The economics of evolution: Henry Ford and the Model T

For the past 30 years evolutionary biologists have used a fictional tale about engineer and businessman Henry Ford to help illustrate the undesirability of over-design. Thus, on discovering that kingpins were rarely damaged in scrapped Model T automobiles, Henry Ford is alleged to have concluded that the kingpins were unnecessarily durable and asked that they be built to a cheaper specification. The general lesson that has been drawn from this tale is that natural selection will act to equalize the mortality risks accruing from damage to each part of the organism's body. Yet it is well known, at least as far as humans are concerned, that death is more likely to be attributed to a failure of some organs compared to others. To understand why this might be so, we use some graphical and mathematical models to show that even if all body organs were equally important to survivorship, then the optimal investment solution that maximizes whole-organism longevity will typically not be the solution that equalizes the fail-times of the individual organs. As with natural organisms, the key to any optimal investment policy in multi-component systems is understanding what 'bang you can get for your buck'. Moreover, we use a specific model to show that, even following selection to ameliorate the effects of damage, those body parts that receive more damage are still more likely to be the ultimate cause of death – that is, there is 'under-compensation'. Therefore, the decision to make the kingpins more cheaply should not have been based only on the fact they rarely cause car failure compared to other car components. Such arguments wrongly assume that if one car part (or body part) is less durable than the others, then it will always be the reason for any future breakdown (or death).     

Compound leaf development and evolution in the legumes

Across vascular plants, Class 1 KNOTTED1-like (KNOX1) genes appear to play a critical role in the development of compound leaves. An exception to this trend is found in the Fabaceae, where pea (Pisum sativum) uses UNIFOLIATA, an ortholog of the floral regulators FLORICAULA (FLO) and LEAFY (LFY), in place of KNOX1 genes to regulate compound leaf development. To assess the phylogenetic distribution of KNOX1-independent compound leaf development, a survey of KNOX1 protein expression across the Fabaceae was undertaken. The majority of compound-leafed Fabaceae have expression of KNOX1 proteins associated with developing compound leaves. However, in a large subclade of the Fabaceae, the inverted repeat-lacking clade (IRLC), of which pea is a member, KNOX1 expression is not associated with compound leaves. These data suggest that the FLO/LFY gene may function in place of KNOX1 genes in generating compound leaves throughout the IRLC. The contribution of FLO/LFY to leaf complexity in a member of the Fabaceae outside of the IRLC was examined by reducing expression of FLO/LFY orthologs in transgenic soybean (Glycine max). Transgenic plants with reduced FLO/LFY expression showed only slight reductions in leaflet number. Overexpression of a KNOX1 gene in alfalfa (Medicago sativa), a member of the IRLC, resulted in an increase in leaflet number. This implies that KNOX1 targets, which promote compound leaf development, are present in alfalfa and are still sensitive to KNOX1 regulation. These data suggest that KNOX1 genes and the FLO/LFY gene may have played partially overlapping roles in compound leaf development in ancestral Fabaceae but that the FLO/LFY gene took over this role in the IRLC.     

A model for the evolution of pathogen-induced leaf shedding

A size-structured plant population model is developed to study the evolution of pathogen-induced leaf shedding under various environmental conditions. The evolutionary stable strategy (ESS) of the leaf shedding rate is determined for two scenarios: i) a constant leaf shedding strategy and ii) an infection load driven leaf shedding strategy.
The model predicts that ESS leaf shedding rates increase with nutrient availability. No effect of plant density on the ESS leaf shedding rate is found even though disease severity increases with plant density.
When auto-infection, that is increased infection due to spores produced on the plant itself, plays a key role in further disease increase on the plant, shedding leaves removes disease that would otherwise contribute to disease increase on the plant itself. Consequently leaf shedding responses to infections may evolve. When external infection, that is infection due to immigrant spores, is the key determinant, shedding a leaf does not reduce the force of infection on the leaf shedding plant. In this case leaf shedding will not evolve.
Under a low external disease pressure adopting an infection driven leaf shedding strategy is more efficient than adopting a constant leaf shedding strategy, since a plant adopting an infection driven leaf shedding strategy does not shed any leaves in the absence of infection, even when leaf shedding rates are high. A plant adopting a constant leaf shedding rate sheds the same amount of leaves regardless of the presence of infection.
Based on the results we develop two hypotheses that can be tested if the appropriate plant material is available.     

Transport dependence of leaf evolution in dicots

The diversity of tissue and cell organization in the leaves of dicots is explained as the mutual effect of light and water fluxes distribution. Equally with certain data about the role of light distribution, the same influence of water flux distribution on the leaf structure is recognized. Dorsiventral leaves of woody plants have an adequate to structure dorsiventral ring of water circulation. Rising flux from the xylem allocates via leaf apoplast with intermediate accumulation in upper epiderma. Descending flux starts and returns to bundle moving from cell to cell along the symplast (ER) of spongy parenchyma, bundle sheath and terminal complexes of the phloem. Isolateral leaves of herbs have a concentric pathway of solute circulation corresponding to the structure. Xylem flux allocates via symplast with water and nitrogen accumulation in paraveinal parenchyma. Water returns to phloem by transit via the apoplast in parallels with phloem exudate formation. Structural features correlated with the model of water circulation in the leaf are described. Numerous lines of leaf evolution well-known for dicots collect to two main topics which are typical for woody and herbaceous forms of dicots. The mechanisms of cell and tissue differentiation under the control of transport fluxes are discussed with special attention to ontogenetic and phylogenetic trends.     

The inheritance and evolution of leaf pigmentation and pubescence in teosinte

To investigate the genetic mechanisms that underlie morphological evolution in natural populations, we employed QTL mapping to dissect the inheritance of leaf sheath characters that distinguish Chalco from Balsas teosinte. Abundant macrohairs (trichomes) and intense anthocyanin accumulation are found in Chalco teosinte sheaths whereas Balsas teosinte leaf sheaths are green and glabrous. These character states may represent adaptations to the cooler highland (Chalco) vs. warmer middle-elevation (Balsas) climates. QTL mapping in multiple populations revealed a mix of major- and minor-effect QTL affecting both sheath color (anthocyanin) and macrohair abundance. The major QTL for macrohairs accounts for 52% of the parental difference. Epistatic interactions were detected between the major-effect QTL and multiple other QTL for both traits, accounting for substantial portions of phenotypic variance. Developmental analyses suggest that regulatory program changes underlie the phenotypic differences. Sheath anthocyanin QTL are clearly associated with b1 and a3, both of which are regulators of anthocyanin biosynthesis. Our findings suggest that changes in a small number of QTL can lead to morphological evolution by modulating existing developmental programs.     

Chlorophyll fluorescence emission spectrum inside a leaf.

Chlorophyll a fluorescence can be used as an early stress indicator. Fluorescence is also connected to photosynthesis so it can be proposed for global monitoring of vegetation status from a satellite platform. Nevertheless, the correct interpretation of fluorescence requires accurate physical models. The spectral shape of the leaf fluorescence free of any re-absorption effect plays a key role in the models and is difficult to measure. We present a vegetation fluorescence emission spectrum free of re-absorption based on a combination of measurements and modelling. The suggested spectrum takes into account the photosystem I and II spectra and their relative contribution to fluorescence. This emission spectrum is applicable to describe vegetation fluorescence in biospectroscopy and remote sensing.     

The genetics and evolution of the mariner transposable element in Drosophila simulans: worldwide distribution and experimental population dynamics

We have studied both the frequency and biogeographical distribution of the transposable DNA element mariner in natural populations of Drosophila simulans and the short-term evolutionary characteristics of mariner in experimental populations. The mariner element has been identified in natural populations of D. simulans from Africa, Europe, the Middle East, Japan, Australia, several Pacific islands, North America, and South America. Only four lines out of 296 were devoid of active mariner elements, as measured by the presence of functional mariner transposase. A slight correlation was found between the latitudinal coordinate of the collection sites and the level of mariner activity in the population; this correlation became highly significant in Australia where a cline in mariner activity was observed along the eastern coast of the continent. We also observed that wild-type laboratory strains kept for several years as small populations might lose mariner activity over time. Using experimental populations, we modeled what might happen when naturally occurring populations exhibiting high and low levels of mariner activity encounter one another. We found that active mariner elements either will tend to lose their activity over time and gradually become inactive or possibly will be lost from the population; in either case, this will lead to the pattern seen in this experiment of a significant loss of mariner activity over time. This revised version was published online in July 2006 with corrections to the Cover Date.     

The impact of plasticity and maternal effect on the evolution of leaf resin production in Diplacus aurantiacus

Our previous quantitative genetic study of leaf resin production in Diplacus aurantiacus revealed large environmental and maternal effects on variation in resin production, which suggests the possibility of a genotype×environment interaction for this trait when plants grow in heterogeneous environments. Our objectives in this study were to observe the genetic variation in plasticity of resin production under field and chamber conditions, compare phenotypic correlations of resin content with growth traits under these two environmental conditions, and distinguish the possible basis of the maternal effect on resin production using parents and half-sib progeny. A significant genotype×environment interaction (P<0.0001) in leaf resin production was found, which suggests a potential for the evolution of plasticity of these secondary metabolites under heterogeneous environments. The phenotypic correlation between resin content and growth rate also exhibited plasticity. In addition, the resin content of dam half-sib families grown in the chamber had a closer relationship with their maternal parents in the field (r=0.65, P=0.059) than in the chamber (r=0.39, P=0.34), suggesting an environmentally based maternal effect on the secondary chemicals. We suggest that the maternal environmental effect may act as a contributor to plasticity of resin production and, while it may not diminish the appearance of the genotype×environment interaction, the heritable variation of plasticity of resin production may be confounded.     

Daniel Bernoulli (1738): evolution and economics under risk

Conclusion In this 300th anniversary of Daniel Bernoulli's birth, this essay traces the influence of one of his works usually regarded by mathematicians and physicists as too minor to mention. From this source has flowed much of our understanding of how to deal with risk in economics and evolution. The concepts introduced by Bernoulli help us to think about the evolution of reproductive lifespan, dormancy and diapause, sexual versus asexual reproduction, and population dynamics. In economics they form the foundation of portfolio and insurance theory. The 1738 paper was definitely not minor.     

Angiosperm leaf vein evolution was physiologically and environmentally transformative

The veins that irrigate leaves during photosynthesis are demonstrated to be strikingly more abundant in flowering plants than in any other vascular plant lineage. Angiosperm vein densities average 8 mm of vein per mm² of leaf area and can reach 25 mm mm⁻², whereas such high densities are absent from all other plants, living or extinct. Leaves of non-angiosperms have consistently averaged close to 2 mm mm⁻² throughout 380 million years of evolution despite a complex history that has involved four or more independent origins of laminate leaves with many veins and dramatic changes in climate and atmospheric composition. We further demonstrate that the high leaf vein densities unique to the angiosperms enable unparalleled transpiration rates, extending previous work indicating a strong correlation between vein density and assimilation rates. Because vein density is directly measurable in fossils, these correlations provide new access to the physiology of extinct plants and how they may have impacted their environments. First, the high assimilation rates currently confined to the angiosperms among living plants are likely to have been unique throughout evolutionary history. Second, the transpiration-driven recycling of water that is important for bolstering precipitation in modern tropical rainforests might have been significantly less in a world before the angiosperms.     

Origin and evolution of the unusual leaf epidermis of Caryodaphnopsis (Lauraceae)

We studied leaf epidermal anatomy of Caryodaphnopsis Airy Shaw, a genus disjunct between tropical Asia and tropical America, using light microscope and scanning electron microscope. We sampled 10 species of Caryodaphnopsis and 52 species of other Lauraceous genera. Our observations suggest that this genus possesses a unique lower leaf epidermis. Compared with other leaves of Lauraceae, this genus has an additional layer covering the lower leaf epidermis and the stomatal apparatus. The additional layer is either closed or poriferous/reticulate. The outer periclinal walls of the lower leaf epidermis protrude outside forming hollow domes or columns, and the distal endings of these domes or columns are expanded and fused, which forms the additional covering layer. Three different types are recognized in the genus: (1) the middle portion of the protuberances is not contracted and the distal endings are free or adnate to each other, with only limited space between the two layers; (2) the middle portion is slightly contracted in the outer part, the distal endings are fused, with more space between the two layers; (3) the middle portion is conspicuously contracted and elongated into columns, the distal endings are fused and it is roomy between the two layers. The structure of the leaf lower epidermis is reconstructed and illustrated for the first time. This unusual leaf lower epidermis of Caryodaphnopsis is derived in Lauraceae and is an autapomorphic character. Outward protrusions of the outer periclinal walls forming papillate protuberances surrounding the stomatal apparatus are also found in a few other genera including Neocinnamomum, a genus closely related to Caryodaphnopsis, but the distal endings of these protrusions are not expanded and connected. We hypothesize that the periclinal wall of the lower leaf epidermis has been gradually modified in Lauraceae, from a smooth pattern in most genera, to papillate pattern (e.g. Neocinnamomum), and to the double layered lower leaf epidermis in Caryodaphnopsis. The origin and evolution of this unique lower epidermis might have been related to the climatic cooling and aridification since the late Eocene.