# Volatiles in Berries: Biosynthesis, Composition, Bioavailability, and Health Benefits
## 5 Health Benefits of Berry Volatiles
### [PERSON] et al. demonstrated that geraniol inhibits human prostate cancer cell PC-3 proliferation in in vitro and in vivo xenograft mice models [164]. Geraniol at 0.5 and 1 mM significantly suppressed the cell growth of PC-3 by increasing cell cycle arrest and apoptosis. Balb/C nude mice inoculated with PC-3 cells took intratumoral geraniol injection daily for 38 days at 0, 12, 60, or 300 mg/kg. The mice treated with 60 or 300 mg/kg geraniol showed significantly decreased tumor volume and weight. Injection of geraniol at 20 mg/kg also sensitized the chemotherapeutic agent Docetaxel (2 mg/kg) in the xenograft mice model. In [PERSON] et al., geraniol inhibited prostate cancer growth by a down-regulating E2F8 transcription factor and inducing G2/M phase cell cycle arrest [165]. In gastric adenocarcinoma AGS cells, geraniol showed cytotoxicity by inhibiting the JNK/ERK signaling pathway [166]. Lavender essential oil, its active compounds linalool and linalyl acetate [167], and ethyl acetate fraction of Ajwa dates [168] also inhibited PC-3 cell proliferation by increasing apoptosis and cell cycle arrest. Linalool also reduced tumor growth in PC-3 xenograft mice [167] and the 22 Rv1 xenograft mice model [169]. Ethyl acetate exerted an anti-proliferative activity on human breast cancer MCF7 and SKBR3 cells [170] and human cervical cancer HeLa cells [171]. \(\alpha\)-terpineol [172] and linalool [173] also showed strong cytotoxicity on HeLa cells with apoptosis and cell cycle arrest. In human acute myeloid leukemia U937 cells [173], human oral cancer cells [174; 175], and lung adenocarcinoma A549 cells [176], linalool significantly suppressed the cell growth. D-limonene [177] and d-limonene-rich blood orange volatile oils [178] inhibited the proliferation of lung cancer A549 and H1299 cells and human colon adenocarcinoma cells SW480 and HT-29 cells, respectively. Limonene (9 mM) significantly reduced the proliferation of human bladder cancer cell T24 after 24 h, showing induced apoptosis with increased G2/M cell cycle arrest and apoptotic markers (Bax, and cleaved caspase-3, 8, and 9) [179]. In [PERSON] et al., d-limonene induced apoptosis and autophagy-related genes in a lung cancer model [177]. In an acetic acid-induced gastric ulcer rat model, 7-day oral administration of (-)-myrtenol at 50-100 mg/kg increased the healing of the ulcer [180]. Myrcene640 mM significantly decreased the proliferation of SCC9 oral cancer cells after 24 h [181]. Myrcene also showed increased apoptosis with the concentration of 5-20 mM and significantly suppressed the migration of SCC9 cells at 10 mM myrcene treatment. The effects of volatile compounds rich in berries on cancer models were summarized in Table 3.  
\begin{table}
\begin{tabular}{c c c c} \hline \hline
**Volatile Compound** & **Cancer Model** & **Effect** & **References** \\ \hline \multirow{3}{*}{Geraniol} & Human prostate cancer PC-3 cells, & Cell proliferation\(\downarrow\) & \\  & in vitro and in vivo xenograft & Cell cycle arrest and apoptosis\(\uparrow\) & [164] \\ \cline{1-1}  & mice model & Tumor volume and weight\(\downarrow\) & \\ \cline{1-1}  & & Docetaxel sensitization\(\uparrow\) & \\ \hline \hline \end{tabular}
\end{table}
Table 3: The effect of volatile compounds rich in berries on cancer models.