Anthocyanins are a class of water-soluble flavonoids widely present in fruits and vegetables. Dietary sources of anthocyanins include red and purple berries, grapes, apples, plums, cabbage, or foods containing high levels of natural colorants. Cyanidin, delphinidin, malvidin, peonidin, petunidin, and pelargonidin are the six common anthocyanidins. Following consumption, anthocyanin, absorption occurs along the gastrointestinal tract, the distal lower bowel being the place where most of the absorption and metabolism occurs. In the intestine, anthocyanins first undergo extensive microbial catabolism followed by absorption and human phase II metabolism. This produces hybrid microbial–human metabolites which are absorbed and subsequently increase the bioavailability of anthocyanins. Health benefits of anthocyanins have been widely described, especially in the prevention of diseases associated with oxidative stress, such as cardiovascular and neurodegenerative diseases. Furthermore, recent evidence suggests that health-promoting effects attributed to anthocyanins may also be related to modulation of gut microbiota. In this paper we attempt to provide a comprehensive view of the state-of-the-art literature on anthocyanins, summarizing recent findings on their chemistry, biosynthesis, nutritional value and on their effects on human health.

The recent progresses in the research on proline will be described, focusing on plants and covering proline metabolism and signal transduction as well as the role of this imino acid in stress response. Furthermore, the recently described developmental role of proline in flowering and reproduction will be illustrated and discussed.

In addition to its role in protein synthesis and the plant cells' response to environmental stresses, circumstantial evidence suggest that proline may also play a role in flowering and development both as a metabolite and as a signal molecule. Although there is a growing consensus that proline is of special importance throughout the reproductive phase (from flower transition to seed development) a general agreement on the molecular and genetic mechanisms proline is involved in, is yet to be established. In this paper we shall review and critically discuss most of the evidence supporting a role for proline in plant development, paying special attention to the recently reported role of proline in flower transition.

Overexpression of the proline biosynthetic gene P5CS1 results in early flowering in Arabidopsis. However, the p5cs1 loss-of-function mutant exhibits a modest delay in flowering, suggesting that P5CS2, a duplicated P5CS1 gene present in the Arabidopsis, may also play a role in flower transition. In situ mRNA hybridizations and quantitative reverse transcription-polymerase chain reaction (RT-PCR) revealed that P5CS1 and P5CS2 are expressed at similar levels and with the same pattern of expression in vegetative and floral shoot apical meristems as well as in axillary meristems. Arabidopsis lines homozygous for the p5cs1 mutant and simultaneously heterozygous for the p5cs2 mutation showed a stronger late-flowering phenotype than p5cs1 single mutants, confirming that also P5CS2 plays a role in flower transition and supporting the notion of overlapping functions of the two P5CS genes in this developmental process. P5CS1 and P5CS2 have identical messenger RNA (mRNA) distributions also in embryos, but only p5cs2 mutant embryos exhibit alterations of the cellular division planes and consequently stop developing. This suggests a specific role of P5CS2 in embryogenesis and an involvement of proline in cell division. Accordingly, exogenous proline accelerated organ growth and meristem formation, and stimulated expression of the cell cycle-related protein CYCB1;1.

We reported previously that the plant oncogene rolD anticipates and stimulates flowering in Nicotiana tabacum, and encodes ornithine cyclodeaminase, an enzyme catalysing the conversion of ornithine to proline. To investigate on the possible role of proline in flowering, we altered the expression of AtP5CS1, encoding the rate-limiting enzyme of proline biosynthesis in plants. Accordingly we characterized a mutant line containing a T-DNA insertion into AtP5CS1 and introduced in Arabidopsis thaliana AtP5CS1 under the control of the CaMV35S promoter. As expected homozygous p5cs1 mutants behaved as late flowering. In addition p5cs1 mutants exhibited a shorter size and contained lower levels of proline, compared to wild type. 35S-P5CS1 plants, manifested, early in development, overexpression of P5CS1 and accumulation of proline, leading to early flowering, both under long- and short-day conditions. Later in development, down-regulation of P5CS1 occurred in 35S-P5CS1 leaves, leading to proline reduction, and, in turn, impaired bolting and stunted growth. Salt-stress restored expression of P5CS1 and proline accumulation in P5CS1-transformed plants, as well as rescuing growth. Our data suggest that proline plays a key role in flower transition, bolting and coflorescence formation.

BackgroundIn crosses between the proline-deficient mutant homozygous for p5cs1 and heterozygous for p5cs2 (p5cs1 p5cs2/P5CS2), used as male, and different Arabidopsis mutants, used as females, the p5cs2 mutant allele was rarely transmitted to the outcrossed progeny, suggesting that the fertility of the male gametophyte carrying mutations in both P5CS1 and P5CS2 is severely compromised.ResultsTo confirm the fertility defects of pollen from p5cs1 p5cs2/P5CS2 mutants, transmission of mutant alleles through pollen was tested in two ways. First, the number of progeny inheriting a dominant sulfadiazine resistance marker linked to p5cs2 was determined. Second, the number of p5cs2/p5cs2 embryos was determined. A ratio of resistant to susceptible plantlets close to 50%, and the absence of aborted embryos were consistent with the hypothesis that the male gametophyte carrying both p5cs1 and p5cs2 alleles is rarely transmitted to the offspring. In addition, in reciprocal crosses with wild type, about 50% of the p5cs2 mutant alleles were transmitted to the sporophytic generation when p5cs1 p5cs2/P5CS2 was used as a female, while less than 1% of the p5cs2 alleles could be transmitted to the outcrossed progeny when p5cs1 p5cs2/P5CS2 was used as a male. Morphological and functional analysis of mutant pollen revealed a population of small, degenerated, and unviable pollen grains, indicating that the mutant homozygous for p5cs1 and heterozygous for p5cs2 is impaired in pollen development, and suggesting a role for proline in male gametophyte development. Consistent with these findings, we found that pollen from p5cs1 homozygous mutants, display defects similar to, but less pronounced than pollen from p5cs1 p5cs2/P5CS2 mutants. Finally, we show that pollen from p5cs1 p5cs2/P5CS2 plants contains less proline than wild type and that exogenous proline supplied from the beginning of another development can partially complement both morphological and functional pollen defects.ConclusionsOur data show that the development of the male gametophyte carrying mutations in both P5CS1 and P5CS2 is severely compromised, and indicate that proline is required for pollen development and transmission.

No abstract

No abstract