conformation change represents a transition from a “pinch” to a “stiff” inclusion, and should be directly detectable from the shape of the scattering peak. Moreover, some amphiphilic helical peptides in contact with a lipid bilayer are known to go from an adsorbed state at low peptide concentration, to an inserted state (a certain number of peptide aggregating to form an hydrophilic pore) at higher concentration. This transition from one state to the other could be interpreted in terms of “stiff” and “soft” (in the case of a pore) inclusions; One could investigate X-ray scattering from such a solution with this transition in mind. Nevertheless, one should always bear in mind that the different states of the amphiphilic peptides are most probably a complex mixed between the types of inclusion proposed in our work, and probably others.

We acknowledge stimulating discussions with W. Urbach, N. Taulier and N. Tsapis (E.N.S Paris), and we would like to thank J.-F. Joanny and A. Johner (I.C.S Strasbourg) for fruitful comments.

Fig.1 The two symmetrical inclusions considered in the paper. A “pinch” inclusion modifies

the layer spacing and broadens the Bragg peak - A “stiff” inclusion increase the local rigidity

and makes the Bragg peak narrower

Fig.2 a) The “tilt” inclusion induces a spontaneous curvature

b) How to make a “tilt” from the combination of “pinch” and “swell”